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The Natural History of Madagascar

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The Natural History of Madagascar The Natural History of Madagascar Edited by Steven M. Goodman and Jonathan P. Benstead Photographs by Harald Schutz FielH THE SCHL[NGER PSKSI ILMJ T Museum FOUNDATION \mm3 *** The University of Chicago Press Chicago and London Steven M, Guodniau i\ Held Biologist ni the Held Museum of Natural History in Chicago, coordinator of the Ecology Training Program of WWF- Madagasca r, and Lecturer at rhe Universke d1 Antananarivo* He- resides in Madagascar and over the pas: 15 years has conducted biological inventories of many unknown or poorly known areas of die island. His main study groups are birds and mammals and he has published widely on the biodiversity of the island. Jonathan R Ben stead is a freshwater ecologist and postdoctoral researcher at The Kcosystems Center of the Marine Biological Laboratory in Woods Hole. He conducted his doctoral research on the ecology of the web-tooted tenrec and the consequences of deforestation for stream ecosystems in eastern Madagascar. He has broad interests in freshwater biodiversity* stream ecology, and semi-aquatic mammals. The University of Chicago Press wishes to gratefully acknowledge The Field Museum of Natural History, QIT Madagascar Minerals, the Sch linger Foundation, and the WWF- Internationa I for their generous contributions toward publication of The Natural History of Madagascar. The University of Chicago Press* Chicago 606.17 The University of Chicago Press* Ltd., London G 20(13 by The University of Chicago All rights reserved Published 2003 Printed in China 12 II 10 09 OK 07 06 05 04 03 12 3 4 5 ISBN 0-226-30106-3 (cloth) Library of Congress Cataloging in Publication Data The natural history of Madagascar / edited by Steven M. Goodman and Jonathan P. Bens re ad \ photographs by Harald Schut£. Includes bibliographical references (p. ). ISBN 0-226-30306-3 (doth : alk. paper! L Natural history — Madagascar. I. Goodman, Steven .VL JL Ben stead, Jonathan P. QH195.M2N38 2003 508.691 — dc2! 2002045581 This hook is printed on acid-free paper. 692 Invertebrates Systematic Accounts ■ True Flies Diptera, True Flies M. E. Irwin, E, I. Schlinger, and F. C. Thompson The class Insecta is extremely rich in species, containing ap- an estimated 60,000 species, is by far the most species-rid proximately 75% of all known life forms. Reasonable es- of the orders, followed by Hymenoptera (36,000 specie^ timates put the total number of insect species somewhere and Coleoptera (26,000 species) (Kosztarab and Schatfl between 5 million and 1 2 million, with about I million spe- 1990). Even given these figures, it is generally agreed [hjjjj cies currently described (May 1990; Gaston 1991; Ode- in tropical environments, beetles arc more specioscthand* gaard 2000). Within Insecta, Diptera, or true flies, is one of other three large holometabolous orders (F.rwm 1'SJ, the four holometabolous (higher insects having separ- 1 997), In North America and generally speaking through ate and distinct egg, larval, pupal, and adult stages) orders out the world, the discovery phase, that is, the part of| with vast numbers of species; the other three orders are timeline during which most of the species in a given en?jj Coleoptera (beetles), Hymenoptera (ants, bees, and wasps), ronment are discovered and described, is quite advance^ and Lepidoprera (moths and butterflies). These four orders for such groups as the beetles, butterflies, bees, andaa% collectively make up the majority of insect species and thus whereas it is still in its early stages for moths and wasps ml comprise an extremely large proportion of the earth's biota. is just beginning for flies. Of these notable insect or&n Of these four orders, Diptera is by far the least known. For Diptera is by far the least known and least understood. example, fewer than 30% of the species in North Amer- ica have been described (Kosztarab and Schacfer 1990; Thompson 1990). It is noteworthy that in North America, Characteristics of Diptera where collecting efforts have been relatively intense and ex- tensive, a greater proportion of the species of other large Diptera is an order of true holometabolous insects. Ch» insect orders are described (90% for Coleoptera, 50% for acteristically, flies have a single pair of wings. The order! Hymenoptera, 80% for Lepidoptera). That trend likely named for this feature: di = two; ptera — wing. Onlyafet holds for other regions of the world, including Madagascar, other orders of insects are characterized by having n» although the relative proportions themselves probably vary wings, of which Strepsiptera is the most notable. Howaq considerably. Within North America, Diptera, containing the wings of Strepsiptera arise on the metathorax, wheroi M. E. Irwin, E. I. Schlinger, and