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

Подождите немного. Документ загружается.

FRESHWATER ANIMAL DIVERSITY ASSESSMENT

Global diversity of craneﬂies (Insecta, Diptera: Tipulidea

or Tipulidae sensu lato) in freshwater

Herman de Jong Æ Pjotr Oosterbroek Æ

Jon Gelhaus Æ Herbert Reusch Æ Chen Young

 Springer Science+Business Media B.V. 2007

Abstract The Tipulidae s.l.—craneﬂies—are one of

the largest groups of the Diptera containing over

15,270 valid species and subspecies. The immatures of

the majority of species live in aquatic or semiaquatic

habitats. Some aquatic species live entirely submerged

and lack functional spiracles, others come to the

surface to take oxygen by using spiracles positioned at

the end of the abdomen. Semiaquatic species occur in a

wide range of habitats. The semiterrestrial and

terrestrial larvae live in environments that are moist

or at least humous. All adult craneﬂies are terrestrial.

Conﬂicting hypotheses on the phylogenetic position of

the Tipuloidea within the Diptera continue to exist:

some authors consider them to represent one of the

oldest lineages of the Diptera, others suppose a close

relationship to the Brachycera, the true ﬂies. Current

systematic knowledge of the Tipuloidea indicates that

the Palaearctic region contains the highest number of

genus-group taxa, while the Neotropical region has the

highest number of species and subspecies. The Afro-

tropical and Australasian regions are relatively poor

respectively in genera and subgenera and in species

and subspecies. The oldest fossils that represent the

Tipuloidea date back to the Lower Triassic at about 240

million years. Present-day general distri bution patterns

of many higher taxa of Tipuloidea probably have a

Pangean or Gondwanan origin.

Keyword Freshwater  Biodiversity  Tipulidae 

Diptera  Zoogeography  Review

Introduction

Systematic position

The Tipulidae sensu lato, also known as the Tipuloidea

and vernacular, as crane ﬂies, constitute one of the

largest groups of the Diptera, with more than 15,270

currently recognized valid species and subspecies that

are contained in 525 genera and subgenera

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

que, H. Segers &

K. Martens

Freshwater Animal Diversity Assessment.

Electronic supplementary material The online version of

this article (doi:10.1007/s10750-007-9131-0) contains supple-

mentary material, which is available to authorized users.

H. de Jong (&)  P. Oosterbroek

Section Entomology, Zoological Museum Amsterdam,

Plantage Middenlaan 64, Amsterdam 1018 DH,

The Netherlands

e-mail: Hjong@science.uva.nl

J. Gelhaus

The Academy of Natural Sciences, 1900 Benjamin

Franklin Parkway, Philadelphia, PA 19103-1195, USA

H. Reusch

BAL – Bureau of Applied Limnology and Landscape

Ecology, Wellendorf 30, 29562 Suhlendorf, Germany

C. Young

Carnegie Museum of Natural History, 4400 Forbes

Avenue, Pittsburgh, PA 15213, USA

123

Hydrobiologia (2008) 595:457–467

DOI 10.1007/s10750-007-9131-0

(Oosterbroek, 2005). The Tipuloidea, as they will be

referred to in the remainder of this text, occur

worldwide, ranging from the arctic to equatorial forests

and from the intertidal zone to over 5,600 m in certain

high mountain ranges (Alexander & Byers, 1981).

They belong to the lower Diptera, but their phyloge-

netic position within the lower Diptera has not yet been

established satisfactorily. Traditionally, the Tipuloidea

were regarded as representing one of the most prim-

itive lineages of the Diptera, if not the most primitive

lineage. Authors supporting this hypothesis are, among

others, Wood & Borkent (1989) and Michelsen (1996) .

An opposing hypothesis claims that the Tipuloidea

belong to the mor e derived lower Diptera and are

closely related to the Brachycera, the higher ﬂies. This

hypothesis, which is based on a digital phylogenetic

analysis of all then available evidence, was ﬁrst

proposed by Oosterbroek & Courtney (1995). Ongoing

research, including phylogenetic analyses of fossil

information, aims to solve this controversy. The mono-

phyly of the Tipuloidea is supported by a sequence of

apomorphies associated with the different life stages

(Wood & Borkent, 1989; Oosterbroek & Theowald,

1991, Oosterbroek & Courtney, 1995). Currently, the

Tipuloidea are considered to contain the families

Cylindrotomidae, Limoniidae, Pediciidae, and Tipuli-

dae sensu stricto.

Habitus

Adult Tipuloidea can be recognized by the presence of

two complete anal veins in the wing, the lack of ocelli,

and the presence of a V-shaped transverse suture on the

mesothorax. They are typically slender-bodied ﬂies,

with long antennae, wings, legs, and abdomen (Fig. 1).

The legs fall off easily. Sizes range from very small

species with a wing length of about 2 mm (e.g.,

Tasiocera) up to large species with wing lengths up to

40 mm (e.g., Holorusia and Leptotarsus). Larvae of

the Tipuloidea can be recognized by the hemicepha-

lous, retractible head capsule, and the usually

metapneustic respiratory system (rarely apneustic).

