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

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One striking aspect is that the areas with the

greatest species diversity are not necessarily those

with the greatest higher-level taxon diversity. The

Neotropical Region, although second highest in

species diversity, is only fourth in generic/

sub-generic diversity after the Nearctic and East

Palaearctic. The subfamily Leptocerinae is most

diverse in generic diversity in the Afrotropical

Region, but records the highest number of species

in the Oriental Region (de Moor 2002a). These

discrepancies might indicate rapid speciation in

recent times after the formation of rainforest ecosys-

tems in the Late Caenozoic.

Areas with great concentrations of endemic species

or with high numbers of species (hotspots of endemism

or hotspots of species diversity) can be identiﬁed as

being in temperate and tropical mountain regions or in

areas of high rainfall. The northern Andes, the Amazon

basin and mountains of southern and southeastern

Brazil are represented in the Neotropics (Flint et al.,

1999); the mountain fold belt of the south western and

southern Cape in South Africa, the central African

highlands and West Africa are represented in the

Afrotropics. Numer ous endemics are found in the

Mediterranean area including Turkey, the southern

Caucasus and the Alps (Malicky, 1983, 1986; Malicky

& Sipahiler, 1993) as well as the Iberian peninsula in

WP. There are smaller centres of endemism in Middle

Asia (from Pamir to the Tyan-Shan mountain ranges);

in the Altai Mountains, mountain ridges of the Far East

and around Lake Baikal. A large amount of endemism

occurs in Korea, China and Japan, although the

Chinese biogeography is too poorly studied to provide

signiﬁcant detail of smaller regions. Large numbers of

local endemic species are found throughout SE Asia,

with signiﬁcant concentrations along the Himalayan

slopes and the meridional mountain ridges of Indo-

china, and also on the islands of Indonesia. The

endemism here correlates with the monsoon intensity

providing the humid conditions necessary for Trichop-

tera development.

Repetitive patterns of shared taxa between bio-

geographical regions suggest possible centres of

origin, routes of dispersal or vicariant events. These

are apparent from the database of generic/subgeneric

distribution and species abundance prepared from the

TWC (see intern et documents and Morse, 1999). In

summarising the foregoing discussions the following

biogeographical patterns are proposed:

1. Temperate Gondwana link between Australian

and Neotropical Regions with some relict survi-

vors at superfamily level (Sericostomatoidea)

also found in the southernmost part of Africa and

Madagascar. Several families and closely related

genera show this pattern.

2. A shared distribution pattern between the Orien-

tal and Holarctic Regions. This has two

alternative routes of dispersal both indicated by

species abundances of closely related taxa; the

ﬁrst via the East Palaeartic to the Nearctic

Region, and the second to the West Palaearctic

Region (the Rhyacophilidae indicate both these

patterns).

3. A tropical Gondwana link between Africa,

Madagascar and India (genera of Hydropsychi-

dae, Hydroptilidae and Dipseudopsidae). There

are also a few instances where Brazilian linkages

with West Africa can be discerned with some

taxa even extending to the Australian region

(family Atriplectididae, genus Ecnomus and

some Leptocerinae and Hydroptilidae genera).

4. A two-way exchange of Neotropic and Nearctic

faunas, indicated by some shared Hydroptilidae

and Leptoceridae genera.

5. A more recent link between the Australian

and Oriental Region with a number of taxa

crossing the Wallace/Weber lines. This is mostly

of tropical origin into temperate Australia, but

there are some temperate Triplectidinae that have

apparently moved into the Oriental realm more

recently.

