Hydrobiologia (2008) 595:365377

DOI 10.1007/s10750-007-9031-3

FRESHWATER ANIMAL DIVERSITY ASSESSMENT

Global diversity of stoneflies (Plecoptera; Insecta)

in freshwater
Romolo Fochetti
Jose Manuel Tierno de Figueroa

Springer Science+Business Media B.V. 2007

Abstract Plecoptera, or stoneflies, is a small order Zealand (104 species) are relatively well-known,
of hemimetabolous insects: according to our data, while our knowledge of the Plecoptera of Central and
more than 3,497 species have been described so far in South America (95 and 378 species respectively) is
the world. The total number of species has enor- poor and still not representative of the real diversity.
mously increased in the last 30 years (2,000 species Africa has a reduced stonefly fauna (126 species).
estimated in 1976) and, if the trend continues, then it Asian stonefly diversity (approximately 1,527 spe-
will nearly double in the near future. The order is cies) is much greater than that of Europe or North
divided into the suborders Arctoperlaria and Antarct- America despite the fact that, except for Japan and
operlaria, and includes 16 families: 12 arctoperlarian Asiatic Russia that have been well studied, our
and 4 antarctoperlarian. The Arctoperlaria account knowledge of the remaining Asiatic areas is extre-
for a total number of 3,179 species, and Antarctop- mely poor. Even though our data indicate the
erlaria, only 318 species. The total number of genera Holarctic Region as the diversity hot-spot for the
is 286. We give in this article the estimated number order, the analysis of the specific diversity divided by
of species for each family. The fauna and diversity of family suggests also an important role of tropical
stonefly in North America (650 species reported) and stoneflies.
Europe (426 species) are best known. Nevertheless,
in the last 25 years, a mean of 2.6 Plecoptera species Keywords Plecoptera
Distribution

per year were described in Europe. Stonefly-faunas of Diversity
Zoogeographical regions
Australia (191 species, Tasmania included) and New

Introduction
Guest editors: E.V. Balian, C. Leveque, H. Segers &
K. Martens Plecoptera is a small order of hemimetabolous
Freshwater Animal Diversity Assessment insects, commonly called stoneflies, with more than
3,497 described species (this article). Stoneflies are
R. Fochetti (&)
Dipartimento di Scienze Ambientali, Universita della distributed over all continents except Antarctica, and
Tuscia, Largo dellUniversita snc, 01100 Viterbo, Italy constitute a significant ecological component of
e-mail: fochetti@unitus.it running water ecosystems. Their ecological require-
ments greatly limit the dispersal capacity of the
J. M. Tierno de Figueroa
Departamento de Biologia Animal, Facultad de Ciencias, nymphs and, because adults have reduced flight
Universidad de Granada, 18071 Granada, Spain ability, stoneflies show a high percentage of

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366 Hydrobiologia (2008) 595:365377

endemism. Due to the growing pollution and alter- present, gills are located on different parts of the
ation of water courses and to the high stenoecity of body.
Plecoptera, numerous stonefly species are reduced to Plecoptera nymphs are aquatic and live mainly in
small isolated populations and many others have cold, well-oxygenated running waters, although
already gone extinct. Stoneflies are probably one of some species can also be found in lakes. Few
the most endangered groups of insects (Fochetti & species are adapted to terrestrial life in Sub-Antarc-
Tierno de Figueroa, 2004). tic areas: Hynes (1976) reported the tendency of
Plecoptera can be easily recognized by several southern hemisphere nymphs to leave the water.
morphological characters: soft body, three segmented However, the increasing number of stoneflies
tarsi, elongate filiform antennae, mandibulated described from the tropics and their high rate of
mouthparts, two compound eyes, two or three ocelli, endemicity can modify the common belief that
two usually long cerci, 10-segmented abdomen with Plecoptera are cold-water specialists, and suggest
vestiges of the eleventh segment. Adults have two instead that the true hot-spot for Plecoptera diversity
pair of membranous large wings (sometimes reduced are tropical areas (Zwick, pers. com.). The life cycle
or absent), and subequal fore and hind wings (hind of stoneflies lasts for one or more years, but there
wings slightly wider) that fold horizontally over and are also bi- or tri-voltine species. Nymphal or egg
around abdomen when at rest (hence the name: diapause is not uncommon. The nymphs can moult
plecos = folded; pteros = wings) (Fig. 1A). Nymphs up to 33 times before emerging. They feed on
are similar to adults (Fig. 1B), with a closed tracheal animal or vegetable matter as collectors, scrapers,
system with or without filamentous gills. When shredders or predators.

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

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Hydrobiologia (2008) 595:365377 367

