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

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some attention is nowadays paid to that problem

(Mogi et al., 1999).

Apart from these marginal relationships between

manhood and water beetles, the applied relevance of

aquatic Coleoptera can be summarized in three main

groups.

Environmental indication

Water beetles, especially Elmidae, are increasingly

gaining recognition as biological indicators for (1)

water quality (saprobial index), (2) habitat types, (3)

biological functionality and (4) species and habitat

conservation (red lists) (see Moog, 2003; Ja

ch et al.,

2005b). A comprehensive list of saprobic valencies of

the Austrian elmids was published by Moog & Ja

(2003).

The IUCN Red List of Threatened Species (see

http://www.redlist.org/ ) contains also a number of

water beetles. However, this list has obviously not

been compiled by water beetle experts and it is

therefore dearly in need of revision. Two New Cal-

edonian dytiscids, Rhantus alutaceus and R.

novaecaledoniae, both listed as extinct in the 2004

IUCN Red List of Threa tened Species, deﬁnitely

suffer from ongoing habitat loss, but were otherwise

still collected during expert ﬁeldwork recently. On

the other hand, some speci es, which have repeatedly

been recorded as being probably extinct, e.g.,

Fig. 2 Global diversity of water beetles by zoogeographic

regions. Borders between realms arbitrary. Species from the

Antarctic Region (ANT) and the Paciﬁc Region (PAC) have

been included to adjacent realms. AU: Australia, AT:

Afrotropical, NA: Nearctic, NT: Neotropic, OL: Oriental,

PA: Palearctic, PAC: Paciﬁc

Fig. 3 Estimation of global diversity of water beetles by

biogeographic regions

Hydrobiologia (2008) 595:419–442 435

123

Hygrobia davidi from China and Hydraena sappho

from Greece (see Audisio et al., 1996), have not been

included in the IUCN Red List so far. Numerous

coleopterists (e.g., P. Audisio, M. Hess, U. Heckes,

M.A. Ja

ch, A. Komarek, H. Schill hammer, H.

Scho

nmann) have vainly searched for the latter spe-

cies in the last two decades.

In general, water beetle communities all around

the world suffer from desertiﬁca tion, irrigation,

eutrophication induced by lifestock, man, and agri-

culture, as well as overall loss of primary habitats.

Species living in aquatic habitats surrounded by

tropical rain forests seem to be especially vulnerable

to deforestation, which is devastatingly effecting

chemical and physical characters of freshwater

biotopes. Furthermore, in numerous water beetles

at least one stage of the life cycle is spent outside

the water. Therefore, any disturbance of the imme-

diate surroundings of an aquatic habitat must be

considered a major threat for its water beetle

communities.

Pest and pest control

Several species of Phytophilous Water Beetles are used

to control water plants that are regarded as pests while

others are destructive to plants of economic importance.

The South American Agasicles hygrophila Selman

& Vogt (Chrysomelidae: Alticinae) was introduced to

several countries on various continents to control

Alligatorweed (Alternanthera). The Neotropical

Cyrtobagous salviniae (Erirhinidae) is used as control

agent for the Water Fern (Salvinia) and has been

introduced, for instance, to India, Thailand and

Australia. Two species of Neochetina, N. eichhorniae

and N. bruchi (Erirhinidae), natives of South America,

were released in the USA and Asia where they are

most effective in reducing the ﬂowering and potential

growth of waterhyacinth (Eichhornia crassipes), the

world’s most important aquatic weed spreading at

an alarming rate. The Waterlettuce Weevil, Neohydro-

nomus afﬁnis (Erirhinidae), from South America was

released in Florida and Australia for biological control

Fig. 4 Global diversity of water beetle families; number of

described species (pale grey/blue) and estimation of unde-

scribed (dark grey/purple) species in 12 beetle families;

paraquatic and terrestrial members not included; 1—

Dytiscidae, 2—Hydraenidae, 3—Hydrophilidae, 4—Elmidae,

5—Scirtidae, 6—Gyrinidae, 7—Psephenidae, 8—Dryopidae,

9—Noteridae, 10—Hydrochidae, 11—Haliplidae, 12—

Helophoridae

436 Hydrobiologia (2008) 595:419–442

123

of Pistia stratiotes.SpeciesofNanophyes (Nanophy-

idae) are feeding in the stems of invasive plant pests,

such as the Water Primrose (Ludwigia).