The discovery of Eocene Baltic Amber fossil

records of Helicophidae and psyllobetine Hydrobios-

idae—groups presently restricted to Australia and

New Zealand (Australian Region) and Chile

(Neotropical Re gion)—suggests a very different

alternative hypothesis. Possibly these presently-

restricted, temperat e Gondwana taxa (Southern

Hemisphere) had a much wider distribution, previ-

ously extending into the Northern Hemisphere, and

became extinct in most of their former range

(Botosaneanu & Wichard, 1983). The Northern

Hemisphere continents in the last 3 million years

(Pleistocene) have been exposed to glaciation events

which have largely left the southern hemisphere

continents unscathed. It is likely that the glaciations

led to the local and regional extinction of many

Hydrobiologia (2008) 595:393–407 403

123

widespread taxa that were unable to disperse. Relict

populations probably survived in more hospitable

climates. Baltic and Dominican Amber fossils also

reveal that certain taxa (Leptoceridae: Triplectidinae)

were more widespread than indicated by present-day

records (Botosaneanu & Wichard, 1981). Some

populations of the more plesiomorphic Philopotami-

dae and Hydrobiosidae—found presently only in

temperate Gondwana regions—may therefore also

represent a relict fauna of previously widespread taxa

that became extinct in other regions in the Northern

Hemisphere (Eskov, 1984).

From the above discussion it is suggested that an

alternative map highlighting regions on present day

continents reﬂect a somewhat different biogeograph-

ical pattern to the generally accepted seven

biogeographical Regions. Detailed distribution pat-

terns of Trichoptera and possibly other amphibiotic

insects cannot be expound ed here, but what is clear is

that Gondwana needs to be divided into several more

closely related biogeographic al regions reﬂecting a

temperate southern region incorporating Australia,

New Zealand, Patagonia and southern Chile and

possibly the southern tip of Africa and southernmost

part of Madagascar; and a tropical region incorpo-

rating parts of the Neotropical, African, Madagascar

and the Indian-part of the Oriental regions. The

proposed map highlighting distinct biogeographic

regions and indicating close relationships illustrates

these patterns (Fig. 4).

The Northern Holarctic Region also presents a

different pattern with at least ﬁve clearly discern-

ible divisions (Fig. 4). Thus, the distribution of

Hydrobiosidae in S. America shows great extra-

tropical diversity in Patagonia similar to that in

New Zealand and Southern Australia, whereas the

slopes of the Andes further to the north have

increasingly less diversity (Schmid, 1989). The

distribution of Trichoptera in Central Asia shows

a distinct gap in Eastern Iran and Western

Afghanistan between the generally western Euro-

pean fauna of Iran and the Oriental fauna of

Afghanistan (Malicky, 1986). The distribution of

the ‘‘Beringian’’ species encompasses Eastern Sibe-

ria, but generally avoids Western Siberia

(Levanidova, 1982).

Fig. 4 Preliminary outline for Trichoptera biogeographical

regions. 1. West Nearctic, 2. East Nearctic, 3. West Palaearctic,

4. Boreal Palaearctic, 5. South Palaearctic, 6. East Palaearctic,

7. Beringian, 8. Neotropical, 9. Afrotropical, 10. Cape South

African, 11. Oriental, 12. Australasian, 13. Patagonian. Similar

colours indicate close biogeographical relationships

404 Hydrobiologia (2008) 595:393–407

123

Human related issues

Trichoptera are abundant in all types of natural

aquatic ecosystems. They are apneustic and rely on

dissolved oxygen for respiration. With a high diver-

sity of species with both case- and shelter-

constructing larvae, they are useful indicators of

organic pollution. Trichoptera have been used exten-

sively in biomonitoring assays with indicator species ,

selected communities or assemblages of species or

more broadly based family level identiﬁcations of

species being used to assess the health status of

aquatic ecosystems. Trichoptera, in conjunction with

other aquatic insect orders, have also been used to

assess aquatic biodiversity EPT (Ephemeroptera,

Plecoptera and Trichoptera) or ETS (Ephemeroptera,

Trichoptera, Simuliidae: Diptera) and habitat

diversity (Hannaford & Resh, 1995; Hewlett, 2000;

de Moor , 2002b).