Adults are terrestrial or sub-aerial, with the Very few species are shared among zoogeograph-
exception of Capnia lacustra Jewett, whose adults ical regions (mainly between Nearctic and Palaearctic
have been collected only at a depth of 6080 m in the and between Palaearctic and Oriental) or among
Lake Tahoe (USA) (Jewett, 1963), and the adult continents, i.e. at borders between Africa and Europe
females of Zapada cinctipes (Bank) that can remain or at the borders between Europe and Asia. Australia
under water for a long time (2060 min) for ovipos- and New Zealand are obviously isolated; deserts or
iting (Tozer, 1979). Adults lifespan is short, from a dry lands prevent stonefly migration from North to
few days to weeks; they display complex behaviours Central and South America, and from the Palaearctic
related to mate searching (such as the use of to the Ethiopic Regions. Thus, the total number of
intersexual vibrational communication in Arctoper- described species should not be too far from the
larian stoneflies) and oviposition (Stewart, 1994). The reported one.
adult feeds on pollen, lichens, cyanobacteria, etc., In comparison to the papers by Hynes (1976) and
although the adults of the largest species do not seem Zwick (1980) who reported about 2,000 described
to feed (Tierno de Figueroa & Fochetti, 2001). stonefly species, the total number has enourmously
Females produce 253,000 eggs, and lay their egg- increased in the last 25 years and, if the trend
clutches, or rarely each egg separately, on the water. continues then it will nearly double in the near future.
Parthenogenesis and ovoviviparity have been also The fauna and diversity of stonefly in North
described for some species. 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
Species diversity Plecoptera species per year were described in Europe
(Fochetti & Tierno de Figueroa, 2005). Stonefly-
According to our data, approximately 3,500 species faunas of Australia (Tasmania included) and New
of Plecoptera have been reported so far in the world Zealand are relatively well-known, while our knowl-
(Tables 1, 2). They are divided as: North America edge of the Plecoptera of Central and South America
650 species (Stark & Baumann, 2005), Central is poor and still not representative of the real
America 95 species, South America 378 species diversity.
(Heckman, 2003), Europe 426 species (Fochetti & Asian stonefly diversity is much greater than that
Tierno de Figueroa, 2004), Africa 126 species (31 of Europe or North America. This is true despite the
from Mediterranean Africa, with only 12 endemic fact that, except for Japan and Asiatic Russia that
species, 50 from tropical Africa, 33 from South have been well studied, our knowledge of the
Africa and 12 from Madagascar), Asia approximately remaining Asiatic areas is extremely poor. Our
1,527 species [(about 784 from South-East Asia = 31 greatest lack of knowledge and the highest uncer-
Taiwan, 32 Korea, 25 Philippines, 39 Borneo, 36 tainty regard Asiatic faunas occurs in the Oriental
Indonesia, 17 Malaysia, 29 Thailand, 20 Vietnam, Region. For instance, according to Yuzhou & Junhua
two Laos, three Myanmar, 28 Bhutan, two Bangla- (2001) over 230 species of Perlidae, the most
desh, 10 Sri Lanka, 112 India, 63 Nepal, 20 Pakistan, speciose family in the country, have so far been
nine Afghanistan, 306 Japan, no species from Cam- described from China. The identity of many of these
bodia; Sivec & Yang 2001), China 350 (?) species species is questionable and cannot be checked
(deduced from Yuzhou & Junhua, 2001, who report because most of the types were lost or destroyed;
231 species of Perlidae), West Asia about 114 species nonetheless, intensive future research will certainly
(two Syria, two Israel, 25 Iran, 15 Lebanon, 70 lead to the discovery of hundreds of new species in
Asiatic Turkey), Asiatic Russia 279 species (179 this country. In fact, a few intensive collecting trips to
Nemouroidea and about 100 Perloidea). Although a Taiwan almost doubled its stonefly-fauna (Sivec &
small number of species are shared by several Yang, 2001).
countries, we consider the total sum as a good In Africa, a reduced number of species were
approximation)], Australia 191 species (Michaelis & collected, but we do not expect a great increase in
Yule, 1988), New Zealand 104 species (McLellan, Plecoptera diversity in the future. The results of
2006). recent studies on the systematics of the tropical tricky

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

NAm CAm SAm Eu AF ASa 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
a
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)

genus Neoperla might increase the species list and

Table 2 Number of Plecoptera species in each family and
each zoogeographical region some new species could be described from Mediter-
ranean countries (Morocco, Algeria, Tunisia), but the
PAa NA NT AT OLb AU World
total stonefly diversity will not increase to the level of
Eustheniidae 2 19 21 the other continents.
Diamphipnoidae 5 5 As we expected, the analysis of Plecoptera diver-
Austroperlidae 4 11 15 sity and distribution shows their well-known
Gripopterygidae 73 204 277 ecological requirements as: they are absent from arid
Pteronarcyidae 10 1 13 or semi-arid areas everywhere in the world. There are
Peltoperlidae 22 46 almost no records from West Australia, no records at
Styloperlidae 8 all for Saudi Arabia, Yemen, Oman, Emirates,
Perlodidae 63 125 4 311 Kuwait and Iraq in Asia, and stonefly diversity in
Perlidae 18 82 348 52 1,049 Africa is greatly limited by deserts and dry lands.
Chloroperlidae 20 95 2 206
The order is divided into the suborders Arctoper-
Scopuridae 8
laria and Antarctoperlaria (see the phylogeny chapter
Taeniopterygidae 42 34 142
below), and includes 16 families: 12 arctoperlarian
and four antarctoperlarian. The Arctoperlaria account
Notonemouridae 17 43 61 121
for a total number of 3,179 species, and Antarctop-
Nemouridae 136 71 9 633
erlaria, only 318 species. They are listed in Table 2
Capniidae 21 156 8 315
with the estimated number of species for each family.
Leuctridae 138 55 1 327
The total number of genera is 286 (Table 3). Even
Total 438a 650 474 95 350? 295 3,497
though Table 1 is incomplete because it lacks details
a
Palaearctic Asia not included. PA: Palaearctic Region, NA: on the distribution of Chinese families and Far East
Nearctic Region, NT: Neotropical Region, AT: Afrotropical
Russia species, Perlidae is the most diverse family,
Region, OL: Oriental Region, AU: Australasian Region. b The
distribution of Oriental Plecoptera cannot be unequivocally with more than 1,000 described species. Since the last
established estimate (Zwick, 1980), the number of perlid species