The Rice Water Weevil, Lissorhoptrus oryzophilus

Kuschel (Erirhinidae), is a most destructive pest of rice

(Oryza sativa). It originates from Amer ica, where it is a

particularly severe pest in Louisiana. Larvae of L.

oryzophilus are responsible for the main damage as

they feed on leafs for a short period and then crawl

down to the roots; they have paired dorsal hooks to

pierce the roots for obtaining oxygen. The fourth larval

instar forms a mud-coated cocoon attached to the roots.

Serious crop losses are reported in all countries where

this species occurs (up to 60% in Japan). Lissorhoptrus

oryzophilus has been spread accidentally to all conti-

nents except Africa and Australia. To Europe (Italy) it

was introduced rather recently (Caldara et al., 2004).

Lissorhoptrus oryzophilus is not only a pest of rice, but

it also attacks many other wild grasses and sedges

(Poaceae and Cyperaceae) which serve as alternative

hosts for adults in or near rice ﬁelds, rendering pest

management most difﬁcult. Furthermore, resistance to

some insecticides has also been reported. In Asia and in

California only parthenogenetic females are known.

Other Erirhinidae, e.g., Lissorhoptrus simplex (Say)

and Afroryzophilus djibai Lyal, are causing heavy

damages too.

Cuisine and medicine

The medical/nutritive relationships between water

beetles and mankind have obviously been more

diverse in the past, but they have survived to some

extent. Even today, various species are used for human

consumption (both as medicine and as confection).

Eating water beetles is still practised in Madagascar,

New Guinea and Asia, for instance in China, where two

genera, Hydrophilus Geoffroy (Hydrophilidae) and

Cybister Curtis (Dytiscidae) are offered for sale in

Guangdong markets and restaurants. Remarkably,

Cybister is more expensive than Hydrophilus (Ja

ch,

2003). Ding et al. (1997) published a list of medicinal

insects in China and mentioned that Cybister was

consumed in order to improve blood circulation.

According to Hoffmann (1947) water beetles are

considered as an anti-diuretic.

More than a century ago, Austrelmis condimen-

tarius (Philippi) (Elmidae), was used as seasoning for

food in South America, and it was reported to have

considerable commercial value (Philippi, 1864; Ne-

tolizky, 1916).

In the early 19th century, Aulonogyrus strigosus F.

(Gyrinidae), was roasted and eaten by Australian

aborigines (Mjo

berg, 1916; Ochs, 1924) and approx-

imately at the same time, gyrinids were used in

Europe as an aphrodisiac for cows and mares

(Netolizky, 1916, 1919; Ochs, 1966).

Rubbing Gyrinidae (Dineutus, Gyrinus, etc.) and

Dytiscidae (Hydaticus, Rhantus) onto young girls

breasts for stimulating their growth is a unique

traditional practice still alive and wide-spread in East

Africa (Kutalek & Kassa, 2005). In their prothoracic

defence glands dytiscids produce steroids resembling

human hormones! However, there is insufﬁcient

evidence to conclude that the chemical defence

mechanism of the water beetles is in fact responsible

for a possible growth of the breasts.

Acknowledgements The following persons are sincerely

thanked for providing useful information and comments: M.

Alonso-Zarazaga, R. Angus, R. Caldara, E. Colonnelli, M.-L.

Jeng, J. Kodada, A. Komarek, A.S. Konstantinov, D. Kovac, C.-F.

Lee, P. Mazzoldi, A.N. Nilsson, I. Ribera, W. Ru

cker, R. Schuh,

W.D. Shepard, C.H.S. Watts and H. Yoshitomi. Special thanks

are due to M. Alonso-Zarazaga, R. Angus, R. Caldara,

E. Colonnelli, M.-L. Jeng, A. Komarek, A.S. Konstantinov,

C.-F. Lee, P. Mazzoldi, A.N. Nilsson and H. Yoshitomi for

providing updated species numbers from their personal databases.

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123

FRESHWATER ANIMAL DIVERSITY ASSESSMENT

Global biodiversity of Scorpionﬂies and Hangingﬂies

(Mecoptera) in freshwater

Leonard C. Ferrington Jr.

 Springer Science+Business Media B.V. 2007

Abstract Seven species and one sub-species of

Mecoptera are aquatic as larvae and pupae. All

aquatic species are classiﬁed in two genera of the

family Nannochoristidae and have very restricted

geographic ranges, with three species conﬁned to

extreme southern South America, three species and

one sub-species conﬁned to New South Wales or

Tasmania in Australia, and one species conﬁned to

the South Island of New Zealand.