Morphology

Adult

Head. Usually with short and inconspicuous rostrum

in the Cylindrotomidae, Limoniidae and Pediciidae,

rostrum were well-developed in Tipulidae s.str. and

often with a nasus at tip. Rostrum elongate and

slender in Elephantomyia, Helius, Toxorhina, in some

species as long as head and thorax combined;

mouthparts elongate in Geranomyia, forming long

rostrum-like structure. Mouthparts are usually well-

developed, palpi generally ﬁve-segmented, although

the number of segments can be reduced. Ultimate

palpal segment short in most Cylindrotomidae,

Limoniidae and Pediciidae, elongate and slender in

Tipulidae s.str. Antennae usually long and slender,

consisting of 14–16 segments remarkably elongate—

up to four times the body length, e.g., in certain

species of Hexatoma, Rhabdomastix, Leptotarsus,

and Megistocera. Flagellomeres usually cylindrical in

both sexes, but pectinate in (males of) e.g., Dic-

ranomyia (Zelandoglochina), Clytocosmus,

Ctenophora and allies, Elnoretta, Platyphasia, and

Ptilogyna s.l. Compound eyes large, often touching

or nearly touching on ventral side of head. Eyes

usually bare, but provided with macrotrichia in

between ommatidia in most Pediciidae (absent in

Nipponomyia). Ocelli absent.

Thorax with well-developed mesothorax; protho-

rax and metathorax small were reduced to strip-like

elements. Legs long and slender, tips of tib iae

without spurs or with one or two spurs. Tarsal claws

simple or toothed with one tooth or several teeth,

arolium between claws. Halter with long shaft and

distinct knob. Wing usually well-developed but

sometimes reduced in female or both sexes,

Fig. 1 Tipula (Acutipula) maxima Poda, 1761, male. Euro-

pean species with larva in marshy soil or freely aquatic in small

rivulets, adult in wet woodland along streams. Germany, photo

rgen Peters

458 Hydrobiologia (2008) 595:457–467

123

especially in species living in isolated localities like

islands and mountain ranges, or in cold-ada pted

species. Venation variable, characteristic for major

taxonomic units. Always two complete anal veins are

present (excepting Stibadocerella, the species of

which have one complete anal vein).

Abdomen long or very long, usually slender.

External sexual organs are at the tip of the abdomen.

Terminalia of male with structural adaptation s of the

last tergites, sternites and their genital processes.

Structures of male terminalia considered species-

speciﬁc. Female termin alia generally forming an

ovipositor consisting of sclerotized, blad elike cerci

and hypogynial valves, usually less characteristic for

individual species, but showing d ifferences at level of

higher taxonomic groups.

Larva

Elongate, cylindrical, tapering toward anterior tip,

posterior end often more or less truncate, with

ﬂattened spiracular disc. Head capsule usually dis-

tinct, retracted into anterior thoracic segments, deeply

incised ventrally and often dorso-laterally more

reduced. Abdominal segments smooth, or with trans-

verse rows of hairs. Creeping welts or ﬂeshy

projections sometimes present. Spiracular disc usu-

ally glabrous and surrounded by lobe-like

projections. Membranous anal lobes usually present.

Pupa

Pupa obtect, elongate. Eyes prominent. Mesothoracic

horns usually simple, ranging from short sessile to

very elongate. Antennal sheaths long. Tarsal sheaths

arranged side by side, not superimposed, lateral tarsal

sheaths partly covered by wing sheaths. Abdomen

parallel-sided or almost so, more or less smooth, or

provided with welts or spines; prominent marginal or

other abdominal spines especially in Limnophilinae

and Tipulidae s.str. Abdominal end truncate in male,

pointed in female. Anal segment usually with spines.

Identiﬁcation

There are many publications available for the iden-

tiﬁcation of adult Tipuloidea, although they are rather

scattered in the literature. Good starting points are

Alexander & Byers (1981) for the Nearctic region,

Dienske (1987), Mannheims (1951–1968), Peus

(1952), Savchenko (1961–1983 (Fauna SSSR),

1982–1986 (Fauna Ukrainy), 1983, 1989), Savchenko

& Krivolutskaya (1976), and Theowald (1973–1980)

for the Palaearctic region, Oosterbroek (1998) for

Malaysia, Alexander (1929) for the Neotropical

region, Dobrotworsky (1968–1974) and Theischinger

(1996) for Australia, and Alexander (1956, 1964) and

Wood (1952) for the Afrotropical region.

The identiﬁcation of pre-adult Tipuloidea can be

attempted with the aid of Alexander & Byers (1981),

Byers (1996), and Gelhaus (1986, 2000, 2002) for the

Nearctic region, Brindle (1960, 1967), Brinkmann

(1997), Gelhaus & Byers (1994), Hofsvang (1997),

Peus (1952), Reusch & Oosterbroek (1997), and

Theowald (1957, 1967) for the Palaearctic region,

Young (2004) for Malaysia, and Wood (1952) for the

Afrotropical region.