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almost doubled, due to the increase of faunistic Capniidae and Leuctridae include a similar num-
research in Asia and, above all, to the improved bers of species, 311 and 321 respectively, both with
knowledge of the systematics of the tropical genera mainly Holarctic distribution and a strict systematic
Neoperla and Anacroneuria. The family Perlidae affinity. The most diverse genera are the Holarctic
spreads over Holarctic, Oriental, Afrotropical and Capnia (110 species), Allocapnia (Nearctic; 43
Neotropical Regions. The most diverse perlid areas species), Mesocapnia (Nearctic and East Palaearctic,
are Asia, and Central and South America. Presently the 19 species) among Capniidae, and Leuctra (Holarctic,
Chinese perlid species represent about 25% of the reaching Northern India; 190 species) and Rhopalop-
world Perlidae, making China one of the most impor- sole (East Palaearctic and Oriental; 20 species)
tant areas of the world for perlid diversity (Yuzhou & among Leuctridae. The genus Megaleuctra presently
Junhua, 2001). The Neotropical perlid fauna currently includes six North American and one North Korea
includes more than 350 species: Anacroneuria is the species; it is also known the fossil record in North
dominant and most diverse stonefly genus, (Stark, America and in Baltic amber.
2001) with almost 300 described species. Notonemouridae (121 species), is considered the
Perlodidae presently includes about 300 species, sister taxon of Nemouridae (but see below), and thus
mainly in the Holarctic region. The most speciose is an Arctoperlarian family, but shows a disjunct
perlodid genus, Isoperla, has a Holarctic distribution South African (included Madagascar), South Amer-
and has more than 100 species, a third of the total. A ican and Australian (included New Zealand)
similar distribution is shown by the ancient genus distribution. The family may not be monophyletic
Arcynopteryx [Aubert (1959) stated it is an Ercinic (Zwick, 1990) and its biogeography is not fully
relict] and by the genus Diura. A. compacta understood yet. Austrocercella (15 species) is the
(McLachlan) is distributed in North America and most diverse genus.
Europe, as well as D. bicaudata (Linnaeus) and D. Among the remaining Arctoperlaria, the small
nanseni (Kempny). family (eight species) of wingless stoneflies Scopu-
Chloroperlidae include more than 200 species ridae is known only from Japan and Korea, and the
mainly in the Holarctic region. The genera Sweltsa family Pteronarcyidae (13 species) is present in North
and Alloperla have a Nearctic and East Palaearctic America and East Asia. Peltoperlidae has a disjunct
distribution and includes almost 80 species. In North American and East Asian distribution (44
Europe, only four genera and 19 species are present. species). Styloperlidae a recently instituted family
Taeniopterygidae is a family present in the (eight species), erected from aberrant Peltoperlidae,
Nearctic and Palaearctic Regions, with almost 150 is presently known only from China and Taiwan.
species. The genus Taeniopteryx (20 species) is Among Antarctoperlaria, Gripopterygidae is the
present in Europe and North America (even though most diverse family, with 277 described species. It
the morphology of the nymphs slightly differs in the has a disjunct South American and Australian
two continents), as well as the genus Oemopteryx. (included New Zealand) distribution, but no genera
The genus Doddsia is shared by North America and are shared between the two continents. Dinotoperla
Japan. The West Palaearctic genus Brachyptera (33 species), Leptoperla (28 species), Riekoperla (26
includes more than 30 species. species) are the most diverse Australian genera,
Nemouridae was considered the most diverse Zelandobius (27 species) is the most diverse genus of
family, with almost 400 described species (Zwick, New Zealand. The remaining Antarctoperlaria fam-
1980). The number currently is 633 species, but it is ilies are scarcely diversified: Diamphipnoidae (five
now the second most diverse family due to the species) is a small South American family, Eusthe-
enormous increase of Perlidae. The family has an niidae includes two genera in South America (two
Holarctic distribution with some genera reaching the species) one genus in New Zealand (four species) and
Oriental Region; the genera Amphinemura (Holarc- three genera in Australia (15 species), while
tic; more than 100 species), Nemoura (Holarctic; 175 Austroperlidae includes 10 species in Australia (five
species) and Protonemura (Palaearctic; 135 species) genera), a single species, Austroperla cyrene
account for 2/3 of the whole family. Nemoura arctica Tylliard, in New Zealand, and four species (three
Esben-Petersen has a circum-arctic distribution. genera) in South America.

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Fig. 2 Plecoptera Eustheniidae

EUSS
phylogeny according to EU
Diamphipnoidae
Zwick (2000). AN: AA
NN
Antarctoperlaria; AR: Austroperlidae
Arctoperlaria; EUS: GGR
R
GR Gripopterygidae
Eusthenioidea; GR:
Gripopterygoidea; EU: Scopuridae
Euholognatha; NE:
Nemouroidea; SY: Taeniopterygidae
Systellognatha; PT: EE
UU
Pteronarcyoidea; PE: Capniidae
Perloidea. (redrawn from NN
EE
Zwick, 1990) Leuctridae

