B I O D I V E R S I T Y 3 ( 4 ) 1

2002

Policy of RespectTo engender and foster respect for all living species,this journal capitalizes the common names of allformally named species.

BiodiversityVolume 3 Number 4 November 2002

ARTICLESExploring the diversity of flies (Diptera)Jeffrey H. Skevington and P.T. Dang (Editors) . . . . . . . . . . . 3

Introducing the ubiquitous Diptera by J.R. Vockeroth (3)Audacious predacious lifestyles by Scott E. Brooks (6)Intimate neighbours: Parasitoids and parasites by Jeffrey H.

Skevington (8)Cleaning up the world: Dipteran decomposers by Jade Savage (12)Going vegetarian: Plant and fungus feeding by Stéphanie Boucher

and Terry A. Wheeler (15)Flowers, pollination, and the associated diversity of flies by Peter

G. Kevan (16)Flies as vectors of disease by Desmond H. Foley (18)When being a maggot is a good thing: The role of Diptera in forensic

science by Andrew McDowell (20)Molecular systematics of flies (Diptera) by Shaun L. Winterton (21)Dipteran glow-worms: Marvellous maggots weave magic for tourists

by Claire H. Baker (23)

FORUMAn opportunity for innovation in managing fisheries: Internationalfishery closures and marine protected areas on the Grand Banks . . . 38Jon Lien

IN EVERY ISSUEEDITOR’S CORNERFunding priorities for species and ecosystems research . . . . . . . . . . . i.

LETTERS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

SPECIES BY SPECIESGiant Chacoan Peccary (Catagonus wagneri) . . . . . . . . . . . . . . . . . 28William Toone and Michael WallaceBIODIVERSITY NEWSStories include: Global estimate of insect diversity now reduced; GreatBear Rainforest (British Columbia); Sei Whales included in researchwhaling (Japan); Cloning pandas (China); backgrounder on “biodiversityrights legislation” . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32

BOOK REVIEWSConservation Of Exploited Species (Reynolds et al, eds); MountainGorillas (Robbins et al, eds); Parasitism (Bush et al); FreshwaterAlgal Flora, British Isles (D.M. John et al) . . . . . . . . . . . . . . . . . . . . . 38ANNOUNCEMENTS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44

EditorTED MOSQUIN, PhD Botany, Canada

Honorary EditorMAURICE STRONGUnder-Secretary-General, UNSpecial Advisor to the UN Secretary-GeneralSecretary-General, UN Conference on

Environment & Development,Biological Diversity Convention, Brazil, 1992.

Associate EditorsP. BALAKRISHNA, PhD Molecular Biology, Sri LankaWILLIAM C.G. BURNS, International Wildlife Law, U.SAPAUL CATLING, PhD Botany, CanadaJOHN LAMBERT, PhD Medicinal Plants, World BankBERNARD LANDRY, PhDEntomology, SwitzerlandGEOFF SCUDDER, D. Phil. Zoology, CanadaMICHAEL SHARKEY,PhD Entomology, U SAVO TONG XUAN, PhD Agronomy, Viet NamIAN SMITH, PhD Arachnids, Freshwater Arthropods, CanadaMANUEL ZUMBADO, Curator - InBIO, Costa RicaManaging EditorCATHERINE RIPLEY

Assistant EditorRICHARD VOCKEROTH, D. Phil.

Book Review EditorK.G. ANDREW HAMILTON, PhD

News EditorSTEPHEN AITKENResearch & Development DirectorROBERT McFETRIDGE

IllustratorROELOF IDEMALayoutH.T. MAIEditorial SubmissionsManaging Editorc/o Tropical Conservancy (see address below)m-editor@tc-biodiversity.orgSubscription & Charitable DonationsV. ChungTropical Conservancy94 Four Seasons DriveOttawa, Ontario, Canada K2E 7S1Tel: 1-613-224-9518admin@tc-biodiversity.org; URL: http://tc-biodiversity.org

ISSN 1488-8386

Publication Date: 20 September 2002

This issue is supported in part by the InternationalDevelopment Research Centre (Canada).

INDEX /VOLUME 3 (2002 ) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46

○ ○ ○ ○ ○ ○ ○

The images of flies on the front cover illustrate some of the remarkable variation among the 125,000described species belonging to the order Diptera in the class Insecta. Flies show countless differencesnot only in appearance but also in genetics, physiology, behaviour, habits, ecological function, anddistribution. The three illustrated species include: 1. [center] Lasia sp. (Acroceridae – small-headedflies) – Species of Lasia occur only in the New World. As with all small-headed flies, their larvae areparasitoids of spiders; 2. [left] Giraffomyia willeyi (Platystomatidae – signal flies) – The genus, withfive species, occurs only in New Britain and the Solomon Islands. The outgrowths from the headoccur only in the males and are presumably used in mating rituals. Unlike eyestalks, found in eightfamilies of flies (see back cover), they are not rigid processes but have a weak flexible base. Larvaeare unknown; and 3. [right] Trichopoda pennipes (Tachinidae – tachinid flies). – A native of temperateNorth America, this species has been introduced into many parts of the world as a biocontrol agentagainst pest species of stink bugs. The family Tachinidae has 9,200 species worldwide. All areparasitoids of arthropods. – Illustrated by Roelof Idema. More fly photos on page 44 & back cover.

B I O D I V E R S I T Y 3 ( 4 ) 3

T R O P I C A L C O N S E R V A N C YA R T I C L E SA R T I C L E SA R T I C L E SA R T I C L E SA R T I C L E S

Coordinated and edited byJeffrey H. Skevington and P.T. DangSystematic EntomologyAgriculture and Agri-Food Canada (AAFC),Ottawa, ON K1A 0C6Canadajskevington@cdfa.ca.gov or jeff_skevington@yahoo.cadangp@agr.gc.ca

ABSTRACT. Flies (Diptera) are an important but under-appreciated part of our planet’s biodiversity. With over124,000 described species, and countless more awaitingdiscovery, they are one of the most diverse groups oforganisms on Earth. This series of ten papers explores thediversity of Diptera. Several authors describe the diversityof dipteran lifestyles and behaviours, both as larvae andadults. They also reveal the various roles that these animalsplay in the ecological interactions of the planet—countlessnumbers of flies feed on plants, control pest arthropods(including other flies!), break down rotting vegetation andexcrement, pollinate flowers, provide food for other species,and of course, spread diseases. Indeed, because of theirrole as vectors of disease, flies have almost single-handedlyprevented the economic development of countries in tropicalAfrica and South America. But flies are used in positive waysby humans, too, and several authors describe their use inforensic science, molecular research, and even as “mainattractions” in the tourism industry. The intent of this seriesof papers is to encourage a broader interest in Diptera that,ideally, will lead to further research and conservation efforts.