Life history

As in all holometabolic insects, the Tipuloidea have

four life stages: egg, larva, pupa and adult. The egg

stage usually lasts 6–14 days and is followed by four

larval stages of variable duration. The pupal stage

lasts 5–12 days and the adult usually is short-lived.

The majority of species living in temperate environ-

ments are univoltine or bivoltine, i.e., they have one

or two generations per year. The life cycle of others

can be as short as six weeks, or, in the case of some

arctic species, can last up to ﬁve years. The long life

cycle of arctic species principally is caused by the

shortness of the arctic summer, rather than by the low

temperatures (Pritchard, 1983).

Current main habitats

Adults of the Tipuloidea are exclusively terrestrial,

whereas the preadult stages of the majority of species

are aquatic or semiaquatic. Adult crane ﬂies usually

occur in moist environments in the damp vegetation

along the borders of lakes and streams. Certain

species or higher taxonomic groups are adapted to

open meadows, fairly dry rangelands, and even

deserts.

Hydrobiologia (2008) 595:457–467 459

123

Larvae of crane ﬂies occupy a wide variety of

habitats, ranging from strictly aquatic to entirely

terrestrial. Fully aquatic larvae can occur in fresh-

water, especially in rapidly ﬂowing streams, or in

brackish water or intertidal zones. Some of these

species live entirely submerged, are without func-

tional spiracles, and take oxygen by gas-exchange

through the cuticle. Others come to the surface for

oxygen by using the spiracles positioned at the

posterior end of the body.

A numberof aquatic species occur in the benthic zone

(epibenthic to hyporheic) of fast ﬂowing rivers and

streams. Specialized aquatic species occur on wet cliff

and rock faces, piles or bridge piers in or beneath a scum

of algal growth, mosses, or liverworts. Others are marine

and live on rocks or earth in algal mats that become

submerged by the tide. Species are known from cold

springs, while others live in phytotelmata, such as water-

ﬁlled tree-holes, or water-containing axils of leaves of

such plants as Bromeliaceae and Liliaceae.

Semi-aquatic species occur in wet or saturated

mats or cushions of mosses and liverworts, in leaf

drift and rich organic mud along the edges of streams,

lakes, ponds, and other waterbodies. Semi-terrestrial

and terrestrial species are known from moist to

saturated humous soils in woods, meadows, and open

lands. Some species live in decaying plant material in

various stages of putrefaction, in fungi that are often

in advanced stages of decay, and in dead wood,

ranging from wet, decayed wood to relatively sound

wood. A few species are leafminers, some live in dry

moss cushions, and others can be found in relatively

dry soil (Alexander, 1931; Alexander & Byers, 1981).

The majority of larvae feed on decaying plant

material and its associated microﬂora (algae and

fungi), others on mosses and liverworts. Species of

the Pediciinae (Pediciidae) and most Limnophilinae

(Limoniidae) are predacious.

Pupation usually takes place in dryer places near

the larval habitats.

Species diversity

Numbers of species and subspecies

An overview of the genera, subgen era and number of

species and subspecies of the Tipuloidea for each of

the biogeographical regions is given in Tables 1, 2.

The information is derived from Pjotr Oosterbroek’s

Catalogue of the Craneﬂies of the World, the contents

of which is based on the examination of virtually the

entire world literature dealing with the taxonomy,

systematics, faunistics and biogeography of the

Tipuloidea. The classiﬁcation employed in the tables

is in accordance with the classiﬁcation used in the

Table 1 Numbers of species and subspecies of the families of the Tipuloidea according to biogeographic region

Family PA NA NT AT OL AU Total world # valid species

Cylindrotomidae 27 8 1 0 31 8 71

Limoniidae 1579 885 2728 1038 2324 2114 10430

Pediciidae 202 144 13 0 132 5 489

Tipulidae sensu stricto 1280 573 805 339 925 385 4188

Higher level classiﬁcation of the Tipuloidea according to Oosterbroek (2005). All data derived from Oosterbroek (2005)

PA: Palaearctic; NA: Nearctic; NT: Neotropical; AT: Afrotropical; OL: Oriental; AU: Australasian

Table 2 Numbers of the genera and subgenera of the families of the Tipuloidea according to biogeographic region

Family PA NA NT AT OL AU Total world # valid genera

Cylindrotomidae 6 4 1 0 7 2 9

Limoniidae 137 104 121 90 121 116 314

Pediciidae 17 13 3 0 17 1 26

Tipulidae sensu stricto 45 38 36 23 45 30 115

Higher level classiﬁcation of the Tipuloidea according to Oosterbroek (2005). All data derived from Oosterbroek (2005)

PA: Palaearctic; NA: Nearctic; NT: Neotropical; AT: Afrotropical ; OL: Oriental; AU: Australasian

460 Hydrobiologia (2008) 595:457–467

123

Catalogue of the Craneﬂies of the World. The

catalogue is available on-line via http://ww w.science.

uva.nl/zma/.