Nemouridae

Notonemouridae

Pteronarcyidae
PP
T
PT
T Styloperlidae
R
AAR Peltoperlidae

SS
YY
Perlidae

PP
EE ? Chloroperlidae

Perlodidae

Among Antarctoperlaria Eustheniidae are preda- intersegmental ventrolongitudinal muscles are pres-
tors, as are Perloidea in the Northern hemisphere ent that support laterally undulating swimming (this
Arctoperlaria; Austroperlidae are shredders as are the last character also occurs in Zygoptera). Two other
Arctoperlaria Pteronarcyidae. Many Gripopterygidae probably derived characters have been reported for
are scrapers, as are Brachypterainae in the Arctoper- Plecoptera include the absence of an ovipositor
laria fauna; other Gripopterygidae genera are (although it can secondarily appear in some taxa)
ecologically similar to genera and families of and the existence of an accessory circulatory organ
Nemouroidea. (cercus heart) in Perlidae and Perlodidae, that can be
present in the other families as well.
The order includes 16 families whose relationships
Phylogeny and historical processes have been studied by several authors: we will follow
the more recent and widely accepted classification by
Stoneflies are Pterygota, Neoptera, Exopterygota Zwick (2000) (Fig. 2), which recognizes two large
insects. Although they are usually included among groups: Antarctoperlaria, present only in the Southern
Polyneoptera (possibly they are the sister-group of Hemisphere, and Arctoperlaria, distributed mostly in
the remaining Polyneoptera), the relationships of the Northern Hemisphere. The first taxon includes
Plecoptera with other orders of this subdivision are four families, the latter includes 12 families belonging
poorly understood (Gullan & Cranston, 2005). to two subgroups (Systellognatha and Euholognatha)
Among Polyneoptera, Plecoptera can be identified with six families each one.
by a combination of mostly primitive characters. The monophyly of Antarctoperlaria is supported
Moreover, only a few derived characters support the by morphological characters, while monophyly of
Plecoptera monophyly (Zwick, 2000): in the adult, Arctoperlaria is supported by ethological and mor-
gonads form loops with the anterior ends medially phological characters related to mate finding: the
fused; in the male, there is a complex arrangement of intersexual vibrational communication (drumming,
two superimposed seminal vesicles each of which tremulation, rubbing or combinations of them) and/or
forms an anterior loop; in the nymph, strong oblique the existence of structures associated to it in the male

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(lobes, hammers, etc.) that are widely present in all Australia. Before the isolation of Madagascar, Africa
families of this group except Scopuridae, and absent was for sure colonized by one family of Antarctop-
in Antarctoperlaria (Zwick, 2000). Among the exist- erlaria, the Notonemouridae (Zwick, 1990).
ing families, only Notonemouridae seems to be a The Arctoperlaria families mainly have aHolarctic
polyphyletic assemblage (McLellan, 2000). distribution (with the exception of Notonemouridae
According to Rohdendorf & Rasnitsyn (1980), which are exclusive of the Southern Hemisphere),
recent stoneflies derived from Plecopteroidea from with taxa (even species) in common between North
the Permian Period (including three orders: fossils America and East Asia. The European stonefly-fauna
Protoperlaria and Paraplecoptera, possibly not mono- shares only a few genera with Asia and very few with
phyletic, and Plecoptera). The oldest known North America. This may be due to Pleistocenic
Plecoptera fossils are from the early Permian Period, events, such as the temporal separation of Europe and
258263 m.y. ago (Sinitshenkova, 1997). According Asia by the Turgai Strait, the extinctions that
to this author, the origin and evolution of the group is occurred in Europe (families Pteronarcyidae and
linked to temperate areas; during the Mesozoic Peltoperlidae) and the faunistic interchanges through
Period and the role of this group in lenthic ecosys- the Bering Strait. The presence of arctoperlarian taxa
tems was much more important than it has been in the Southern Hemisphere could be due to invasions
afterwards. from the Northern Hemisphere (Zwick, 1981). The
According to Illies (1965), Banarescu (1990) and distribution of the very distinct Notonemoura group
Zwick (1990, 2000) Arctoperlaria and Antarctoper- excludes recent dispersal. It appears that the present
laria originated as independent lines at the splitting of distribution results from dispersal across Antarctica
Pangea and the subsequent separation of Gondwana- (Zwick, 1981, 1990). Nevertheless, Zwick (2000)
land and Laurasia, at the end of Triassic Period reported two alternative hypotheses: extinction of
(Fig. 3). Antarctoperlaria possibly began their diver- Antarctoperlaria in the Northern Hemisphere, and of
sification before the continents separated, producing Arctoperlaria in the Southern one. These hypotheses
some sister-groups distributed in South America, are based on the fact that, although stoneflies are an
Australia and New Zealand. The absence of Antarct- ancient insect order, their extant families may not
operlaria from South Africa and India may be necessarily be old, revealing recent and numerous
interpreted as a later extinction event. These lands speciation and extinction events, more than dispers-
became warmer and drier during their northward sive processes. Moreover, the scarce flight ability of
journey after separation from Antarctica and these insects makes improbable long distance

Fig. 3 Plecoptera families distribution; Arctoperlaria = continuous line; Antarctoperlaria = dotted line (modified from Zwick, 1990)

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Fig. 4 Distribution of
freshwater Plecoptera
species and genera by
zoogeographical region
(species number/genus
number). PAPalaearctic;
NANearctic; NT
Neotropical; AT
Afrotropical; OLOriental;
AUAustralasian; PAC
Pacific & Oceanic Islands,
ANT: Antarctic

dispersal (Zwick, 2000). Also, according to Sinit- Antarctoperlaria. As stated before, Anctartoperlaria
shenkova (1997), the Northern and Southern faunas must have been present in Gondwana before Africa
were isolated from the late Permian Period, when the (and India) split from it; climatic changes are thought
Pangea still existed. to have caused their disappearance from these lands
(Banarescu, 1990; Zwick, 2003). Only two Arctop-
erlaria families (Perlidae and Notonemouridae) are
Present distribution and main areas of endemicity present today in the Afrotropical Region, and with a
low specific diversity; one of them (Perlidae) is
The diversity for the various zoogeographical regions absent in Madagascar. Only the genus Neoperla
can be only roughly estimated; it should be approx- Needham occurs throughout tropical Africa, includ-
imately as follows: Nearctic 650 species, Palaearctic ing South Africa, with an uncertain number of
1,628 species (it arbitrarily includes also Southern species. They vary, according to the different authors,
Chinese species) (Holarctic 2,278), Neotropical 474, from only one [Neoperla spio (Newman)] to at least
Afrotropical (Ethiopic) 95 species, Oriental 350? 50 (Zwick, 1998). The originally Oriental genus
species (Southern Chinese species not included), Neoperla reached Africa probably via the same route
Australian 295 species (Table 3, Fig. 4). Even though as the mammalian Ungulata during the Tertiary
these data indicate the Holarctic Region as the Period (Zwick, 1998). About 31 species of Notone-
diversity hot-spot for the order, the analysis of the muridae, belonging to six genera (Aphanicerca
specific diversity divided by family suggests also an Tillyard; Aphanicercella Tillyard; Balinskycercella
important role of tropical stoneflies (see below). Stevens & Picker; Afronemura Illies; Aphanicercop-
sis Barnard, and Desmonemoura Tillyard) besides
two yet undescribed Neoperla species are present in
The Afrotropical Region (Ethiopian) South Africa (Villet, 2000). Only the endemic
Notonemourid genus Madenemura Paulian, with the
As reported by Zwick (1998), although Africa (as subgenera (considered as genera by some authors)
well as India) was part of Gondwana, its plecopte- Madenemura and Tsaranemura (Paulian, 1951), is
rafauna lacks members of the Gondwanian present in Madagascar. Unfortunately, only the