INTRODUCING THEUBIQUITOUS DIPTERA

J.R.VockerothFlies (Diptera) are among the most ubiquitous and widelydistributed insects. Their close association with humans hasled them to be perceived as annoying and unpleasant crea-tures, and indeed some flies are the cause of millions ofdeaths and illnesses among human populations. Yet flies arealso among the key components in most ecosystems andare beneficial in many ways.WHAT ARE FLIES?Flies are insects with only one pair of functional wings;the hind wings are reduced to a pair of stalked knobs, calledhalters, which are used for balance during flight. The adultmouthparts are modified for sucking liquids; mandibles,used in mostinsects for

chewing, are usually absent. When present (e.g.in mosquitoes), mandibles are in the form of slen-der stylets that can pierce the skin of larger ani-mals. In some families the mouthparts may be non-functional or even absent. The eyesare often large and in extremecases, such as big-headed flies(Pipunculidae), they cover a lmos tthe entire surface of thehead. In other groups,such as stalk-eyedflies (Diopsidae), the eyes are located at theends of long stalks. In a few families thewings and halters are often reduced orabsent. Some bat parasites of the familyStreblidae show one of the greatest mor-phological reductions found in insects –the mature female consists of an egg-producing sac embedded under theskin of the bat wing.Adults range in length fromabout 0.5 mm (some biting midges, Ceratopogonidae) toabout 60 mm with a wingspan up to 75 mm (some Austra-lian robber flies, Asilidae). There are three main groups offlies, each with a different and characteristic body shapeand antennal structure:(1) The Nematocera (Figure 1), the oldest group, usu-

ally have a slender body and long slender antennaewith 16 apparent segments; in some males these seg-ments have long dense hairs; common groups arecrane flies (Tipulidae), mosquitoes (Culicidae),midges (Chironomidae), and blackflies (Simuliidae);

(2) The lower Brachycera, including horse and deer flies(Tabanidae), are more robust and have shorter andstouter antennae with three to ten apparent segments;

(3) The higher Brachycera, including muscid flies (Muscidae)and blow flies (Calliphoridae), typically have a short broadbody and antennae with three broad segments.

Larvae, commonly known as maggots, (at least whenreferring to the higher flies), are extremely varied in shapeand structure. Their most conspicuous common feature isa lack of segmented legs (Figure 2). Primitive familiesusually have both the head capsule and the mouthparts welldeveloped. Aquatic larvae of several families have themouthparts modified into a pair of dense brushes of hairsor slender rods for filtering microorganisms from the water(Figure 2). In more specialized flies, the head capsule be-comes more and more reduced and retracted into the bodyuntil only the apices of the mandibles are protrusible andare used for rasping (Figure 3).Larvae move using a variety of surface structures, usuallyprolegs (fleshy, leg-like processes) or creeping welts (cov-

Exploring the diversity of flies (Diptera)

Figure 1.Phantom crane fliessuch asBittacomorphaclavipes(Ptychopteridae) aremembers of theNematocera.Immature phantomcrane flies are aquaticand the adults can beseen along smallstreams in all bioticregions exceptAustralasia. B.clavipes is unusual inthat the firsttarsomere of each legis dilated and filledwith air, enablingindividuals to drift inthe wind. (Illustrationcourtesy of AAFC[from McAlpine et al1981, p. 325]).

Figure 2.A typical Dipteranlarva, Notiphila sp.(Ephydridae), lackssegmented legs andpossesses creepingwelts for moving.This aquatic specieshas a flexible airtube that allows it totake in surface airfor respiration.(Illustration courtesyof AAFC [McAlpineet al 1981, p.1044]).

T R O P I C A L C O N S E R V A N C Y4

ered with many small spines). Many aquatic larvae, such asmosquito larvae, swim vigorously with movements of

the whole body. All larval net-winged midges(Blephariceridae) and a few moth flies (Psy-chodidae) live in flowing water and have oneto six ventral sucking discs used for locomo-

tion and anchoring. Respiration is usually bymeans of paired spiracles (breathing pores). In

most aquatic larvae no functional spiracles arepresent. A few of these have processes that act asgills; mosquito larvae have a rigid apical tube that

takes in surface air and many other aquatic larvaehave a telescopic tube with the same function.

DIVERSITY AND ABUNDANCEFlies are among the most diverse group of organisms with

128 families and about 124,000 described species(Brown 2001). The crane fly family (Tipulidae)is the largest with about 14,000 species of which

an American entomologist, C.P. Alexander,described 12,000 between 1911 and 1963. Other

very large families of flies include the midges(Chironomidae) with 5,000 species, the bitingmidges (Ceratopogonidae) with 5,300 species, the

gall midges (Cecidomyiidae) with 4,600 species, therobber flies (Asilidae) with 5,600 species, the beeflies (Bombyliidae) with 4,800 species, the long-

legged flies (Dolichopodidae) with 5,100 species, the hoverflies (Syrphidae) with 5,800 species, and parasitic flies(Tachinidae) with 9,200 species.Unlike the families of birds and mammals, most familiesof Diptera are nearly worldwide in distribution. Sub-Sa-haran Africa, which is a major region with no direct pastconnections with the Nearctic, has 95 families; 8 of these,with about 156 species, are not native to North America.Hölldobler and Wilson (1994) concluded that the ants(Formicidae) are far more numerous as individuals thaninsects of any other family. I believe that midges(Chironomidae) are probably just as abundant. Enormousmating swarms of midges form in cool temperate andarctic regions throughout most of the summer. On calmdays on the arctic tundra of Canada swarm after swarmof many midge species are nearly continuous and far out-number all other insects.SURVIVAL STRATEGIESReproduction. The life cycle of Diptera includes four stages- egg, larva, pupa, adult - and the length of this cycle variesfrom 30 days or less in Drosophila to several years in higharctic species. All flesh flies (Sarcophagidae) give birth toliving larvae rather than laying eggs. In tsetse flies(Glossinidae) and three families of external parasitic flies re-stricted to birds and mammals (collectively known asPupipara), one larva at a time is nourished by glands in theuterus of the female fly. When mature, the larva is ejectedand pupates outside the female body; this type of larval de-velopment does not occur in other insects. Paedogenesis (re-production by immature stages) is found in several gall midges(Cecidomyiidae); here the larvae produce more young.