From an ecol ogical perspective, Trichoptera are

important processors of organic matter and provide a

keystone taxon in the development of the river

continuum concept (RCC) (Vannote et al., 1980). As

processors of organic matter, collectively known as the

functional feeding groups (FFG) of animals, they

display the full array of feeding modes (Cummins,

1973). In lotic water ﬁlter-feeding, shelter-construct-

ing species are important predators of blackﬂy larvae

and help keep population levels of pest species at

acceptable levels (de Moor, 1992). In some instances

excessive construction of shelters by collector/ﬁlter-

feeding larvae have caused the clogging of pipes under

bridges and storm-water pipes, creating problems

when ﬂooding occurs (McCafferty, 1981). Trichoptera

larvae, pupae and adults also form an important link in

the food chain and they have also been used extensively

by trout ﬁshing enthusiasts as models for ‘‘ﬂies’’

(McCafferty, 1981).

Development and man’s activities have consider-

ably reduced Trichoptera species diversity and the

need for remedial measures to prevent further dam-

age to ecosystems and loss of Trichoptera species

have been discussed by Botosaneanu (1981) and

Ivanov (2000). Red data lists for certain countries

have been produced and can be used to manage

ecosystems to protect vulnerable or rare species of

Trichoptera. Anthropogenic development threatens

biodiversity and the value of Trichoptera as important

functional components of aquatic ecosystems,

indicator species of deteriorating conditions and

custodians of environmental protection are realised.

Acknowledgements We would like to thank Dr. Koen

Martens and Ms. Estelle Balian for organising and

facilitating the workshop where discussions of the project

and the sharing of the views of all the disciplines presented by

the invited coordinators was made possible. The Royal Belgian

Institute for Natural Sciences is thanked for providing the

funding for air fare and accommodation that allowed F. C. de

Moor to attend the meeting in Mechelen in October 2005. Prof.

John Morse (Clemson University, USA) and Dr. Alice Wells

(Australian Biological Resources Survey) are thanked for their

critical evaluation of the manuscript and for providing

information and thanks are also due to two anonymous

reviewers for suggestions and corrections. Irene de Moor is

thanked for editing the manuscript in various stages. V. D.

Ivanov is thankful to the Russian Federal Programme for

support of the leading Scientiﬁc Schools project NSH-

7130.2006.4. The Directorate of Eastern Cape Museums is

thanked for supporting F. C. de Moor in the research into

aquatic biodiversity at the Albany Museum.

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Hydrobiologia (2008) 595:393–407 407

123

FRESHWATER ANIMAL DIVERSITY ASSESSMENT

Global diversity of dobsonﬂies, ﬁshﬂies, and alderﬂies

(Megaloptera; Insecta) and spongillaﬂies, nevrorthids,

and osmylids (Neuroptera; Insecta) in freshwater

Matthew R. Cover Æ Vincent H. Resh

 Springer Science+Business Media B.V. 2007

Abstract The insect orders Megaloptera and Neu-

roptera are closely related members of the superorder

Neuropterida, a relict lineage of holometabolous

insects that also includes the Raphidoptera. Megalop-

tera, composed of the fam ilies Sialidae and Corydal-

idae (including subfamilies Chauliodinae and

Corydalinae), has fully aquatic larvae that occur in a

wide variety of lotic and lentic habitats, includi ng

temporary streams. In total, 2 of 17 families of

Neuroptera have aquatic larvae: Nevrorthidae live in

the benthos of fast-ﬂo wing streams and Sisyridae

reside on freshwater sponges. A third family of

Neuroptera, Osmylidae, contains some water-dependent

species that reside under leaves and rocks along the

margins of waterbodies. We recognize 328 extant,

described species of Megaloptera (composed of 116

species of Chauliodinae, 131 species of Corydalinae,

and 81 species of Sialidae) and 73 species of aquatic

Neuroptera (composed of 12 species of Nevrorthidae

and 61 species of Sisyridae). Additionally, we estimate

that 45 species of Osmyl idae are water-dependent,

although the ecology of this group is poorly understood.

Chauliodinae and Corydalidae are both found in the

New World, the Oriental region, and South Africa, but

are absent from Europe, the Middle East, Central Asia,

tropical Africa, and boreal regions. Chauliodinae is

quite speciose in Australia, whereas Corydal inae is

absent. Sialidae is most speciose in temperate regions,

and is absent from tro pical Africa and portions of the

Oriental region. Sisyridae and Osmylidae are nearly

cosmopolitan, but the relict family Nevrorthidae is

limited to Japan, the Mediterranean, and Australia.