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Table 3 Number of Plecoptera genera in each family and each zoogeographical region and total genera number
PA NA NT AT OL AU World

Eustheniidae 2 3 5
Diamphipnoidae 2 2
Austroperlidae 3 6 9
Gripopterygidae 24 25 49
Pteronarcyidae 1 2 1 2
Peltoperlidae 4 6 4 11
Styloperlidae 2 2
Perlodidae 29 30 2 1 51
Perlidae 20 15 11 1 24 52
Chloroperlidae 14 12 2 2 17
Scopuridae 1 1
Taeniopterygidae 11 6 12
Notonemouridae 4 7 (8) 12 23
Nemouridae 8 12 2 6 21
Capniidae 13 10 3 17
Leuctridae 7 9 1 2 12
Total 108 102 57 8(9) 41 46 286
PA: Palaearctic, NA: Nearctic, NT: Neotropical, AT: Afrotropical, OL: Oriental, AU: Australasian

females of some species were described; the disap- Perlodidae (one genus), Chloroperlidae (two genera)
pearance of holotypes and the lack of description of and Leuctridae (two genera) complete the list.
some material from collections make it difficult to Unfortunately, the poor and fragmentary knowledge
estimate the total number of species living in of this region makes it difficult to estimate the real
Madagascar (Elouard, pers. com.). The totally diversity and the exact number of Plecoptera species
endemic notonemourid fauna of Madagascar together that must be much larger than what we know. In fact,
with the absence of Neoperla, may be explained by the Chinese stonefly fauna is almost unknown and
the early isolation of Lemuria. hundreds of species are probably still undescribed
(Sivec & Yang, 2001). These authors estimate
approximately 350 Plecoptera species in the countries
The Oriental Region forming the Oriental Region, Southern China
excluded.
Seven Arctoperlaria families are present in the The Oriental stonefly fauna was colonized from
Oriental Region, including 41 genera. The absence close Palaearctic areas (Zwick, 1980), as suggested
of Antarctoperlaria in India may be explained by the by the decrease in species numbers towards the south.
same reasons of its absence in Africa (see above). The first penetration was by members of three
One family, Styloperlidae, composed of two genera familiesNemouridae, Peltoperlidae and Perlidae
and eight species endemic from Taiwan and South that extended down into the rain forest of southeast
China, is endemic to this region. Peltoperlidae (with Asia, thus having somewhat overcome the usual
four Oriental genera) has a Nearctic-Asian distribu- aversion of the order for warm climates.
tion. The remaining families with Oriental
representatives are also widely distributed in the
Holarctic Region; only two of them are well-diver- The Nearctic Region
sified in the Oriental Region: Perlidae (with 24
genera, 12 of which endemic from the Oriental The Nearctic Region includes 650 described species,
Region) and Nemouridae (with six genera). distributed in 102 genera and nine families. Seven of

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these families (Capniidae, Leuctridae, Nemouridae, 1991). On the other hand the tropical Plecoptera
Taeniopterygidae, Chloroperlidae, Perlidae, Perlodi- fauna is also scarce: only the genus Anacroneuria
dae) and 10 genera (Capnia, Leuctra, Amphinemura, (Perlidae) has been able to colonize from the South,
Nemoura, Oemopteryx, Taeniopteryx, Agnetina, Iso- with a few scattered species across the countries (34
perla, Arcynopteryx, Diura) also occur in the in Mexico, eight in Nicaragua, 12 in Costa Rica, 15 in
Palaearctic Region. There is a substantial division Panama, two in Trinidad). However, much research is
of eastern and western North America genera; only a needed to get a reliable knowledge of Central
few of them, in fact, transgress the barrier formed by America stonefly-fauna.
the central prairies region and the Mississippi valley.
Twelve of the western genera are shared with the
Russian Far East or Japan, suggesting a recent South America
invasion from the west. The genus Anacroneuria
invaded from the south, bringing species into Central The stonefly fauna of South America is a complex
America, Texas and Arizona. This occurred after the mixture of arctoperlarian and antarctoperlarian species
formation of the Isthmus of Panama, during the included in six families: Perlidae, Notonemouridae,
Pliocene Period, about 4 millions years ago. Very Diamphinoidae, Eustheniidae, Austroperlidae, and
few species have been able to disperse south into Gripopterygidae (Tables 1, 2, 3). Among Antarctoper-
Central America, which has a mainly Neotropical laria, Eustheniidae (two species) is confined to southern
stonefly-fauna. South America, with two monospecific genera. The
According to Hynes (1988) North America has sister family, Diamphipnoidae (five species), is found
four groups of stoneflies with different origins: only in central Chile, with two genera. Austroperlidae
(four species) also occurs only in Chile, with three
(1) The original eastern fauna that was present
genera. The family Gripopterygidae (73 species), the
before the final break from Europe, a great
largest family of the suborder (24 genera, 10 of which
many of which may have evolved initially in
are monospecific) is more widespread. Unlike the
North America;
former families, the latter is also present in subtropical,
(2) Invasions from the west, possibly over a long
but not in tropical, South America. Only two families
period, while the Cordillera was forming;
of Arctoperlaria are found in this region, Perlidae and
(3) Invasion of one genus from the south after the
Notonemouridae. Perlidae is the most diverse family
formation of the isthmus;
(277 species), including 11 genera: the very speciose
(4) Invasion by several species from the Bering
genus Anacroneuria, restricted to Central America and
Strait within the last few thousand years. These
the tropical South America, includes almost 220
species may have survived the Pleistocene in the
species. Kempnya (29 species) and Macrogynoplax
Alaskan refugium.
(12 species) are diversified genera as well, while the
remaining genera have few species. Finally, Noton-
emouridae (17 species), the only Plecoptera family
The Neotropical Region with a Gondwanian distribution (not circumantarctic
like the Antarctoperlaria), occurs in southern South
Central America America with four genera. However, it must be
reminded that the monophyly of this family is still
The stonefly-fauna of Central America is poorly questioned (Zwick, 2000).
diversified. Deserts and dry areas prevent the immi- It must be noted that, considering the number of
gration of the rich North American Plecoptera fauna: species, the true hot-spot of this area (that is almost
in fact less than 25 Nearctic species are known coincident with the Neotropical Region), is the
from Central America. They came during the Pleis- tropical belt. However, this is due to the presence
tocene Period, when stoneflies were able to cross the of several speciose genera (especially the genus
discontinuity between the southern Rocky Mountains Anacroneuria). Considering generic diversity, the
and the Sierra Madre Occidental and disperse into subtropical and temperate areas are more diversified,
northern, central and southern Mexico (Sargent et al., with many genera including few species.