F. C. Thompson 693 tose of Diptera am on the mesorhorax. In addition, some ficial. Some groups of flies, for instance, are parasitoids of Bale scale insects (Coccoidea) have only two wings and agricultural pests. As an example, larvae of tachinid flies are llBBulohalter on the metathorax. A few dipterous taxa extremely important in lowering populations of pestiferous, lave lost all or part or their single pair of wings. The leaf-feeding moth larvae, and syrphid fly larvae are impor- BBapomorphy (derived character) that defines the flies and tant predators of injurious, sap-sucking aphids. One usu- Bparates them from all other insects is the halters, which ally thinks of the bees as key pollinators of flowering plants, He formed from the extremely reduced hind wings. These including crops, but flies are also important pollinators. A Bructures are miniature gyroscopes, acting to stabilize the large number of fly taxa, including those found in Mada- ^kW) during flight. Adult flies, like adults of a few other gascar, are fungi vores or phytophagous. Moreover, some Orders of insects, have piercing or lapping mouthparts. groups of flies play an economically as well as ecologically Most other adults of holometabolous insect orders have critical role by breaking down dead vegetation and animal iewing (Coleoptera and I lymenoptera) or lapping (Lepi- carcasses; they serve the vital function of recycling organic 4pera and Hymenoptera) mouthparts. products into mulch. Environmental and Ecological Diversity Taxon Richness Itcause holometabolous insects usually occupy different Approximately 150,000 currently valid species of flies have habitats during their larval (feeding) and adult (rcproduc- been described worldwide (table 8.35). These have been fft)stages, they inhabit all, and dominate most, freshwa- placed in a few more than 11,000 valid genera. As of early |tf and terrestrial ecosystems globally, including Madagas- 2001, 19,051 valid fly species have been described in 2,230 Bt-Beyond simple numbers, the Diptera haw an extremely genera in rhe Afrorropical region, representing about 13% ttrsearray of life history traits, encompassing detriti votes of the world's fly species and 24% of the world's fly genera. Saprophytes), herbivores (fungi vores and phytopbages), In Madagascar, which we consider to be a portion of the pedators (carnivores), parasitoids (larva parasitizes and A f rot topical region, 1,796 species in 538 genera have been kiik the host before becoming adult), and parasites. Niche described. Although Madagascar contains a mere 9% of ftersity is considerably less within other speciose holo- the described Afrotropical species (1 % of the world spe- Ifctabolous orders. As a group, flies seemingly perform a cies), it holds 24% of the Afrotropical genera (6% of the wider and more diverse array of ecological functions than world genera). On average, there are more than 16 species (■(Other order in rhe animal kingdom. of flies per genus on a world basis, fewer than 9 species per genus in the Afrotropical region, and slightly more than 3 species per genus in Madagascar. Endemism is relatively high in Madagascar. Of the described taxa, fully 14% of Importance the genera and nearly 80% of the species of flics that occur in Madagascar are endemic to that island. : of their diverse ecological roles, flies have an enor- On an area basis, the Afrotropical region, with approx- flpos impact, not only on the environment but on human- imately 20 million lent, has about 0.001 species and about tgOSwell. Some of these effects can be detrimental, but 0.0001 genus of flies per knr; Madagascar, with 587,041 ■Vft are advantageous. From a human health perspective, km", has about 0.003 species and 0.0009 genus per km-. ttmral fly families are vectors of deleterious and even lethal Therefore, on a per unit area basis, Madagascar is about fere, such as malaria, which is transmitted by anophe- three times richer in described species and nine times richer tattosquitoes (see Duchemin er al„ "Culicidae, Mosqui- in described genera than the Afrotropical region as a whole. #3," this volume). Other flies arc consequential pests of apiculture. The true fruit flies (Tephritidae) are among the ■stserious pests of agricultural crops. The Mediterranean lit fly, Qemtitii capitata (Wiedemarm), causes billions of Classification and Diversity 4&rs of damage to fruits worldwide. A number of fami- hof flies nre external or internal parasites of vertebrate The order Diptera is conventionally divided into two sub- #*,induding humans. They can seriously debilitate live- orders, Nematoccra and Brachyccra (McAlpine ct al. 1981). tod and pets. The screw worm, CochUomyta hominivorax Nematocera, considered the more primitive, is character- Ifcquerell (family Calliphoridac), is an example. Many ized by having
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