A strategy apparently unique among insects is found inone species of dark-winged fungus gnat (Sciaridae)(Steffan 1973, 1975). Two to five larvae form a commoncocoon; both sexes are always present and the immobileadults mate and oviposit in the cocoon. It seems almostcertain that larvae of one or both sexes can recognize lar-vae of the opposite sex.Mating. Most orders of insects have a single or at leasta predominant mating position. Moths and butterflies(Lepidoptera) typically mate tail to tail, with their bodiesupright and facing in opposite directions; beetles (Co-leoptera) face in the same direction with the male abovethe female. In flies the male reproductive structures oc-cupy various positions that are either fixed on emergencefrom the pupa or can be altered at the time of mating. Inmost cases the initial coupling position differs from thefinal mating position, but in each case the dorsal surfaceof the male intromittent organ is opposed to the ventralsurface of the vagina. The five major initial and final po-sitions were illustrated and described by McAlpine (1981).A common mating behaviour among the lower Diptera(Chironomidae and others) is the formation of dense andsometimes enormous swarms. The swarms are generallycomposed of males, and when females enter the swarm,coupling quickly takes place. Picking out a mate is accom-plished by one of several adaptations. For example, the eyesof males of most of these species are enlarged and contigu-ous and the males of many species have numerous long,fine hairs on their antennae that allow them to detect afemale’s wing beat. Other flies may also initiate coupling inthe air, and often recognition is by sight. Male or female orboth of some species secrete pheromones (sex attractants)to bring the sexes together for mating. Individuals may alsouse posturing and display in courtship. In dance flies(Empididae), which often have mating swarms, the malemay present the female with an edible lure or an inediblesubstitute to initiate mating (Cumming 1994).Many Diptera congregate at landmarks, such as hilltops,for the purpose of mating. Hilltopping is a widespreadmating system among insects that appears to have de-veloped in groups that are rare, parasitic, predaceous onephemeral prey, or whose larval foodplants are scatteredor rare (Scott, 1968; Shields, 1967; Thornhill and Alcock,1983). These mobile or rare species are presumably bet-ter able to find each other at landmarks such as hilltops.Diptera commonly encountered on hilltops include small-headed flies (Acroceridae), bee flies (Bombyliidae),tangle-veined flies (Nemestrinidae), bot flies (Oestridae),big-headed fl ies (Pipunculidae), f lesh fl ies(Sarcophagidae), window flies (Scenopinidae), flowerflies (Syrphidae), horse flies (Tabanidae), tachinid flies(Tachinidae), and stiletto flies (Therevidae).Mimicry. This phenomenon is particularly well devel-oped, especially among the flower flies. Many specieslook and behave like wasps or bees and, like the latter,feed on pollen and nectar. Even flower flies of the sub-family Microdontinae, which do not visit flowers, mimic

Figure 3.Typical mosquito

(Culicidae) larvae,such as, have a rigid

apical tube that allowsthem access to surface

air for breathing.(Illustration courtesyof AAFC [McAlpineet al 1981, p. 348]).

B I O D I V E R S I T Y 3 ( 4 ) 5

the flight behaviour and resting position of bees. Broweret al (1960) showed that the resemblance between abumble bee and a robber fly almost certainly protectsthe latter from predation. One species of thick-headedfly (Leopoldius coronatus, Conopidae) has even been ob-served to mimic the zig-zag flight style of its host (awasp) until it is close enough to attack (Raw 1968).IMPORTANCE OF DIPTERAMaintaining Ecosystems and the Earth. Flies are dis-tributed from the northern limit of land in the Arcticsouthward to islands near the coast of Antarctica,where midges (Chironomidae) breeding in pools arethe most southern free-living insects. Because of theirextremely wide range of larval habitats, flies are foundnearly everywhere. The feeding habits of flies haveprofound impacts on ecosystems and the Earth as awhole. Most larvae (perhaps of half the species) arescavengers (page 12) and contribute to the decompo-sition of organic material, which in turn, providesnutrients for plants, space for all organisms, and sup-port for healthy ecosystems and clean environments.Other flies (both larvae and adults) are predators (page6), parasites or parasitoids (page 8), or plant or fun-gus feeders (page 15). Each plays an important rolein maintaining the balance among populations of or-ganisms. Flies serving as pollinators (page 17) anddisease vectors (page 19), for example, contribute tothe propagation of plants and of pathogens (protozo-ans, nematodes, bacteria and viruses).The role of both larvae and adult Diptera as predators isdiscussed below by Brooks but their importance as asource of food for organisms other than Diptera is notmentioned. Many organisms are predacious on larval oradult Diptera, but their feeding is usually unselective.Spiders and several insect orders (beetles [Coleoptera],wasps and their relatives [Hymenoptera], and dragon-flies [Odonata]) include species that are major fly preda-tors. Vertebrates may be the most important predators;of the five major classes the fish probably rank first intheir effect on human ecology. Especially in northernwaters Diptera are an important food source: in a largeArctic lake on Baffin Island, insects (mostly pupae ofmidges [Chironomidae]) made up 96% of the food ofthe Arctic Char, a fish of importance as human food(Oliver 1964). This preponderance of Diptera decreasessouthward as mayflies (Ephemeroptera), caddisflies(Trichoptera), and amphipods increase, but larvae ofmidges remain important in all freshwaters. Fish havebeen used to control “pest” flies; Gambusia has beenwidely distributed as a predator of mosquito larvae. Birdsare major predators of insects and have diverse feedinghabits. Many, such as vireos and some warblers, gleanindividual insects from plants; others, such as flycatch-ers, catch individual insects in flight. Flies, particularlythe wealth of aerial swarming species, are a critical foodfor birds that feed in flight, such as nightjars, swallows,and swifts. Most small carnivorous mammals, such asshrews and their relatives (Insectivora), also consume

flies, and bats (Microchiroptera), most of which feed inflight, are major predators of Diptera.Social and economic aspectsHuman health. Because of the association with detritusand human wastes, many flies (especially house flies)transmit serious illnesses such as dysentery, cholera, andtyphoid. The most important blood-sucking flies, servingas vectors of diseases, are the mosquitoes (Culicidae) (page20). Next come the black flies (Simuliidae) of which many,especially in tropical Africa, transmit nematodes causingriver blindness (onchocerciasis) as well as other patho-gens. And the list continues (Table 6, page 20).

