Hydrobiologia (2008) 595:457–467 DOI 10.1007/s10750-007-9131-0
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
Ó 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
Guest editors: E. V. Balian, C. Le´veˆque, 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 supplementary 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:
[email protected] 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
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 Afrotropical 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 distribution 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
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(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 phylogenetic position within the lower Diptera has not yet been established satisfactorily. Traditionally, the Tipuloidea were regarded as representing one of the most primitive 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 more 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 monophyly 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 Tipulidae 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 hemicephalous, 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,
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Fig. 1 Tipula (Acutipula) maxima Poda, 1761, male. European species with larva in marshy soil or freely aquatic in small rivulets, adult in wet woodland along streams. Germany, photo Ju¨rgen Peters
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 welldeveloped, 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., Dicranomyia (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; prothorax and metathorax small were reduced to strip-like elements. Legs long and slender, tips of tibiae 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,
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especially in species living in isolated localities like islands and mountain ranges, or in cold-adapted 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 adaptations of the last tergites, sternites and their genital processes. Structures of male terminalia considered speciesspecific. Female terminalia generally forming an ovipositor consisting of sclerotized, bladelike cerci and hypogynial valves, usually less characteristic for individual species, but showing differences 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 distinct, retracted into anterior thoracic segments, deeply incised ventrally and often dorso-laterally more reduced. Abdominal segments smooth, or with transverse rows of hairs. Creeping welts or fleshy projections sometimes present. Spiracular disc usually 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 identification of adult Tipuloidea, although they are rather
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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 environments 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.
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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.
Larvae of crane flies occupy a wide variety of habitats, ranging from strictly aquatic to entirely terrestrial. Fully aquatic larvae can occur in freshwater, especially in rapidly flowing streams, or in brackish water or intertidal zones. Some of these species live entirely submerged, are without functional 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 number of 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 waterfilled 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
Species diversity Numbers of species and subspecies An overview of the genera, subgenera 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 Tipulidae sensu stricto
17 45
13 38
3 36
0 23
17 45
1 30
26 115
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
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Catalogue of the Craneflies of the World. The catalogue is available on-line via http://www.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 considerable. 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: Aymaramyia, Gonomyopsis, Jivaromyia, Quathlambia, etc., Pediciidae: Savchenkoiana, Tipulidae s.str.: Austrotipula, Elnoretta, Euvaldiviana etc. Other genera and subgenera hold hundreds of species, e.g., Limoniidae: Cheilotrichia, Erioptera, Gonomyia, Idiocera, Molophilus, Ormosia, Teucholabis, Austrolimnophila, Gynoplistia, Hexatoma, Dicranomyia, Geranomyia, Limonia, Pediciidae: Dicranota, Tipulidae s.str.: Dolichopeza, Nephrotoma, Tipula. These divergent numbers could be related to disparities in the ages of the higher 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 reexamination of taxon clusters containing cryptic species using molecular techniques is expected to increase the number of recognized species considerably.
which is dated at 240 Mya (Krzeminski & Krzeminska, 2003). Other Triassic finds include fossils from the Ladinian-Carnian Madygen Formation in Kyrgyzstan, dated at 230–225 Mya and the NorianRhaetian Tologoi Formation in Kazakhstan, 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. Evenhuis (1994).
Phylogeny and historical processes
Present distribution and main areas of endemism
Fossil record
Distribution
The oldest fossils of Tipuloidea, and of Diptera as such, date back to the Lower Triassic Longtanian of the Gre´s a` Voltzia Formation of Arzviller in France,
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.
Phylogeny Published hypotheses about the phylogeny of the Tipuloidea include Stary (1992) and Oosterbroek & Theowald (1991). Stary (1992) studied the relationships 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 monophyletic 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 Oosterbroek & 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.
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Limoniidae:Limnophilinae Pediciidae Limoniidae:Dactylolabidinae Limoniidae:Austrolimnophila 1 Limoniidae:Epiphragma 1 Limoniidae:Dicranoptycha 2 Limoniidae:Helius 2 Limoniidae:Lipsothrix 2 Limoniidae:Limnophilomyia 3 Limoniidae:Limoniinae
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 majority of higher taxa still has to be substantiated, a few preliminary conclusions can be drawn from the information contained in Table 1.