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Hydrobiologia (2008) 595:365377 375

The Palaearctic Region genera, with only one non-European genus: Afrop-
erlodes Miron & Zwick. Some species are shared
The Palaearctic Region which includes Europe, with Southern Europe or are very closely related.
Northern Africa and a great part of Asia (except Currently 15 genera and about 31 species have been
Southeast), includes ten families and more than 1,600 reported for this area (Sanchez-Ortega & Azzouz,
Plecoptera species. 1998), approximately 1/3 of which are endemic. The
Seven families (Perlidae, Perlodidae, Chloroperlidae, similarity with Southern Europe and particularly with
Taeniopterygidae, Nemouridae, Capniidae and Leuctri- the Southern Iberian Peninsula may be explained by
dae), 35 genera and 426 species constitute the the existence in the past of a land bridge connecting
Plecoptera fauna of the European continent (Fochetti the two areas. In fact, the Betic System (Spain) and
& Tierno de Figueroa, 2004). Thirty-seven percent of the Rif Mountain (North West Africa) were separated
the European genera are monospecific, whereas the four only in the Lower Pliocene, when the Gibraltar Strait
more diversified ones (Leuctra, Protonemura, Nemoura was formed (Sanchez-Ortega & Azzouz, 1998).
and Isoperla) include approximately 70% of the Euro- All Arctoperlaria families, except Notonemouridae
pean stonefly species. This fauna is relatively well and Styloperlidae, are present in Palaearctic Asia.
known from a taxonomical and faunistical point of view. The family Scopuridae is endemic from this region,
Nevertheless, new species continue to be described. and has a small distribution area and includes eight
Although the European Plecoptera fauna is diverse, species belonging to the genus Scopura Ueno, which
considerable extinctions probably occurred during only occurs in Japan and Korea. Pteronarcyidae (only
Pleistocenic glaciations, because the most important one genus with Asiatic species) and Peltoperlidae
European mountain chains are EW oriented: when ice (four Palaearctic genera) have a Nearctic-Asian
advanced southwards they hindered the movement of distribution. The remaining families show a wide
fauna to the South. Moreover, there have been limited Holarctic distribution and are well represented in the
relationship between European and Asian stonefly- Asian part of the Palaearctic Region, some of them
faunas, as reported above. Thus, faunistic interchanges reaching the Oriental Region.
were only possible through Asia Minor and the East Palaearctic (Asian) Plecoptera diversity is
mountain systems, whereas the Central European much greater than all the remaining continents,
plains acted as a barrier, and only a few taxa could despite the poor and fragmentary knowledge of this
move across the Polar Circle (such as Diura Billberg or area: the exact number of Asian Plecoptera species
Arcynopteryx Klapalek) (Zwick, 1980). is difficult to estimate, but hundreds of new species
High numbers of species exist in the Mediterra- will probably be described in the future. Japan with
nean peninsulas (particularly the Western ones) 306 species is an exception, because stoneflies from
including many endemic species. This is due to the this country have been well studied, while the
presence of mountain systems, which represent an Chinese stonefly fauna is almost unknown (Sivec &
optimal environment for stoneflies, and to the effect Yang, 2001). Zhiltzova (1997, 2001) reported 350
of glaciations, that favoured isolation and speciation species in the former USSR (also including the
processes. Also many species occur in Central European part), and noted (1) a marked impover-
Europe mountainous areas (Alpine or Carpathic) ishment of the fauna towards the North; (2) a high
where cold streams are abundant. Nevertheless, the diversity of species in the Caucasus (3) a certain
post-glacial invasion by widespread species has scarcity of Central Asia fauna, with a high degree of
lowered the number of endemic species. On the endemisms at the specific level concentrated in the
contrary, Northern Europe has low species richness. Himalayan region; (4) a rather poor fauna in the
Finally, no species or a few number of them are East Siberian area, even though rich in genera
present in the Atlantic islands as a consequence of the number and endemic species, and (5) a very rich
low dispersion capacity of stoneflies. fauna in the South Russian Far East. Finally, the
Northern Africa Plecoptera are included in the stonefly-fauna of South-West part of Asia is partly
same seven families present in Europe (Perlidae, unknown; due to the general dry climate of the area,
Perlodidae, Chloroperlidae, Taeniopterygidae, Nemo- new species may be described in the future only
uridae, Capniidae and Leuctridae) and in the same from mountain systems.