Medicine. Larvae of several species of blow flies eatonly dead or damaged flesh; maggots of these specieshave been useful in medicine. They are introduced towounds to remove infected tissues and speed healing.Genetic study. Small fruit flies of the genus Drosophila,because of their ease of rearing, short life cycle and pos-session of giant polytene chromosomes in the cells ofsalivary glands, have been the subject of far more ge-netic study than any other group of organisms (page 22).It is a strange chance that they have also had perhaps thegreatest specific diversification in a limited area of anygroup – the genus has developed at least 500 species,with many more likely to be found, in the 20,000 km2 ofthe Hawaiian Islands.Forensic science and archaeology. Because many fliesdevelop in the bodies of dead vertebrates, and becauseseveral species leave behind a heavily sclerotized (hard-ened) puparial case that may persist for centuries, fliesare the most important organisms for forensic study. Theyare particularly valuable in determining the age of ca-davers from a range of a few hours to a few years (page20). They are also significant in the archaeological studyof waste middens.Agriculture and forestry. Flies in only a few familiesare plant pests; of these, three families are of major eco-nomic importance. The fruit flies (Tephritidae) includethe Mediterranean Fruit Fly and many other fruit peststhat have caused enormous losses (page 16). They aresubject to strict quarantine in many countries, especiallythe USA. A few species of root maggot fl ies(Anthomyiidae) are serious pests of root crops and plantseeds. Gall midges (Cecidomyiidae), with many thou-sands of plant-feeding species, can be serious pests ofcereal crops. These midges are also pests of many coni-fers where they cause major damage to needles and todeveloping seeds. Flies also seriously affect livestock.Black flies, screw-worms, bot flies, stable flies, and hornflies are often damaging or deadly to farm animals andrequire expensive control methods (page 12).

ABOUT THE AUTHORRichard Vockeroth is ascientist at theCanadian NationalInsect Collection(Agriculture and Agri-Food Canada inOttawa, Ontario).Much of his researchfocuses on thesystematics of flowerflies (Syrphidae) buthis knowledge extendsthroughout the Diptera.He has published onroot maggot flies(Anthomyiidae),gallmidges(Cecidomyiidae),mosquitoes (Culi-cidae), long-leggedflies (Dolichopodidae),shore flies(Ephydridae),Muscidae, fungus gnats(Mycetophilidae),Opomyzidae,Pachyneuridae, mothflies (Psychodidae),dung flies(Scathophagidae),flower flies(Syrphidae), andTethinidae. You canreach him at J.R.Vockeroth, SystematicEntomology, AAFC,Ottawa, Ontario,Canada K1A 0C6

Flies are found nearly everywhere and theirfeeding habits have profound impacts on eco-systems and the Earth as a whole.

T R O P I C A L C O N S E R V A N C Y6

AUDACIOUSPREDACIOUS LIFESTYLES

Scott E. Brooks

Predacious flies, like other predators, acquire energy bykilling and consuming two or more prey organisms duringtheir lifetime. They play an important ecological role asnatural enemies of a wide variety of organisms. Theircollective prey includes insects, molluscs, crustaceans, andeven vertebrates, in an equally wide variety of habitatsranging from terrestrial to marine. Of the 128 currentlyrecognized families of Diptera, 42 are known to includepredacious members. Most flies exhibiting predaciousbehaviour do so as larvae, the main feeding stage.However, a number of Diptera are predacious as adults.

PREDACIOUS LARVAETable 1 provides a complete list of the 39 families ofDiptera that include predacious larvae as well as noteson their ecology. This table reveals the incredible diver-sity of prey organisms exploited by fly larvae. Severalof these families are of particular interest because theykill economically important or pest species; others areinteresting because of their unusual lifestyles.Predacious fly larvae that have a human or economic impactcan be organized into groups based on their prey organisms.

Beetles: Larvae of several families feed on wood-boring or soil-dwelling beetle larvae. These includelong-legged flies (Dolichopodidae) of the genusMedetera and flutter flies (Pallopteridae), both preda-tors of bark beetle larvae (Scolytidae), and stilettoflies (Therevidae), which attack root-feeding beetlelarvae such as wireworms (Elateridae) and whitegrubs (Scarabaeidae).

Bugs (Sternorrhyncha [Homoptera]): Larvae offlower fl ies (Syrphidae) and aphid fl ies(Chamaemyiidae) are the most significant predatorsof aphids, adelgids, and scales. Several species ofaphid flies have been used as biocontrol agents ofthe Balsam Woolly Aphid in North America. Somegrass fl ies (Chloropidae), small fruit f l ies(Drosophilidae), and scuttle flies (Phoridae) are alsokey natural enemies of various Sternorrhyncha.

Flies: The muscids Myospila meditabunda and Ophyraaenescens are major predators of both House Fly andStable Fly maggots in manure piles (Skidmore 1985).In aquatic habitats phantom midge larvae(Chaoboridae) prey on mosquito larvae, and somedance flies (Empididae) feed on black fly larvae. Interrestrial habitats the larvae of Empididae also ap-pear to prefer Diptera larvae as prey (Cumming andCooper 1993).

Snails: Many species of marsh flies (Sciomyzidae)are voracious predators on the eggs, juveniles, andadults of aquatic snails, including some that harbourparasitic diseases of humans and domestic animals(Berg and Knutson 1978). Each sciomyzid larva maykill and feed on up to 30 snails during its development.

The scuttle fly Megaselia aequalis is predacious onslug eggs.

Eggs: Several families of Diptera include memberswhose larvae prey exclusively on the eggs of otherorganisms. Larvae of Elassogaster linearis(Platystomatidae) prey on the eggs of the migratorylocust. In contrast, certain species of grass flies, smallfruit flies, and scuttle flies are potentially detrimen-tal because they prey on the eggs of beneficialarthropods including dragonflies, mantids, and spi-ders or of vertebrates such as frogs.

WEIRD LARVAL LIFESTYLESOf all the predacious fly larvae, some of the weirdestand most fascinating lifestyles are observed in the ob-scure families Dryomyzidae and Vermileonidae. Take,for example, the relationship between Oedoparena glauca(Dryomyzidae) and intertidal barnacles: Adults deposittheir eggs on closed barnacles during low tide. Eggs hatchduring a subsequent low tide period and larvae enter thebarnacles as they open, when the tide comes in. Duringhigh tide, larvae feed inside the tissues of the submergedbarnacles and in subsequent low tide periods they searchfor new prey. Once a new barnacle is found, the larvauses its mouthparts to anchor itself to the prey’s shelland then waits for the tide to come in to enter the bar-nacle and feed again (Burger et al 1980).Larvae of the family Vermileonidae are commonly calledworm lions because they are fierce predators of ants andother insects. Like the ant lions of the order Neuroptera,worm lions construct pitfall traps in the soil. The wormlion waits in the bottom of the pit for an unfortunatepedestrian to fall in, at which time it attacks. Once it hasfinished feeding the worm lion tosses the victim’s corpsefrom the pit (Wheeler 1930; Teskey 1981b).