The discovery of many new species in recent years,

particularly among Corydalidae in the Neotropics and

China, suggests that our knowledge of aquatic neur-

opterid diversity is far from complete.

Keywords Biodiversity  Species richness 

Megaloptera  Neuroptera  Neuropterida

Introduction

The insect orders Megaloptera and Neuroptera (sensu

stricto), along with Raphidoptera, are closely related

members of the superorder Neuropterida, a relict

lineage of holometabolous insects (Grimaldi &

Engel, 2005). Neuropterida are primarily a terrestrial

group; only the 2 families of Megaloptera and 2 of 17

families of Neuroptera have species with aquatic

larval stages. Neuropterida contains remarkably

heterogeneous taxa, but the relatively low-species

richness (*6,500 species across three orders) and

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

que, H. Segers and

K. Martens

Freshwater Animal Diversity Assessment

M. R. Cover (&)  V. H. Resh

Department of Environmental Science, Policy, &

Management, University of California, Berkeley, 137

Mulford Hall, Berkeley, CA 94720-3114, USA

e-mail: mcover@nature.berkeley.edu

123

Hydrobiologia (2008) 595:409–417

DOI 10.1007/s10750-007-9035-z

relict distributions suggest that the group was much

more speciose and cosmopolitan in the past (Aspo

ck,

2002).

A general account of the biology of the Megalop-

tera and/or Neuroptera is given by New and Theis-

chinger (1993) and New (1986, 1989). The biology of

only a few species of Megaloptera and aquatic

Neuroptera have been described in detail, with most

of the information published on species from Europe,

North America, and Japan.

The Megaloptera is com posed of two extant

families, the Corydalidae and the Sialid ae (alderﬂies).

Corydalidae is further subdivided into the subfamilies

Corydalinae (dobsonﬂies) and Chauliodinae (ﬁsh-

ﬂies), although some have regarded these taxa as

distinct families (e.g., Aspo

ck et al., 1980; Theisch-

inger, 1983). Dobsonﬂy larvae are commonly known

as hellgrammites (although this term is sometimes

applied to larval Chauliodinae or, in a more general

sense, to all large benthic insects used by anglers as

bait). Megalopteran larvae have been reported in a

wide variety of aquatic habitats. Most Corydalidae

are found in fast-ﬂowing rifﬂes under gravel and

cobble, but some corydalids are found on snags and

sand in low-gradient streams. Some species of

Chauliodinae have been reported from lentic habitats,

including ponds and swamps (e.g., Dolin & Tarter,

1981). Sialidae are also found in a wide range of

freshwater habitats, both lotic and lentic, and are

usually associated with soft sediments. Megalopteran

larvae have also been reported from several remark-

able locations that suggest adaptations to otherwise

inhospitable habitats. For example, the larvae of both

Sialidae and Chauliodinae have been reported in

water-ﬁlled vegetation, including tree holes and the

purple pitcher plant (Sarracenia purpurea) (Fashing,

1994; Hamilton et al., 1996; Pittman et al., 1996).

Five species of ﬁshﬂies from California (USA)

representing three genera (Neohermes, Protochauli-

odes, and Dysmicohermes), and two species of

dobsonﬂies (Corydalus luteus and C. afﬁnis) in the

Neotropics have been found in dry streambeds

(Evans, 1972; Contreras-Ramos, 1998). Larvae of

Neohermes dig burrows beneath large rocks as

temporary streams in California dry up, where they

remain for 6 months or more until water begins to

ﬂow. Many ﬁshﬂy larvae possess a pair of highly

modiﬁed abdominal spiracles to facilitate aerial

respiration (Fig. 1), whereas dobsonﬂy larvae possess

ventral tufts of tracheal gills. Co-occurring species of

Megaloptera can be ecologically separated by differ-

ences in larval habitat related to substrate or ﬂow

conditions (Evans, 1972; Hayashi, 1989).