Table 2 shows that the Cylindrotomidae contain

71 valid species and subspecies, the Limoniidae

10,430 species-group taxa, the Pediciidae 489, and

the Tipulidae s.str. 4,188. As these numbers show ,

differences in species richness between the major

groups are consider able. The numbers of species and

subspecies included in the various genera and

subgenera of the Tipuloidea also show striking

differences. A number of genera are monotypic, e.g.,

in Cylindrotomidae: Stibadocerina, Limoniidae: Ay-

maramyia, Gonomyopsis, Jivaromyia, Quathlambia,

etc., Pediciidae: Savchenkoiana, Tipulidae s.str.:

Austrotipula, Elnoretta, Euvaldiviana etc. Other

genera and subgenera hold hundre ds of species,

e.g., Limoniidae: Cheilotrichia, Erioptera, Gonom-

yia, Idiocera, Molophilus, Ormosia, Teucholabis,

Austrolimnophila, Gynoplistia , Hex atoma, Dic-

ranomyia, Geranomyia, Limonia, Pediciidae:

Dicranota, Tipulidae s.str.: Dolichopeza, Nephro-

toma, Tipula. These divergent numbers could be

related to disparities in the ages of the highe r taxa

and the speciation rates in the different taxonomic

groups. Otherwise, variance could be an artifact of

the current traditional classiﬁcation with more

phylogenetic studies needed to distinguish between

natural and apparent species diversity.

More than 11,000 of the presently known 15,250

plus species and subspecies of the Tipuloidea were

described between 1910 and 1981 by a single

author, the renowned Charles P. Alexander, and

certainly many more species still await description.

Virtually all currently known species and subspecies

are recognized on morphological traits, and re-

examination of taxon clusters containing cryptic

species using molecular techniques is expected

to increase the number of recognized species

considerably.

Phylogeny and historical processes

Fossil record

The oldest fossils of Tipuloidea, and of Diptera as

such, date back to the Lower Triassic Longtanian of

the Gre

Voltzia Formation of Arzviller in France,

which is dated at 240 Mya (Krzeminski & Krze-

minska, 2003). Other Triassic ﬁnds include fossils

from the Ladinian-Carnian Madygen Formation in

Kyrgyzstan, dated at 230–225 Mya and the Norian-

Rhaetian Tologoi Formation in Kazakhs tan, dated

216–208 Mya (Shcherbakov et al. 1995). Since the

Triassic, a continuous record of fossil Tipuloidea is

available through the different geological epochs.

The literature dealing with the taxonomy of fossil

Diptera up to the early 1990’s is summarized in the

Catalogue of the fossil ﬂies of the World

by Neal L.

Evenhui

(1994).

Phylogeny

Published hypotheses about the phylogeny of the

Tipuloidea include Stary (1992) and Oosterbroek &

Theowald (1991). Stary (1992) studied the relation-

ships at the family level within the Tipuloidea, using

characters of the adults. He concluded that the

monophyletic Limoniidae are the sistergroup of a

clade containing the Pediciidae, Cylindrotomidae,

and Tipulidae s.str., of which the Pediciidae are the

sister group of the Cylindrotomidae + Tipulidae

s.str. Oosterbroek & Theowald (1991) studied the

then available information on pre-adult Tipuloidea at

the genus level. According to their results, the

Pediciidae, Cylindrotomidae and Tipulidae are mono-

phyletic groups, while the Limoniidae are

paraphyletic. Part of the Limoniidae are considered

to form a monophyletic clade together with the

Cylindrotomidae and Tipulidae, which are regarded

as sistergroups (Fig. 2).

The results of these studies by Stary and Oosterb-

roek & Theowald were primarily based on

information on species occurring in the northern

hemisphere, and their hypotheses need to be tested by

incorporating additional taxa of the Southern

hemisphere.

Present distribution and main areas of endemism

Distribution

Figure 3 plots the number of genera and subgenera

and the number of species and subspecies of the

Tipuloidea for each of the biogeographic regions.

Hydrobiologia (2008) 595:457–467 461

123

According to the present classiﬁcation, these numbers

are the following: Palaearctic region, 205 genera and

subgenera, 3088 species and subspecies; Oriental

region, 190 and 341 2; Nearctic region, 159 and 1610;

Neotropical region, 161 and 3547; Afrotropical

region, 113 and 2512; Australasian region, 149 and

1377.

Our present understanding of the Tipuloidea shows

that the Palaearctic region is the richest region for the

taxa at the genus-group level (205), while the

Neotropical region contains the highest number of

species and subspecies (3547). The lowest number of

genera and subgenera is found in the Afrotropical

region (113), and the Australasian region has the

lowest number of species and subspecies (1377).

Although the current classiﬁcation of the Tipuloidea

is based primarily on typological taxon concepts and

the monophyly of the major ity of higher taxa still has to

be substantiated, a few preliminary conclusions can be

drawn from the information contained in Table 1.