PREDACIOUS ADULTSCompared to the many families that include predatorylarvae, relatively few families of flies have developedpredacious habits as adults. Of the ten families listedbelow, the majority of predatory adults are found in theAsilidae, Dolichopodidae, and Empididae: Root-maggot flies (Anthomyiidae): Most adult an-

thomyiids feed on honeydew, nectar, dung and decay-ing organic material; however, the genusParaprosalpia has been reported to be predacious(Smith 1978).

Robber flies (Asilidae): Adult robber flies are large-to moderate-sized flies equipped with strong legs tocapture prey and syringe-like mouthparts to injectparalyzing venom and digestive saliva. Prey itemsinclude the adults of most insect orders as well assome other arthropods, which are usually captured inflight (Wood 1981). In general, robber flies tend tospecialize on larger prey than other predatory flies,occasionally taking insects over twice their own size(Platt and Harrison 1995). Net-winged midges (Blephariceridae): The females

of most species of this obscure family have well-de-

Table 1.(Opposite page).

Families of Dipterawith predacious

larvae including noteson their ecological

roles. Unlessotherwise indicated,

the informationpresented here was

extracted fromMcAlpine et al (1981,

1987), Ferrar (1987),and references cited in

these sources.

ABOUT THE AUTHORScott Brooks is a PhD

student at McGillUniversity and is

studying thesystematics and higher

classification of long-legged flies

(Dolichopodidae). Hehas also published

articles on gall-inducing wasps of thefamily Cynipidae. You

can reach him atLyman EntomologicalMuseum, Department

of Natural ResourceSciences, McGill

University, Ste-Anne-de-Bellevue, QC H9X

3V9 Canada;scottebrooks@hotmail.com

B I O D I V E R S I T Y 3 ( 4 ) 7

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T R O P I C A L C O N S E R V A N C Y8

veloped mouthparts that they use to slash open thebody of their prey. Typical prey includes midges andsmall crane flies (Hogue 1981).

Biting midges (Ceratopogonidae): Many female bit-ing midges that are not blood feeders have become pre-dacious in order to acquire the protein needed to de-velop their eggs. These females hunt in the male-domi-nated mating swarms of other flies (mainly othermidges) and mayflies (Ephemeroptera) (Downes 1978).

Long-legged fl ies (Dolichopodidae): Mostdolichopodids feed primarily on small soft-bodied in-sects and other invertebrates including Diptera lar-vae, springtails, aphids, thrips, mites, and smallworms. Species of the large and abundant genusDolichopus are notable predators of mosquito larvae.Dolichopodids do not typically capture prey in flightbut instead “graze” in areas where slow moving orconfined prey are abundant (such as mosquito ormidge larvae in small pools), often grabbing prey di-rectly with the labellum of their mouthparts.

Dance flies (Empididae) (Figure 4): Adult danceflies specialize on small prey, mainly the adults ofswarming or emerging flies. Most empidids capturetheir prey in flight and feed by sucking out the semi-fluid tissues through punctures made in the cuticle.Adults of the subfamily Tachydromiinae are impor-tant natural enemies of a variety of agricultural pestsand show strong potential to be used as biocontrolagents (Cumming and Cooper 1993).

Shore flies (Ephydridae): In contrast to the adults ofmost shore flies that feed mainly on microscopic algae,the genus Ochthera is predacious and has large, mantid-like raptorial forelegs that are used to capture midge lar-vae and other small adult insects (Wirth et al 1987).

Stilt-legged flies (Micropezidae): Smith (1978) re-corded predacious behavior in some adult micropezids.

Muscid flies (Muscidae): Predacious behaviourin adult muscids is restricted to a few genera in thesubfamily Coenosiinae, which prey mainly on otherflies including midges, sand flies, mosquito larvae,mil ichi ids , anthomyiids , and other muscids(Skidmore 1985). Dung flies (Scathophagidae): All adult dung flies

are predacious on insects and other invertebrates.

PREDATION AND SEX IN ADULT DIPTERAOf all the adult flies that have developed predacious ten-dencies, the most fascinating behaviours are seen among

the members of the Empididae and Ceratopogonidae.The adults of both dance flies and biting midges

form swarms where the sexes meet tofind a mate; however, for someof these flies, sex and preda-tion have become intimatelyentwined. In a few special-ized genera of dance flies,

males fly into the matingswarms bringing uneatenprey items (usually small

midges) that they offer as nuptial gifts to the females inexchange for sex (Cumming 1994). In an equally bizarrescenario, the females of some biting midges have evolvedcannibalistic tendencies and hunt in the mating swarmsof their own males. Once in copula, the female piercesthe male’s head with her mouthparts, injects digestiveenzymes, and sucks the male dry. Throughout this or-deal, the genitalia of the male remain firmly attached tothe female (often permanently) and eventually the restof his dried carcass breaks away (Downes 1978).

INTIMATE NEIGHBOURS:PARASITOIDS AND PARASITES

Jeffrey H. Skevington

The parasit ic Diptera exhibit some of the mostspecialized behaviours in the insect world and displayevery behaviour imaginable. This is the stuff that horrorfilms are made of! Unlike the amazingly successfulparasitic Hymenoptera that appear to have evolvedparasitic behaviour only once, flies have had the plasticityto develop parasitic lifestyles over 100 times (Feener andBrown 1997). Thirty-one families of flies have at leastsome species that have adopted parasitic existences(Tables 2 and 3).

Before discussing some of these lifestyles, it is necessaryto define some terms. Parasitic Diptera are usually dividedinto two groups, the true parasites and the parasitoids:True parasites live in intimate association with a host

from which they obtain food and usually other ben-efits (such as shelter and transportation) at the host’sexpense. They cause some degree of overt damage butusually do not kill their host.

Parasitoid larvae feed upon living host tissues in anorderly sequence until the host is killed, with death tothe host occurring only after larval development ofthe parasitoid is complete.