Keyword Nannochoristidae

Introduction

The Mecoptera are represented by nine families

(Fig. 1), four of which are restricted to the southern

Neotropical and Australasian regions (Riek, 1954).

Immature stages of many species are unknown or

poorly described, but known larvae of one family,

Nannochoristidae, inhabit ﬂowing water and can be

collected in springs and smaller streams (Byers,

1989; Pilgrim, 1972; Winterbourn & Gregson, 1989).

Adults of aquatic species are not abundant, but when

collected are commonly found near the natal habitat

as a consequence of their low vagility. Adults of

some species in other genera [e.g., Apteropanorpa

tasmanica Carpenter (Apteropanorpidae), Merope

tuber Newman (Meropeidae)] are also predictably

encountered on or among riparian or aquatic vege-

tation, mosses or bryophytes, and the unknown

immature stages, when discovered, may conform to

the deﬁnition of ‘‘aquatic’’ used in this publication.

Consequently, the list of taxa provided here repre-

sents a minimum estimate for aquatic species of

Mecoptera.

Species diversity

Two genera consisting of seven species and one sub-

species comprise the family Nannochoristidae

(Table 1). Nannochoristidae are unusual among Me-

coptera in that they have an elateriform larva, with

slightly reduced sclerotization of abdominal seg-

ments, that probably represent morphological

adaptations to aquatic habits. Larvae of Microchor-

ista philpotti (Tillyard), have been recovere d from

ﬁne organic deposits in areas shielded from current in

small woodland streams of South Island, New

Zealand (Winterbourn, 1982; Winterbourn & Greg-

son, 1989). However, larvae of an unidentiﬁed second

species have been collected repeatedly from lateral

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

que, H. Segers &

K. Martens

Freshwater Animal Diversity Assessment

L. C. Ferrington Jr. (&)

Department of Entomology, University of Minnesota,

Saint Paul, MN 55108-6125, USA

e-mail: ferri016@umn.edu

123

Hydrobiologia (2008) 595:443–445

DOI 10.1007/s10750-007-9133-y

margins of rifﬂe habitats in fast-ﬂowing streams in

Tasmania (Ferrington, unpublished data). Larvae of

M. philpotti are predatory, feeding on larval Chiro-

nomidae and other aquatic Diptera (Winterbourn,

1982). It has been assumed that larvae of all

Nannochoristidae species share similar aquatic and

trophic habits (Hynes, 1984).

Phylogeny and historical processes

Two fundamental competing hypotheses related to the

phylogenetic position of Nannochoristidae exist in

recent literature. An earlier phylogeny of Mecoptera

proposed by Willmann (1987) is shown in Fig. 1.

According to this hypothesis, based primarily on

external and internal morphological characters, the

Nannochoristidae plus the fossil family Robinjohniidae

form the suborder Nannomecoptera, which is the

primitive sister group to the suborder Pistillifera,

containing all the other families of Mecoptera (Will-

mann, 1987). An alternative hypothesis more recently

proposed by Whiting (2002) was developed by

integrating DNA sequence data. Although phyloge-

netic analyses of single genes across Mecoptera and

Siphonaptera was not sufﬁcient to unambiguously

resolve the position of Nannochoristidae, a single

fully resolved topology was obtained when four genes

were analyzed. The analysis supports the view that

Mecoptera as deﬁned by Willmann (1987) is para-

phyletic, but with two major lineages. The total

evidence molecular tree based on 18S + 28S + EF1-

a + COII with all characters weighted equally

resulted in two clades, where the Nannochoristidae

plus Boreidae and Siphonaptera form the basal group

to all remaining families of Mecoptera.

Fig. 1 Phylogenetic

relationships of Mecoptera

families (modiﬁed from

Willmann 1987). Extant

families shown in bold,

extinct families in

parenthesis

Table 1 Aquatic

Mecoptera and their known

distributions

Species Author Distribution

Microchorista philpotti (Tillyard) 1917 New Zealand, South Island

Nannochorista andina Byers 1989 Argentina, Chile

Nannochorista dipteroides Tillyard 1917 Australia: Tasmania

Nannochorista dipteroides eboraca Tillyard 1917 Australia: New South Wales

Nannochorista edwardsi Kimmins 1929 Argentina, Chile

Nannochorista holostigma Tillyard 1917 Australia: Tasmania

Nannochorista maculipennis Tillyard 1917 Australia: Tasmania

Nannochorista neotropica Nava

s 1928 Argentina, Chile

444 Hydrobiologia (2008) 595:443–445

123