Endemicity and distribution patterns

The greater majority of the species and subspecies of

Tipuloidea are restricted to a single biogeographic

region and thus underscore the endemicity of these

areas, and neighboring biogeographic regions usually

share the presence of only a small percentage of

species and subspecies. The Palaearctic and Nearctic

regions have 88 species and subspecies in common,

the Palaearctic and Oriental regions 176, the Nearctic

and Neotr opical regions 48, and the Oriental and

Australasian regions 44. Other combinations of

biogeographic regions with shared species or subspe-

cies are the Palaearctic and Neotropical regions (3

species-group taxa), the Palaearctic and Afrotropical

Fig. 3 Numbers of species

and subspecies/genera and

subgenera of the Tipuloidea

for the biogeographic

regions. PA—Palaearctic;

NA—Nearctic; NT—

Neotropical; AT—

Afrotropical ; OL—

Oriental; AU—

Australasian; PAC—Paciﬁc

Oceanic Islands; ANT—

Antarctic

Limoniidae:Limnophilinae

Pediciidae

Limoniidae:Dactylolabidinae

Limoniidae:Austrolimnophila

Limoniidae:Epiphragma

Limoniidae:Dicranoptycha

Limoniidae:Helius

Limoniidae:Lipsothrix

Limoniidae:Limnophilomyia

Limoniidae:Limoniinae

Cylindrotomidae

Tipulidae

Fig. 2 Cladogram showing the relationships of the subfamilies

and families of the Tipuloidea, modiﬁed from Oosterbroek &

Theowald (1991). 1: Currently assigned to Limnophilinae;

2: currently assigned to Limoniinae; 3: currently assigned to

Chioneinae. (see Table 1)

462 Hydrobiologia (2008) 595:457–467

123

regions (5), the Palaearctic and Australasian regions

(9), the Nearctic and Afrotropical regions (2), the

Nearctic and Oriental regions (6), the Nearctic and

Australian regions (3), the Neotropical and Afrotrop-

ical regions (3), the Neotropical and Oriental regions

(2), the Neotropical and Aus tralasian regions (3), the

Afrotropical and Orien tal regions (9), and the Afro-

tropical and Aus tralasian regions (5). In most of these

cases, the taxa in common are ubiquitous species and

subspecies such as Atypophthalmus (Atypophthalmus)

umbratus, Symplecta (Trimicra) pilipes pilipes, Tip-

ula (Tipula) oleracea, and Conosia irrorata irrorata,

i.e., taxa of which the range probably or certainly was

accidentally extended by man.

Looking at the distribution of the genera and

subgenera of the Tipuloidea over the six biogeograp hic

regions recognized, a set of unique distribution patterns

can be distinguished (Table 2). Regarding number of

endemic genera and subgenera, the Neotropical and

Australasian regions are the richest with 61 endemic

genus level taxa each. The Afrotropical region contains

35 endemic genera and subgenera, the Palaearctic 29,

and the Oriental and Nearctic regions each 25. Other

unique distribution patterns with a high frequencies of

repetition are the combinations of Holar ctic: Palaearc-

tic—Nearctic (25), Eurasia: Palaearctic—Oriental

(25), Palearctic—Oriental—Nearctic (33) (i.e.,in

addition to the occurrence of the combinations Palae-

arctic—Nearctic and Palaearctic—Oriental),

Cosmopolitan: Palaearctic—Oriental—Nearctic—

Neotropical—Australasian—Afrotropical (22), Orien-

tal—Australasian (16), Palaearctic—Oriental—

Nearctic—Neotropical (13), Neotropical—Austral-

asian (12). All other distribution patterns occur with a

frequency of fewer than 10 times (Table 3).

Palaeogeography

As stated above, the ﬁrst evidence of Diptera in the

fossil record date back to the Lower Triassic (240

Mya), during which the terrestrial domain of the earth

consisted of a single landmass, Pangaea. The oldest

fossil Diptera include species that deﬁnitely represent

ancestral lineages of the present-day Tipuloidea.

Recent distributions of the genera and subgenera of

the Tipuloidea to some extent reﬂect the palaeaog-

eographic history of the earth from Triassic times

onward.

Pangaea existed from the Upper Carboniferous

(320 Mya) to the Upper Jurassic (180–160 Mya),

when it split into the Northern continent Laurasia and

the Southern continent Gondwana.

Since its origin in the Upper Jurassic, Laurasia

fragmented into four main landmasses—correspond-

ing to the present-day western Nearctic, eastern

Nearctic, western Palaearctic, and eastern Palaearc-

tic—that collided in various combinations during

different geological periods (cf. Enghoff, 1995).

Gondwana started to fragment at about 150–130

Mya when Africa drifted away from the remainder of

Gondwana. Since then a sequence of vicariance

events led to a fragmentation history that can be

summarized in the following pare nthetical notation:

(Africa (India (New Zealand (Australia (South

America Antarctica))))).