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)
According to the present classification, these numbers are the following: Palaearctic region, 205 genera and subgenera, 3088 species and subspecies; Oriental region, 190 and 3412; 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 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
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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 Neotropical regions 48, and the Oriental and Australasian regions 44. Other combinations of biogeographic regions with shared species or subspecies are the Palaearctic and Neotropical regions (3 species-group taxa), the Palaearctic and Afrotropical
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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 Afrotropical regions (3), the Neotropical and Oriental regions (2), the Neotropical and Australasian regions (3), the Afrotropical and Oriental regions (9), and the Afrotropical and Australasian 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, Tipula (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 biogeographic 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 Holarctic: Palaearctic—Nearctic (25), Eurasia: Palaearctic—Oriental (25), Palearctic—Oriental—Nearctic (33) (i.e., in addition to the occurrence of the combinations Palaearctic—Nearctic and Palaearctic—Oriental), Cosmopolitan: Palaearctic—Oriental—Nearctic— Neotropical—Australasian—Afrotropical (22), Oriental—Australasian (16), Palaearctic—Oriental— Nearctic—Neotropical (13), Neotropical—Australasian (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 palaeaogeographic history of the earth from Triassic times onward.
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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—corresponding to the present-day western Nearctic, eastern Nearctic, western Palaearctic, and eastern Palaearctic—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 parenthetical 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 relatively 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 needed to distinguish between global distributions in the Tipuloidea that should be regarded as Pangaean or that result from historical processes that date back to more recent geological times. The number of genera and subgenera that exclusively 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 Australian 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 Pangaea into Laurasia 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
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464 Table 3 Frequency of distribution patterns of the genera and subgenera of the Tipuloidea across biogeographic region
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x = presence of genus/subgenus PA
NA
NT
AT
OL
AU
Frequency of pattern
Count of regions involved
0
0
x
0
0
0
61
1
0
0
0
0
0
x
61
1
0 x
0 x
0 0
x 0
0 x
0 0
35 33
1 3
x
0
0
0
0
0
29
1
x
x
0
0
0
0
25
2
0
x
0
0
0
0
25
1
x
0
0
0
x
0
25
2
0
0
0
0
x
0
25
1
x
x
x
x
x
x
22
6
0
0
0
0
x
x
16
2
x
x
x
0
x
0
13
4
0
0
x
0
0
x
12
2
x
0
0
x
x
x
9
4
0
x
x
0
0
0
7
2
x
x
0
0
x
x
7
4
x
x
x
x
x
0
6
5
x
x
0
x
x
0
6
4
x 0
0 0
x 0
x x
x x
x 0
5 5
5 2
x
x
0
x
x
x
4
5
x
x
x
x
0
0
4
4
x
0
0
x
x
0
4
3
x
x
x
0
x
x
3
5
0
x
x
x
x
x
2
5
0
x
x
0
0
0
2
2
x
0
x
0
0
0
2
2
x
0
0
x
0
0
2
2
0
0
x
x
x
x
2
4
0
0
x
x
0
0
2
2
x
x
0
0
x
x
1
4
x
x
0
x
0
0
1
3
x
x
x
0
0
0
1
3
0 0
x x
0 x
0 x
x 0
0 x
1 1
2 4
Higher level classification of the Tipuloidea according to Oosterbroek (2005). All data derived from Oosterbroek (2005)
0
x
x
0
0
x
1
3
0
x
x
x
0
0
1
3
x
0
x
0
x
x
1
4
x
0
x
0
x
0
1
3
PA: Palaearctic; NA: Nearctic; NT: Neotropical; AT: Afrotropical ; OL: Oriental; AU: Australasian
x
0
x
x
0
0
1
3
0
0
x
0
x
x
1
3
0
0
x
x
0
x
1
3
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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, Australia, 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 hemisphere, including the Tipuloidea. Lack of information on the phylogenetic relationships of the taxa involved impeded an actual association of distribution 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 hemisphere. The relatively isolated position of the Afrotropical region probably reflects its early separation from the remainder of Gondwana. Taxonomic diversification patterns at the speciesgroup 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 specialized and have restricted habitat requirements and can, therefore, be used to indicate
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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, amphibia, birds and mammals, to spiders and predacious insects. Tipulid larvae are a component in aquatic biomonitoring—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.
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