As with most ecological definitions, these definitionsbreak up a biological continuum. Parasitoid behaviourscharacterize the range of feeding habits that are inter-mediate between the parasitic and predacious ends of thiscontinuum. For example, how should we classify someof the snail-feeding marsh flies (Sciomyzidae)? Theseflies exhibit every range of behaviour: some species arepredators that quickly kill and consume several snailswhile other species are true parasitoids that develop inonly one host and do not kill it for several days. Someintermediate species cannot easily be shoehorned intoeither ecological guild. Despite some difficulties like this,there are advantages to treating the groups separately andwe have done our best to partition these behaviours.In this paper, I will not be treating transient blood-suck-ing species that are occasionally classified as parasites(Athericidae, Ceratopogonidae, Culicidae, Glossinidae,Muscidae [most Stomoxyinae], Psychodidae,Rhagionidae, Simuliidae, and Tabanidae) orkleptoparasites that steal their hosts from other species

Figure 4.Hemerodromia

rogatoris, a dance fly(Empididae). Larvae

feed on black flies(Simuliidae) and other

small aquaticinvertebrates; adults

are also predatory anduse their enlarged

front legs forcapturing prey in the

same way thatpreying mantids do.

(Illustration cour-tesyof AAFC[from

McAlpine et al 1981,p. 607]).

B I O D I V E R S I T Y 3 ( 4 ) 9

(Anthomyiidae [Eustalomyia, Leucophora], Braulidae,Phoridae and Sarcophagidae [Miltogramminae]).PARASITOIDSDipteran parasitoids include about 16,000 species, whichis equal to about 20% of the total number of insect parasi-toids (Feener and Brown 1997). Wasps and their relatives(Hymenoptera) account for 78% of the parasitoid species.Parasitoidism occurs to a smaller degree in three otherinsect orders: beetles (Coleoptera), butterflies and moths(Lepidoptera), and lacewings and their relatives(Neuroptera). Twenty-four families of flies contain at leastone parasitoid species (Table 2). Dipteran parasitoids donot paralyze their hosts or arrest their development withvenom as do many Hymenoptera (Feener and Brown1997). A unique developmental feature of some speciesinvolves the survival of their hosts. The hosts of sometachinid fly species, for example, survive and produce vi-able offspring (English-Loeb et al 1990). Although rare,this non-lethal parasitoidism may have important evolu-tionary consequences for both the hosts and parasitoids.Twenty-two orders (five phyla) of hosts are attacked byfly parasitoids, more than in any other group of parasi-toids (Eggleton and Belshaw 1992, 1993; Ferrar 1987). Incontrast, hosts of the more diverse parasitic Hymenopteraare restricted to 19 orders, all of them arthropods. Hostassociations unique to parasitoid Diptera include flatworms(Tricladida), earthworms (Haplotaxida), freshwater andterrestrial pulmonate snails (Basommatophora andStylommatophora), woodlice (order Isopoda), scorpions,and termites (Feener and Brown 1997). All of these un-usual hosts are associated with substrate-zone habitats andreflect the importance of these habitats in the evolution ofparasitoid lifestyles in Diptera.The broadest host use occurs in scuttle flies (Phoridae),flesh fl ies (Sarcophagidae), and tachinid fl ies(Tachinidae). This diversity of host use can be explainedin the former two families because of their many evolu-tionary origins of parasitism. Diversification of host usein tachinids presumably followed acquisition of parasi-toid lifestyle since all tachinids are parasitoids. Tachin-ids also exploit insect herbivores to a greater extent thanany other group of dipteran parasitoids and this may haveled to an explosive increase in opportunities for host uti-lization (Feener and Brown 1997).Most families are restricted to a smaller range of hosts. Big-headed flies (Pipunculidae) (Figure 5) and scarab flies(Pyrgotidae) are the most restricted, possibly because of theindependently evolved, piercing ovipositor used to insert eggsinto host bodies. This specialized structure may limit theiropportunities for host range expansion (Feener and Brown1997). Big-headed flies are the hummingbirds of the insectworld. They are accomplished hoverers and sometimes evenfly backwards for short distances. With huge eyes that aidboth their flight style and host searching, females pounce onnymphs of their hosts (leafhoppers and their relatives[Homoptera, Auchenorrhyncha]). They then often fly into theair with them before ovipositing and dropping them (May

1979; Williams 1919). Pyrgotids are also specialized andparasitise June beetles (Scarabaeidae). Onespecies attacks flying beetlesthat quickly close theirwings and fall toground withthe fly stilla t tacheda n dt h eo v i -positor wedged betweentheir protective fore-wings (elytra). An-other species ovipo-sits in the anal opening ofbeetles while they are on the ground (Ferrar 1987). Both fami-lies are important natural regulators of their hosts’ numbers.

Finding A Place To Stay…And Eat Other host-findingmethods employed by adult flies are almost as varied as theflies themselves. In general, the flies are drawn by signalsfrom the hosts’ microhabitat, activities of the hosts, or di-rectly from the hosts themselves. Female flies often usesignals associated with sexual communications of hosts. Awide range of dipteran parasites use this strategy of hostlocation whereas few hymenopteran parasitoids have evolvedthe means to exploit the communication systems of theirhosts (Vinson 1984). One of the most impressive breachesof host communication involves species of Ormia(Tachinidae). Through a specialized tympanal hearing or-gan, Ormia females are attracted by the mating songs ofmale crickets (Gryllidae) and deposit fully developed eggson or near these animals (Robert et al 1992). A parallel ex-ample is that of the flesh fly Colcondamyia auditrix femalesorienting themselves to the mating song of the cicadaOkanagana rimosa (Soper et al 1976).

A more common way of detecting hosts is by detectingchemical cues. For example, Trichopoda pennipes(Tachinidae) is attracted to the aggregation pheromoneof its stinkbug host (Nezara viridula, Pentatomidae)(Aldrich et al 1989). Scuttle fly parasitoids of ants alsoappear to use olfactory signals as orientation cues andare often attracted to nest sites, recruitment trails, or evenalarm pheromones of their hosts (Brown and Feener1991; Feener et al 1996).

Associative learning is probably the most common methodof host location in Hymenoptera. For example, if femaleMicroplitis croceipes braconids discover host frass (excre-ment) in association with a volatile chemical from a foodplant, they subsequently orient towards this new stimulus(McCall et al 1993). This strategy is likely common indipteran parasitoids but so far has only been reported inDrino bohemica (Tachinidae). A few flies exhibit the samebehaviour as antbirds that hunt displaced prey animals aroundthe periphery of army ant swarms. As advancing army antcolumns flush potential victims from their hiding places,Calodexia (Tachinidae), Androeurops (Tachinidae), and

Figure 5.Big-headed flies(Pipunculidae) such asPipunculus luteicornisare internal parasi-toids of variousAuchenorrhyncha(Homoptera) families,especially leafhoppers(Cicadellidae),delphacidplanthoppers(Delphacidae) andspittlebugs(Cercopidae).(Illustration courtesyof AAFC [fromMcAlpine et al 1987,p. 745 – misidentifiedas Pipunculus ater]).