Pangaea, Laurasia and Gondwana patterns

The Northern and Southern hemispheres share the

presence of 166 genus- group taxa. Widespread genera

such as Baeoura, Rhabdomastix, and Styringomyia

take basic positions in the cladograms by Oosterbroek

& Theowald (1991), which could reﬂect their rela-

tively old age and, with some reservation, their

distribution could be called Pangaean. Worldwide

distributions are also found in relatively derived taxa,

such as Antocha, Molophilus, and Orimarga. More

detailed historical biogeographic studies are neede d to

distinguish between global distributions in the Tipu-

loidea that should be regarded as Pangaean or that

result from histor ical processes that date back to more

recent geological times.

The number of genera and subgenera that exclu-

sively occur in the northern hemisphere (the

Palaearctic and Nearctic regions and the combination

of both), is 79, the number of genera and subgenera

occurring exclusively in the Southern hemisphere

(the Oriental, Neotropical, Afrotropical, and Austra-

lian regions and all combinations of these) is 221.

Both these ﬁgures indicate the relative isolation of

these two composite areas and could be correlated

with the break-up of Pangae a into Lau rasia and

Gondwana.

The Neotropical and Australasian regions share 12

genus-group taxa. The shared occurrence of genera and

subgenera in the Neotropical and Australasian faunas

Hydrobiologia (2008) 595:457–467 463

123

Table 3 Frequency of

distribution patterns of the

genera and subgenera of the

Tipuloidea across

biogeographic region

Higher level classiﬁcation

of the Tipuloidea according

to Oosterbroek (2005). All

data derived from

Oosterbroek (2005)

PA: Palaearctic; NA:

Nearctic; NT: Neotropical;

AT: Afrotropical ; OL:

Oriental; AU: Australasian

x = presence of genus/subgenus

PA NA NT AT OL AU Frequency of pattern Count of regions involved

00x000 61 1

00000x 61 1

000x00 35 1

xx00x0 33 3

x00000 29 1

xx0000 25 2

0x0000 25 1

x000x0 25 2

0000x0 25 1

xxxxxx 22 6

0000xx 16 2

xxx0x0 13 4

00x00x 12 2

x00xxx 9 4

0xx000 7 2

xx00xx 7 4

xxxxx0 6 5

xx0xx0 6 4

x0xxxx 5 5

000xx0 5 2

xx0xxx 4 5

xxxx00 4 4

x00xx0 4 3

xxx0xx 3 5

0xxxxx 2 5

0xx000 2 2

x0x000 2 2

x00x00 2 2

00xxxx 2 4

00xx00 2 2

xx00xx 1 4

xx0x00 1 3

xxx000 1 3

0x00x0 1 2

0xxx0x 1 4

0xx00x 1 3

0xxx00 1 3

x0x0xx 1 4

x0x0x0 1 3

x0xx00 1 3

00x0xx 1 3

00xx0x 1 3

464 Hydrobiologia (2008) 595:457–467

123

of Tipuloidea, as such led Alexander (1929) to suggest

a former Antarctic land connection. A closer look at the

taxa involved reveals more intricate patterns: four

genus-group taxa are present in New Zealand, Austra-

lia, and the Neotropical region, four taxa are

exclusively found in New Zealand and the Neotropical

region, and four taxa are exclusively shared by

Australia and the Neotropical region. Most of the

Neotropical species concerned occur in the south of

that region. In 1960 Hennig published an analysis of

the distribution patterns of Diptera occurring in the

southern hem isphere, including the Tipuloidea. Lack

of information on the phylogenetic relationships of the

taxa involved impeded an actual association of distri-

bution patterns with the geological events that caused

the break-up of Gondwana. Hennig suggested a range

of possible studies on promissing and rewarding

groups of Tipuloidea occurring in the area, but since

then only a few of such studies have been carried out

(cf. De Jong, 1989).

The many endemic genus-group taxa in the

Afrotropical region and the relatively few taxa this

region shares with other biogeographical regions

substantiate the relatively isolated position of the

continent. Only a few patterns indicate an exclusive

afﬁnity of the Afrotropical region with one or more of

the other regions of the Southern hemisphere. Most of

the inter-regional patterns involving the Afrotropical

region suggest a relationship with one or more of the

biogeographical regions from the northern hemi-

sphere. The relatively isolated position of the

Afrotropical region probably reﬂects its early sepa-

ration from the remainder of Gondwa na.

Taxonomic diversiﬁcation patterns at the species-

group level in the Tipuloidea often can be correlated

with Tertiary palaeaogeographic and climatic processes

(e.g., De Jong, 1998; Oosterbroek, 1980; Oosterbroek &

Arntzen, 1992; Tangelder, 1988; Theowald, 1984).

Human related issues

Research

Due to their high species-richness, the Tipuloidea are

an excellent study group for looking at patterns of

local and regional diversity and endemism. Many

species are special ized and have restricted habitat

requirements and can, therefore, be used to indicate

habitat quality and to identify regions of priority for

conservation efforts (Falk, 1991; Oosterbroek, 1994).

Tipuloidea species are often extremely abundant and

they play key roles in many ecosystems, including

aquatic systems, either as consumers or as prey for a

wide variety of predators ranging from ﬁsh, amphi-

bia, birds and mammals, to spiders and predacious

insects.