123
376 Hydrobiologia (2008) 595:365377

From a zoogeographical point of view, the distribu- Human related issues

tion of Pteronarcyidae, Peltoperlidae and several taxa
within other families demonstrates that these evolved Stoneflies represent a very important component of
in the area that encompasses East and North Asia as rivers and streams, both as biomass and as diversity
well as North America, when the two continents were of ecological roles, acting as primary or secondary
still joined together (Banarescu, 1990). The presence of consumers and as prey for other macroinvertebrates
the genus Perlodes Banks only in Europe and Japan, and fishes, including those of economic importance.
may be explained by its extinction in Siberia during the From a scientific point of view, they have been used
Ice Age (Banarescu, 1990). as biogeographical indicators and in evolutionary
researches (for instance in the insect wings and
respiratory proteins evolution). Their importance as
The Australian Region human food is scarce, although some big species are
included in the diet of some cultures. Stoneflies
Australia currently includes 191 species from 26 have been used for centuries in the sport of fly-
genera and four families. Gripopterygidae is shared fishing, and fishermen have a good general knowl-
with South America while Notonemouridae occurs in edge of them. Plecoptera are frequently employed as
both South America and South Africa. All 191 bioindicators of water quality: the highest values as
Plecoptera species and 24 of the 26 Australian genera indicators of good ecological and environmental
are endemic (Theischinger 1991). Tasmanian Ple- quality are normally assigned to this group. Stone-
coptera includes 47 species, 41 of which are endemic flies do not cause economic damages to human
to the island. Plecoptera can be found throughout crops although some damage to fruit trees and
Australia, except the Northern Territory. Their ornamental plants by adult Taeniopterygidae have
distribution in Western and South Australia is been reported.
restricted, due to the arid conditions. The genera Unfortunately, water quality degradation and
Crypturoperla, Tasmanoperla (Austroperlidae), Tas- physical alteration of streams and rivers have caused
manocerca (Notonemouridae) and Cardioperla a decrease in the number of Plecoptera species,
(Gripopterygidae) are limited to Tasmania. Many particularly those inhabiting lowland rivers of indus-
genera in all families are confined to mainland trialized countries (such as many Central European
Australia. Only the family Gripopterygidae occurs ones), or highly populated countries like the Asiatic
in Western Australia. ones. Taeniopteryx araneoides (Klapalek) and Oe-
New Zealand has 104 described species (McLe- mopteryx loewi (Albarda), once common in large
llan, 2006), and the same four families as Australia. Central Europe rivers, are now extinct (Zwick, 2004).
All genera are endemics to this island system, except These are among the very few documented cases of
Notonemoura and Stenoperla, that are shared with extinction in insects. Many undescribed species,
Australia, even though species of this genera are mainly from Asia, will probably go extinct before
endemic in both countries. A considerable proportion they are discovered (Sivec & Yang, 2001). Never-
of the entire stonefly-fauna is found in the high theless, few Plecoptera species are included in the
precipitation region of the Southern Alps and its official lists of threatened species. Only one
subsidary mountain ranges (McLellan, 2000), and species, the chloroperlid Alloperla roberti Surdick
endemisms are most numerous in South Island. endemic from Illinois (USA), is reported as extinct in
At least one species of Neoperla occurs in New the IUCN Red List of Threatened Species, (IUCN,
Guinea, while no Plecoptera at all are present in New 2004).
Caledonia.
The total Plecoptera species of the Australian Acknowledgements We would like to thank P. Zwick
Region is 295. This number is remarkably small if (Schlitz, Germany) and Ian McLellan (Westport, New
Zealand) for providing material and useful suggestions. B. P.
compared to the Holarctic and Oriental faunas, which
Stark (Mississippi, U.S.A.) and K. W. Stewart (Texas, U.S.A.)
are of entirely different composition, sharing no commented and improved an early version of the paper and are
families with Australia (Zwick, 1981). heartly thanked.