ABOUT THE AUTHORJeff Skevington is aNatural Sciences andEngineering ResearchCouncil of Canadapostdoctoral fellowwith Agriculture andAgri-Food Canada inOttawa, Ontario andMcGill University inMontreal, Quebec.His research focuseson the systematics ofbig-headed flies(Pipunculidae) andmorphological andmolecularphylogenetics ofhigher flies. Jeff hasalso published paperson robber flies(Asilidae), stilettoflies (Therevidae),flower flies (Syr-phidae), dragonfliesand damselflies (Odo-nata), and birds.Address and emailsgiven on page 3.

T R O P I C A L C O N S E R V A N C Y10

Stylogaster (Conopidae) (Figure 6) attack cockroaches andrelated insects. The associated melee is impressive to watchas ants boil over the ground, hidden animals emerge fromcover and flee, hundreds of parasitoid flies swirl and dart inall directions, and birds whirl around.Letting The Larvae Do The Work In contrast to theabove-mentioned flies, small-headed flies (Acroceridae), beeflies (Bombyliidae), blow flies (Calliphoridae), tangle-veined flies (Nemestrinidae), rhinophorid flies(Rhinophoridae), and many tachinid flies (Tachinidae) makeno attempt to oviposit directly on their hosts. Instead theysimply broadcast their eggs in huge numbers in habitats mostlikely to be occupied by potential host species and rely ontheir larvae to do the work. For example, each female small-

headed fly lays up to 4,000 eggs, and in order to locate theirspider hosts, the larvae crawl or jump. On finding a suitablehost, the larvae burrow through its exoskeleton and attachto a book lung so that they can breath outside air. They re-main there from four months to several years in diapause(arrested development). After breaking diapause, they usu-ally feed voraciously and develop quickly before finally kill-ing their host (Schlinger 1981). An advantage of this host-finding strategy is that it allows access to hosts that are in-accessible to adult flies (for example, those in soil or wood).Some parasitoids (such as other species of tachinids) havelarvae that do not actively seek out a host but instead wait inambush. A remarkable specialization used by one lineage oftachinids (Goniini) relies on the hosts ingesting their eggs.

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;)7891rarreF(secnatsmucricemosnisdiotisarapevitatlucafemocebtahtsregnevacs ihcirdlaaiymodinhcarA tseroFfotnegalortnoclarutanrojamasi(srallipretaCtneT airtssidamosocalaM ;)b7891llewehS() ahpixosealB emosdna agahpocraS yamdna)eadidircA(sreppohssargfosdiotisarapera

.)7891rarreF;1991eniplAcMdnasselloC(lortnoclacigoloibnitnatropmieb

)seilfhsram(eadizymoicS stibahdiotisarapevloveotdednetevahscsullomlairtserretgnikcattaseiceps;srotaderpyllarenegeraeavralsascsullomcitauqakcattatahtseicepS;)7891rarreF( sinnepirginaretpondileP (sedepillimfodiotisarapasi)eaniiymoeahP( suluiotammO .)0991latealaV()

)seilfdinihcat(eadinihcaTkcattaoslayeht;stsohtneuqerftsomehteraaretpoeloCdnaaretpodipeL;stcesniylevisulcxetsomla,sdoporhtrafosdiotisarapylevisulcxeerasdinihcaTseilfrewolf,)aretpohtrO(sevitalerriehtdnasreppohssarg,)atyhpmyS,aretponemyH(seilfwas,)aretporeteH(sgubeurt,)aretpamreD(sgiwrae

;)7891rarreF;1791weksA()adopolihC(sedepitnecdna,)eadilupiT(seilfenarc,)eadinabaT(seilfesrohdnareed,)eadihpryS( sinnepinipsairhtrairT.)c7891dooW(tsepsihtkcattaotaciremAhtroNotnidecudortnineebsahdnasgiwraenaeporuEskcatta

)seilfdicsinihcat(eadicsinihcaT aidnubiB .)7891rarreF()eadiinrutaS(eapuphtomklismorfderaerneebsah

Table 2. Families of parasitoid

Diptera includingnotes on their

ecological roles.Records that are

uncertain are markedwith an asterisk (*).

B I O D I V E R S I T Y 3 ( 4 ) 11

Tiny, microtype eggs are laid in huge numbersby these tachinids in areas frequented by po-tential hosts. The eggs are stimulatedto hatch by a combination ofsaliva, mechanical rupture,and high gut pH of the host.How the Hosts Defend Them-selves. Host defences againstattacking adult flies areoften impressive. Manyant species have unique de-fensive postures or move-ments that make oviposi-tion by phorid parasitoidsdifficult and dangerous. Apocephalus (Phoridae) femalespounce on worker leaf-cutter ants (Atta) in an attempt tolay an egg on their neck. These flies are regularly killedby very small workers that ride on the piece of leaf car-ried by their larger sibling (Askew 1971; Feener and Moss1990). In Neodohrniphora curvinervis, another scuttle flyparasitoid of leaf-cutting ants, oviposition is through theback of the head. Defending ants may damage ovipositingfemales with blows to the body or by pinching the fly be-tween their head and thorax (Feener and Brown 1993).

Of course, getting an egg or larva past the host’s firstline of defences and inside the host is just the first step.The host’s immune system usually responds to internalinvaders by attempting to encapsulate and suffocate theinvader in a layer of blood cells. Some flies have turnedthis to their advantage by building a respiratory funnelfrom the products of the host’s immune response (Salt1968). This funnel gives the parasitoid larva access tofresh air through the host’s tracheal system or through ahole in the host’s exoskeleton. Many fly parasitoids main-tain contact with outside air in this way (e.g. Acroceridae,Bombyliidae, Cryptochetidae, some Calliphoridae,Nemestrinidae, Rhinophoridae, most Tachinidae). Otherflies sidestep the host encapsulation response by movinginto specific tissues that do not elicit immune responses(nerve ganglia, muscles, glands). Larvae remain in theseprotected locations until they are ready to consume thehost. It is not yet known how larvae of Pipunculidae andPhoridae avoid the immune response of their hosts.