Megaloptera larvae are generalist predators of

small invertebrates, and have also been observed to

scavenge and cannibalize. Larvae of Protohermes

grandis rem ained relatively stationary under stones in

rifﬂes for many days at a time, presumably acting as

sit-and-wait predators (Hayashi & Nakane, 1989).

The life cycles of Megaloptera are usually 1–

2 years, although Chauliodinae in cold or seasonally

dry streams may live up to 5 years (Evans, 1972).

Larvae occupying permanent water bodies crawl to

the banks of the stream to build pupal cham bers in the

soil or leaf litter, while larvae in temporary streams

pupate under large stones after streamﬂow ceas es.

Adult Megaloptera live for approximately 1 week

and do not feed, although some have been reported to

drink water or sugar solution s. Adult Corydalidae are

large and conspicuous, especially male dobsonﬂies in

the genera Corydalus and Acanthacorydalis that

possess enormous scimitar-like mandibles. These

mandibles are of unknown function, but are likely a

secondary sexual characteristic selected for male–

male competition or female choice. Females oviposit

Fig. 1 (A) Habitus of freshwater Megaloptera: Neohermes

ﬁlicornis,(B) Habitus of freshwater Neuroptera: Sisyra vicaria

410 Hydrobiologia (2008) 595:409–417

123

on branches, logs, or stones overhanging the stream,

so that freshly hatched larvae fall directly into the

water.

The Neuroptera (lacewings) are primarily a ter-

restrial order. Larval neuropterans all posses sucking

mouthparts composed of modiﬁed maxillae and

mandibles (Fig. 1). Two families have aquatic larvae:

Sisyridae (spongillaﬂies) and Nevrorthidae (often

incorrectly spelled Neurorthidae).

Sisyridae larvae are obligate predators of fresh-

water sponges (Spongillidae), and possibly Bryozo-

ans as well (Parﬁn & Gurney, 1956; Steffan, 1967;

Resh, 1976). Like the Megaloptera, eggs are laid in

small clusters on vegetation overhanging the water.

Hatchling larvae ente r the water and swim until they

ﬁnd a suitable host, where they remain until the end

of the third instar (New, 1986). Pupation occurs on

vegetation, and adults remain near the water for

mating and oviposition. Sisyrids are usually univol-

tine, although multiple cohorts can be present (Resh,

1976).

Nevrorthidae, once considered part of Sisyridae, is

now recognized as a distinct family (Zwick, 1967) or

even suborder (i.e., Nevrorthiformia; Aspo

ck et al.,

2001). Nevrorthid larvae have been found in the

benthos of fast-ﬂowing streams and are believed to be

fully aquatic (Zwick, 1967), although the habits and

morphology have not been described for many

species (Dudgeon, 1999).

Although little is known about the ecol ogy of the

neuropteran family Osmylidae, species in the sub-

families Osmylinae and Kempyninae have been

strongly associated with freshwater habitats (New,

1986). The most well-studied species is Osmylus

fulvicephalus, which is widespread in Europe. Larvae

are found under stones, moss, and leaf litter along the

margins of streams, where they probe the mud in

search of small arthropods (Ward, 1965). Although

not considered aquatic or even subaquatic, diapausing

larvae on the stream bank can survive immersion by

rising ﬂoodwaters. Several other species of Osmyli-

nae and Kempyninae have been found in wet or moist

zones along streams (New, 1986). For the purposes of

this review of freshwater biodiversity, we consider

Osmylinae and Kempyninae to be water-dependent

because they may be restricted to the margins of

aquatic habitats. Other osmylids are known to prefer

terrestrial habitats, including the Australian subfam-

ilies Porisminae and Eidoporisminae, which are

found under the bark of Eucalyptus trees located far

from water (New, 1986). The speciose subfamily

Stenosmylinae most likely has terrestrial larvae as

well (New, personal communication).