Tipulid larvae are a component in aquatic bio-

monitoring—as a group they are considered sensitive

to moderately sensitive to human perturbations (e.g.,

rating of 3 on a 1–10 scale in the Hilsenhoff index,

where 10 is highly tolerant of pollution).

Pest species

A few species of Tipuloidea are of economic

importance in that their larvae feed on roots of crops

or on seedlings of ﬁeld crops. By their sheer numbers,

some of these species can become serious pests

(Blackshaw & Coll, 1999). The larva of Tipula

(Yamatotipula) aino is a chronic aquatic pest of rice

seedlings.

References

Alexander, C. P., 1929. Crane-ﬂies. Diptera of Patagonia and

South Chile 1: 1–240, pl. 1–12.

Alexander, C. P., 1931. Deutsche Limnologische Sunda-

Expedition. The crane-ﬂies (Tipulidae, Diptera). Archiv

r Hydrobiologie, Supplement 9, Tropische Bin-

nengewa

sser 2: 135–191, pl. 2.

Alexander, C. P., 1956. Tipulidae. Ruwenzori Expedition

1934–35 1(7): 129–380, pl. 27, 28.

Alexander, C. P., 1964. Diptera (Nematocera): Tanyderidae,

Ptychopteridae, Tipulidae. South African Animal Life 10:

229–441.

Alexander, C. P. & G. W. Byers, 1981. Tipulidae. In McAl-

pine, J. F., B. V. Peterson, G. E. Shewell, H. J. Teskey, J.

R. Vockeroth & D. M. Wood (eds), Manual of Nearctic

Diptera, Vol. 1. Research Branch, Agriculture Canada,

Monograph 27: 153–190.

Blackshaw, R. P. & C. Coll, 1999. Economically important

leatherjackets of grassland and cereals: biology, impact and

control. Integrated Pest Management Reviews 4: 143–160.

Brindle, A., 1960. The larvae and pupae of the British Tipu-

linae (Diptera: Tipulidae). Transactions of the Society of

British Entomology 14: 63–114.

Brindle, A., 1967. The larvae and pupae of the British Cylin-

rotominae and Limoniidae (Diptera, Tipulidae).

Transactions of the Society of British Entomology 17:

151–216.

Hydrobiologia (2008) 595:457–467 465

123

Brinkmann, R., 1997. Diptera Cylindrotomidae. In Nilsson A.

(ed.), Aquatic Insects of North Europe, a Taxonomic

Handbook, Vol. 2. Apollo Books, Stenstrup: 99–104.

Byers, G. W., 1996. Tipulidae. In Merritt R. W. & Cummins K.

W. (eds), An Introduction to the Aquatic Insects of North

America. Kendall/Hunt Publishing Company, Dubuque,

Iowa: 549–570.

de Jong, H., 1998. In search of historical biogeographic pat-

terns in the western Mediterranean terrestrial fauna.

Biological Journal of the Linnean Society 65: 99–164.

de Jong, H., 1989. Revision of the South American taxa El-

noretta Alexander, Euvaldiviana Alexander and

Valdiviana Alexander (Diptera: Tipulidae), with a dis-

cussion on their phylogeny. Systematic Entomology 14:

243–257.

Dienske, J. W., 1987. An illustrated key to the genera and

subgenera of the western Palaearctic Limoniidae (Insecta:

Diptera), including a description of the external mor-

phology. Stuttgarter Beitra

ge zur Naturkunde, Serie A

(Biologie) 409: 1–52.

Dobrotwosky, N. V. 1968–1974. The Tipulidae (Diptera) of

Australia. Australian Journal of Zoology 16: 459–494;

495–510; Supplement 1: 1–41; Supplement 5: 21–45;

Supplement 16: 1–9; 11–43; Supplement 25: 1–14; 15–30;

31–62; 63–75.

Enghoff, H., 1995. Historical biogeography of the Holarctic:

area relationships, ancestral areas, and dispersal of non-

marine animals. Cladistics 11: 223–263.

Evenhuis, N. L., 1994. Catalogue of the Fossil Flies of the

World (Insecta: Diptera). Backhuys Publishers, Leiden:

1–600.

Falk, S., 1991. A review of the scarce and threatened ﬂies of

Great Britain (Part 1). Research and Survey in Nature

Conservation 39: 1–192.

Gelhaus, J. K., 1986. Larvae of the crane ﬂy genus Tipula in

North America (Diptera: Tipulidae). The University of

Kansas Science Bulletin 53: 121–182.

Gelhaus, J. K. 2000. Manual for the identiﬁcation of aquatic

crane ﬂy larvae for Northwest North America: 1–147. The

Northwest Biological Assessment Workgroup’s Taxo-

nomic Workshop on Diptera, Ellensburg, Washington.

Gelhaus, J. K., 2002. Manual for the identiﬁcation of aquatic

crane ﬂy larvae for Southeastern Uniited States: 1–205.

Carolina Area Benthological Workshop, Durham, North

Carolina.

Gelhaus, J. K. & G. W. Byers, 1994. Tipulidae. In Morse, J. C.,

L.Yang & L. Tian (eds), Aquatic Insects of China Useful

for Monitoring Water Quality: 438–458.