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References Sivec, I. & P. Yang, 2001. Stoneflies of Taiwan within the oriental
stonefly fauna diversity. In Domnguez E. (ed.), Trends in
Aubert, J., 1959. Plecoptera. Insecta Helvetica 1: 1140. Research in Ephemeroptera and Plecoptera. Kluwer Aca-
Banarescu, P. 1990. Zoogeography of Fresh Water. General demic/Plenum Publishers, New York, 401404.
Distribution and Dispersal of Freshwater Animal, Vol. 1. Stark B. P., 2001. A synopsis of the Neotropical Perlidae. In
Aula-Verlag, Wiesbaden. Domnguez E. (ed.), Trends in Research in Ephemerop-
Fochetti, R. & J. M. Tierno de Figueroa, 2004. Plecoptera. tera and Plecoptera. Kluwer Academic/Plenum
Fauna Europaea Web Service. http://www.faunaeur.org Publishers, New York, 405422.
(Fauna Europaea project, ref. EVR1-CT-19992001). Stark, B. P. & R. W. Baumann, 2005. North American Stonefly
Fochetti, R. & J. M. Tierno de Figueroa, 2005. I Plecotteri (Plecoptera) complete list. http://www.mlbean.byu.edu/
europei: distribuzione e stato di conservazione. XX plecoptera/list.asp.
Congresso Italiano di Entomologia. Assisi 1318.VI.2005. Stewart, K. W., 1994. Theoretical considerations of mate
Gullan, P. J. & P. S. Cranston, 2005. The Insects. An outline of finding and other adult behaviors of Plecoptera. Aquatic
Entomology. 3rd ed. Blackwell Publishing Ltd, Oxford. Insects 16: 95104.
Heckman, C. W., 2003. Encyclopedia of South American Theischinger G., 1991. Plecoptera (Stoneflies). In CSIRO (ed.),
Aquatic Insects: Plecoptera. Kluwer Academic Publishers, The Insects of Australia. A textbook for students and
Dordrecht. research workers. Vol. 1, 2nd ed. Melbourne University
Hynes, H. B. N., 1976. Biology of Plecoptera. Annual Review Press, Melbourne, 311319.
of Entomology 21: 135153. Tierno de Figueroa, J. M. & R. Fochetti, 2001. On the adult
Hynes, H. B. N., 1988. Biogeography and origins of the North feeding of several European stoneflies (Plecoptera).
American stoneflies. Memoires Entomological Society Entomological News 112: 130134.
Canada 144: 3137. Tozer, W., 1979. Underwater behavioural thermoregulation in
Illies, J., 1965. Phylogeny and zoogeography of the Plecoptera. the adult stonefly, Zapada cinctipes. Nature (London)
Annual Review of Entomology 10: 117140. 281: 566567.
IUCN, 2004. 2004 IUCN Red List of Threatened Species. Villet, M. H. 2000. The stoneflies (Plecoptera) of South Africa.
http://www.redlist.org. http://www.ru.ac.za./academic/departments/zooento/
Jewett, S. G., 1963. A stonefly aquatic in the adult stage. Martin/plecoptera.html.
Science 139: 484485. Yuzhou, D. & H. Junhua, 2001. Progress on taxonomic study of
McLellan, I. D., 2000. A revision of Cristaperla (Plecoptera: the family Perlidae from China. In Domnguez E. (ed.),
Notonemouridae) and some comments on Notonemouri- Trends in Research in Ephemeroptera and Plecoptera.
dae and its generic groups. New Zealand Journal of Kluwer Academic/Plenum Publishers, New York, 369375.
Zoology 27: 233244. Zhiltzova, L. A., 1997. Zoogeographic features of the Euholog-
McLellan, I. D. 2006. Endemism and biogeography of New natha fauna (Plecoptera) of Russia and adjacent territories
Zealand Plecoptera (Insecta). Illiesia 2: 1523. (within the limits of the former USSR). In Landolt, P. & M.
http://www2.pms-lj.si/illiesia/papers.html. Sartori (eds), Ephemeroptera and Plecoptera: Biology-
Michaelis F. & C. M. Yule, 1988. Plecoptera. In Walton, D. W. Ecology-Systematics. MTL Fribourg, 186192.
& W. W. K. Houston (eds), Ephemeroptera, Megaloptera, Zhiltzova, L. A., 2001. Plecoptera fauna of Capniidae of Russia
Odonata, Plecoptera, Trichoptera. Zoological Catalogue and adjacent territories (within the limits of the former
of Australia, Vol. 6. Canberra, AGPS xi 315 pp (133 USSR). In Domnguez E. (ed.), Trends in Research in
176), (Updated by Andrew A. Calder, CSIRO Entomol- Ephemeroptera and Plecoptera. Kluwer Academic/Plenum
ogy, Canberra, Australia, 2002). Publishers, New York, 423429.
Paulian, R., 1951. Faunes des eaux douces de Madagascar I. Zwick, P. 1980. Plecoptera (Steinfliegen). In Handbuch der
Plecopteres et Megalopteres. Memoires Institut Scientifi- Zoologie, Walter de Gruyter, Berlin, 1115.
que Madagascar 6: 5362. Zwick, P., 1981. Plecoptera. In: Keast A. (ed.) Ecological Bioge-
Rohdendorf, B. B. & A. P. Rasnitsyn, 1980. [Historical ography of Australia, Vol. 2. The Hague, Junk, 11711182.
Development of Insect Class]. Trudy Paleontolo- Zwick, P., 1990. Transantarctic relationships in the Plecoptera.
gischeskogo Instituta 178: 1267 (in Russian). In Campbell I.C. (ed.), Mayflies and Stoneflies, Life
Sanchez-Ortega, A. & M. Azzouz, 1998. Faunistique et History and Biolgy. Kluwer Academic Publishers,
phenologie des Plecopteres (Insecta, Plecoptera) du Rif London, 141148.
marocain (Afrique du nord). Relations avec les autres Zwick, P. 1998. The African Neoperla (Plecoptera: Perlidae)
aires de la region mediterraneenne occidentale. Mitteil- revisited: towards an overdue revision. XIII International
ungen der Schweizerischen Entomologischen Gesellschaft Symposium on Plecoptera, IX International Conference
71: 449461. on Ephemeroptera. Tucuman (Argentina), 92.
Sargent, B. J., R. W. Baumann & B. C. Kondratieff, 1991. Zwick, P., 2000. Phylogenetic System and Zoogeography of the
Zoogeographic affinities of the Nearctic stonefly fauna of Plecoptera. Annual Review of Entomology 45: 709746.
Mexico. Southwestern Naturalist 36: 323331. Zwick, P., 2003. Biogeographical patterns. In Resh V. H. & R.
Shinitshenkova, N. D. 1997. Paleontology of stoneflies. In T. Carde, (eds), Encyclopedia of Insects. Academic Press,
Landolt, P. & M. Sartori (eds), Ephemeroptera and Ple- Elsevier Science, USA, 94103.
coptera, Biology-Ecology-Systematics. MTL Fribourg, Zwick, P., 2004. Key to the West Paleartic genera of stoneflies
561565. (Plecoptera) in the larval stage. Limnologica 34: 315348.

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