The objectives of this chapter are to enumerate the

known worldwide speci es richness of Megaloptera

and aquatic Neuroptera, examine patterns of bioge-

ography and endemicity, and relate these patterns to

what is known of the phylogeny of the groups . The

discovery of many new species in recent years,

particularly among the Corydalidae in the Neotropics

and China, suggests that our knowledge of aquatic

neuropterid diversity is far from complete.

Species diversity

For the Mega loptera we recognize 328 extant,

described species, composed of 116 species of

Chauliodinae, 131 species of Corydalinae, and 81

species of Sialidae (Table 1). For the Neuroptera we

recognize 73 aquatic species, including 12 species of

Nevrorthidae and 61 species of Sisyridae, and 45

species of water-dependent Osmylidae. We have

included all described species in the published

literature, based on the latest taxonomic revisions

(e.g., Contreras-Ramos, 1998; Liu & Yang, 2006a, b),

species lists (e.g., Penny et al., 1997; Con treras-

Ramos, 1999; Oswald, 2006), and distributional

studies (e.g., Bowles, 2006). We excluded, however,

several unnamed species that are only known from

female specimens. Species richness values are

undoubtedly underestimates of true richness, espe-

cially for the tropical and subtropical taxa.

The past quarter century has seen the fastest rate of

new species descriptions of Megaloptera, most nota-

bly in China (by C. Yang, D. Yang, and X. Liu),

Australia (by G. Theischinger), and the Neotropics

(by A. Contreras-Ramos). Although the Megaloptera

of the Holarctic are believed to be well documented,

further work in tropical areas, especially south-east

Asia and portions of South America, is expected to

increase the known richness of Megaloptera to at

least 370 species, and possibly more than 400 species.

There is evidence that approximately 25 undescribed

species of Megaloptera are present in China (X. Liu,

personal communication) and perhaps 15 additional

species are present in the Neotropics (A. Contreras-

Ramos, personal communication).

Hydrobiologia (2008) 595:409–417 411

123

China and south-east Asia are major centers of

species richness of Corydalinae and Chauliodinae

(Table 1, Fig. 2). In China alone, there are 40

described species of Corydalinae and 41 species of

Chauliodinae, the majority of these from the Oriental

region of China. Additional species have been

Table 1 Species richness of Megaloptera and aquatic Neuroptera worldwide and by biogeographic region

PA NT NA AT OL AU PAC ANT Total

freshwater

species

FW WDpt FW WDpt FW WDpt FW WDpt FW WDpt FW WDpt FW WDpt FW WDpt FW WDpt

Chauliodinae 11 0 6 0 18 0 8 0 56 0 23 0 0 0 0 0 116 0

Corydalinae 5 0 54 0 4 0 1 0 71 0 0 0 0 0 0 0 131 0

Sialidae 28 0 10 0 24 0 1 0 8 0 4 0 0 0 0 0 81 0

Total

Megaloptera

44 0 70 0 46 0 10 0 135 0 27 0 0 0 0 0 328 0

Nevrorthidae 9 0 0 0 0 0 0 0 1 0 2 0 0 0 0 0 12 0

Sisyridae 9 0 25 0 6 0 8 0 8 0 6 0 1 0 0 0 61 0

Osmylidae 0 16 0 4 0 0 0 0 0 9 0 15 0 0 0 0 0 45

Total

Neuroptera

34 16 29 4 6 0 8 0 9 9 23 15 1 0 0 0 118 45

‘‘WDpt’’ represents the number of water-dependent species. ‘‘FW’’ refers to the total number of freshwater including fully

aquatic + water dependent species. PA: Palaearctic; NA: Nearctic; NT: Neotropical; AT: Afrotropical; OL: Oriental; AU:

Australasian; PAC: Paciﬁc & Oceanic Islands, ANT: Antarctic

Fig. 2 Distribution of freshwater (aquatic + water dependent)

Megaloptera (M) and Neuroptera (N) species and genera by

zoogeographical region (species number/genus number). PA—

Palaearctic; NA—Nearctic; NT—Neotropical; AT—Afrotrop-

ical; OL—Oriental; AU—Australasian; PAC—Paciﬁc & Oce-

anic Islands, ANT—Antarctic

412 Hydrobiologia (2008) 595:409–417

123

identiﬁed but not yet described, with over 90 species

of Corydalidae having been found in China (X. Liu,

personal communication). China has been well

surveyed relative to the rest of the Oriental region,

so given the high endemicity and restricted distribu-

tions of the Chinese species we expect that many

undescribed, endemic species also occur in India and

Southeast Asia.