Hennig, W., 1960. Die Dipteren-fauna von Neuseeland als

systematisches und tiergeograﬁsches Problem. Beitra

zur Entomologie 10: 221–329.

Hofsvang, T., 1997. Diptera Tipulidae, crane ﬂies. In Nilsson

A. (ed.), Aquatic Insects of North Europe, a Taxonomic

Handbook, Vol. 2. Apollo Books, Stenstrup: 93–98.

Krzeminski, W. & E. Krzeminska, 2003. Triassic Diptera:

descriptions, revisions and phylogenetic relations. Acta

Zoologica Cracoviensia 46: 153–184 Supplement.

Mannheims, B. 1951–1968. Tipulidae. Fliegen der Palaeark-

tischen Region 3(5) 1: 1–320.

Michelsen, V., 1996. Neodiptera: new insights into the adult

morphology and higher level phylogeny of Diptera

(Insecta). Zoological Journal of the Linnean Society 117:

71–102.

Oosterbroek, P., 1980. The western palaearctic species of

Nephrotoma Meigen, 1803 (Diptera, Tipulidae), Part 5,

phylogeny and biogeography. Beaufortia 29: 311–393.

Oosterbroek, P., 1994. Biodiversity in the Mediterranean

region. In Forey, P. L., C. J. Humphries & R. I. Vane-

Wright (eds), Systematics Conservation Evaluation. Sys-

tematics Association Special, Vol. 50. Clarendon Press,

Oxford: 289–307.

Oosterbroek, P., 1998. The families of Diptera of the Malay

Archipelago. Fauna Malesiana Handbooks 1: i-xii,

1–227.

Oosterbroek, P. 2005. Catalogue of the Craneﬂies of the World,

version 17.05.2005. The catalogue is on-line available via

http://www.science.uva.nl/zma/.

Oosterbroek, P. & J. W. Arntzen, 1992. Area-cladograms of

Circum-Mediterranean taxa in relation to Mediterranean

palaeogeography. Journal of Biogeography 19: 3–20.

Oosterbroek, P. & G. Courtney, 1995. Phylogeny of the ne-

matocerous families of Diptera (Insecta). Zoological

Journal of the Linnean Society 115: 267–311.

Oosterbroek, P. & B. Theowald, 1991. Phylogeny of the Tipu-

loidea based on characters of larvae and pupae (Diptera,

Nematocera), with an index to the literature except Tipu-

lidae. Tijdschrift voor Entomologie 134: 211–267.

Peus, F., 1952. Cylindrotomidae. Fliegen der Palaearktischen

Region 3(5) 3: 1–80.

Pritchard, G., 1983. Biology of Tipulidae. Annual Review of

Entomology 28: 1–22.

Reusch, H. & P. Oosterbroek, 1997. Diptera Limoniidae and

Pediciidae, short-palped crane ﬂies. In Nilsson A. (ed.),

Aquatic Insects of North Europe, a Taxonomic Handbook,

Vol. 2. Apollo Books, Stenstrup: 105–132.

Savchenko, E. N., 1961–1983. Crane ﬂies (Diptera, Tipulidae.

Fauna SSSR, Diptera 2 (N.S.) 79: 1–488; 89: 1–503; 105:

1–282; 127: 1–585.

Savchenko, E. N., 1982–1986. Limoniid ﬂies. Fauna Ukrainy

14 (2): 1–340; 14 (3): 1–335; 14 (4): 1–180.

Savchenko, E. N., 1983. Limoniidae of South Primorye. Aka-

demia Nauk Ukrainian SSR. Kiev: 1–156.

Savchenko, E. N., 1989. Limoniidae fauna of the USSR. Ak-

ademia Nauk Ukrainian SSR, Kiev: 1–377.

Savchenko, E. N. & G. O. Krivolutskaya, 1976. Limoniidae of

the South Kuril Islands and South Sakhalin. Akademia

Nauk Ukrainian SSR. Kiev: 1–160.

Shcherbakov,

E., E. D. Lukashevich, & V. A. Blagoderov,

1995 Triassic Diptera and initial radiation of the order.

International Journal of Dipterological Research 6: 75–115.

Stary, J., 1992. Phylogeny and classiﬁcation of Tipulomorpha,

with special emphasis on the family Limoniidae. Acta

Zoologica Cracoviensia 35: 11–36.

Tangelder, I. R. M., 1988. The biogeography of the Holarctic

Nephrotoma dorsalis species-group (Diptera, Tipulidae).

Beaufortia 38: 1–35.

Theischinger, G., 1996. The Limoniinae (Diptera: Tipulidae) of

Australia. VIII. Preliminary key to the genus-group taxa.

Stapﬁa 44: 117–144.

Theowald, B., 1957. Die Entwicklungsstadien der Tipuliden

(Diptera, Nematocera) insbesondere der west-palaearktis-

chen Arten. Tijdschrift voor Entomologie 100: 195–308.

466 Hydrobiologia (2008) 595:457–467

123