A second region of high diversity in Corydalinae is

Amazonia and the Andes Mountains. For example, 18

species of Corydalinae are known from Venezuela

and 16 species are known from Brazil. Chauliodinae

is rather sparse in the Neotropic s, being restricted to

Chile (with one possible undescribed species in

Brazil, A. Contreras-Ramos, personal communica-

tion), but is much more speciose in Australia (23

species) and the Paciﬁc Coast of the US (11 species).

Generic richness follows roughly the same trends as

species richness, although the Nearctic (6 genera) is

the richest bioregion with respect to the Chauliodinae

(Table 2).

Sisyridae and Nevrorthidae have rece ived little

recent attention, and the extent of our knowledge of

the diversity of these families is unclear. The

Sisyridae are particularly speciose in the Neotropics,

with 15 species in Brazi l. The Nevrorthidae have an

extremely disjunct distribution, being limited to the

Mediterranean, Japan, Taiwan, and Australia, and are

clearly a relict taxonomic group.

The number of water-dependent Osmylidae is

highly uncertain because the larvae and biology of

many genera are completely unknown. If all members

of the Osmylinae and Kempyninae occur along

stream margins, the 45 described species in these

subfamilies would be categorized as water-de pendent

osmylids. Water-dependent osmylids are particularly

speciose in Australia (13 endemic species) and east

Asia (16 species) (New, 1983).

Phylogeny

The Neuropterida are a primitive lineage possessing

many apparently pleisomorphic characters of holo-

metabolous insects, such as mobile pupae in

Megaloptera and Raphidoptera. The monophyly of

Neuropterida is generally accepted based upon

inconspicuous characters such as thoracic and

abdominal sutures and the musculature of the

ovipositor (Kristensen, 1999; Aspo

ck, 2002). Mono-

phyly is also supported by molecular (18S rDNA)

data (Whiting, 2002) and combined morphological/

molecular analyses (Wheeler et al., 2001). Neuropte-

rida is hypothesized to comprise the sister group to

Coleoptera, based on morphologic characters (i.e., the

absence of cruciate cervica l muscles, modiﬁcation of

the female terminalia, and characters of the hind-

wing-base; Kristensen, 1999; Hornsch emeyer, 2002)

and some molecular studies (e.g., Wheeler et al.,

2001). Other molecular data, however, does not

support this hypothesis because the monophyly of

Coleoptera has been questioned (Whiting, 2002).

The monophyly of each of the three orders of

Neuropterida is generally accepted, although there is

less certainty about the Megaloptera than the other

two orders. Autapomorphies of Neuropt era are quite

conspicuous, and include complex sucking mouth-

parts and closure of the midgut in the larvae.

Table 2 Generic richness of Megaloptera and aquatic or water-dependent Neuroptera worldwide and by biogeographic region

PA NA NT AT OL AU PAC ANT Total freshwater genera

Chauliodinae 2 6 3 3 4 2 0 0 15

Corydalinae 3 2 3 1 5 0 0 0 9

Sialidae 2 1 1 1 3 2 0 0 7

Total Megaloptera 7 9 7 5 12 4 0 0 31

Nevrorthidae 2 0 0 0 1 1 0 0 3

Sisyridae 1 2 2 2 2 2 1 0 4

Osmylidae 4 0 2 0 1 3 0 0 7

Total Neuroptera 7 2 4 2 5 6 1 0 14

PA: Palaearctic; NA: Nearctic; NT: Neotropical; AT: Afrotropical; OL: Oriental; AU: Australasian; PAC: Paciﬁc & Oceanic Islands,

ANT: Antarctic

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