TRUE PARASITESAs discussed earlier, parasites live in intimate association withtheir host but do not usually kill it. The relative size of thesymbionts is the primary factor that differentiates parasiticfrom parasitoid lifestyles. In general, if the host is consider-ably larger than the parasite, a relationship can developwhereby the host is not killed. Some of these relationshipsare obvious extensions of more familiar relationships. Forexample, louse flies (Hippoboscidae) and bat flies(Nycterobiidae and Streblidae) have simply taken the blood-feeding habits of flies like mosquitoes to a new level. Insteadof taking a quick blood meal and leaving the host as do mos-quitoes, these families have become intimately associated withtheir hosts. Louse flies and bat flies spend most of their adult

lives on their hosts where they feed on blood. Larvaeof all three families develop in the uterus of females

where secretions from glands nourish them until theyare fully developed (Maa and Peterson 1987; Peterson

and Wenzel 1987; Wenzel and Peterson 1987).These larvae immediately form puparia after

being extruded from the adult. Fewer thanten offspring are produced per parent.

Table 3 lists the twelveknown families of flies

that contain para-sitic species.Some of thesefamilies contain

only one lineage or even a single species that has devel-oped this specialized lifestyle. For example, larvae of thebizarre Australian grass flies (Chloropidae) calledBatrachomyia live under the skin on the backs of frogs(Sabrosky 1987b). Another unusual relationship exists withCladochaeta (Drosophilidae) and their spittlebug hosts(Cercopidae). Despite the small size of the host, as manyas three larvae of these tiny flies may feed externally onan individual spittlebug without killing it, much as suck-ing lice feed on a host mammal (Grimaldi & Nguyen 1999).

One of the most successful parasitic dipteran families isthe Oestridae (bot and warble flies). All are parasites ofmammals (and rarely birds) as larvae and most are highlyhost specific (Wood 1987a). Fully developed eggs areusually laid on the host although flies in the tribe Oestrinideposit live larvae into the nostrils of their hosts. Firstinstar larvae tend to be active and migrate to specificbody regions. Later instar larvae tend to be sedentaryand feed at these sites (in the gut wall, respiratory pas-sages, or pockets under the skin). One infamous speciesthat regularly attacks humans has developed very atypi-cal behaviour. The Human Bot Fly (Dermatobia hominis)is common in parts of the Neotropical region and attacksa wide variety of hosts including birds and many mam-mals. Females catch blood-sucking flies, such as mos-quitoes or stable flies, and lay eggs on them. These eggshatch when they are exposed to the body heat of a poten-tial host (i.e. when the mosquito goes for a blood meal).A larva then leaves the mosquito and burrows under theskin of the mammalian or avian host where it developsover the course of about six weeks.

Another infamous group of parasitic flies are the screw-worms – a common name for several unrelated speciesof fly larvae that often enter the body via wounds. Mi-nor injuries, such as tick bites, may quickly becomemortal wounds from the activities of these flies. The bestknown of these is the Primary Screw-Worm, Cochliomyiahominivorax (Calliphoridae). This species attacks a va-riety of mammals including humans and livestock. InNorth America, it was responsible for huge economiclosses in the cattle industry annually and is particularlynoteworthy for having led to the sterile male release tech-nique in insect control developed by Knipling (1960),

Figure 6.Ant flies (Conopidae,Stylogastrinae), suchas this Brazilianspecies of Stylogaster,are parasitoids ofcockroaches(Blattaria), crickets(Gryllidae), andpossibly calyptrateflies (Muscidae andpossibly Calliphoridaeand Tachinidae); theyare often associatedwith army ants,darting after hosts thatflee from the columnof ants.(Illustrated byRoelof Idema).

T R O P I C A L C O N S E R V A N C Y12

which helped to eradicate the species from much of itsNorth American range. In this process, the species wasreared in vast numbers in “screw worm factories,” thepupae were irradiated, and sterilized males were released.Females mate only once and those that mate with sterilemales produce infertile eggs. Through these releases,unviable males quickly overwhelm the healthy males inthe population. A population of 20 million insects (halfmale) can be wiped out in only four generations by re-leasing 20 million sterile males in each generation.Florida was rid of this pest in this way in only two yearsat a cost of eight million dollars (annual damage inFlorida had been over ten million dollars).

SUMMARYAs can be seen from this overview, some of the mostbiologically fascinating and economically significantanimal activities can be attributed to the parasitic Diptera.With their diverse habits and many evolutionary origins,parasitic flies and their hosts offer us unparalleled op-portunities to examine questions in behavioural and evo-lutionary ecology (Feener and Brown 1997). Economicrepercussions of research on Diptera are far reachingand will continue to have impacts as new discoveries aremade. Like most insects, flies are relatively poorly knownand a huge proportion of the diversity has not yet beendescribed let alone studied by ecologists. Undoubtedly,many exciting discoveries remain to be made.

CLEANING UP THE WORLD:DIPTERAN DECOMPOSERS

Jade Savage

Diptera are fundamental participants in the decompo-sition process of terrestrial and aquatic systems (Speightet al 1999). About half of all Dipteran families have lar-vae that feed on decaying organic matter and many moreare indirectly associated with this substrate through pre-dation and parasitism.

While adult flies can often be caught around decompos-ing substances, it is the immature stages that are mostinvolved with the breakdown process. Voracious larvaeeat and fragment large amounts of dead organic mate-rial, extracting energy either directly from the substrateor, more commonly, from the digestion of the microf-lora associated with the substrate (Mason 1977).The role of flies in decomposition is physical rather thanchemical. The surface area of the material is increased byingestion and by boring and tunnelling through it, thus pre-paring it for further decomposition by microorganisms.Active larvae disseminate fungal spores and bacteria toregions of the substrate previously impenetrable to them,while microbial or fungal grazers stimulate the growth andalter the composition of the microbial community throughselective feeding (Moore et al 1988).Detritivores, also called saprophages or scavengers, areoften considered to be unspecialized feeders that in-discriminately consume any detritus available. Whilethe omnivorous maggots of the House Fly, Muscadomest ica , and the Lesser House Fly, Fanniacanicularis (Figure 7), support that assumption, theyare the exception rather than the rule. Instead, mostspecies have very specialized diets.Covering all specialized scavenging habits observedamong flies would be a lengthy business. This review,then, will focus only on those taxa intimately involvedwith the decomposition of three specific types of sub-strates, namely plant material, dung, and carrion. Table4 provides a summary of feeding habits found in all fami-lies with saprophagous species.PLANT MATERIALMany soil-dwelling Diptera play a significant role in therecycling of leaf litter. Primitive families are especiallywell represented among forest floor species, with thelarvae of dark-winged fungus gnats (Sciaridae), midges

YLIMAF SETONeadiiymohtnA )seilftoggam-toor( .)7891ttekcuH(seltrutdnaldnastnedorfosetisaraperasdiiymohtnaemoS

)seilfwolb(eadirohpillaC

;)sisaiym(eussityhtlaehnigniworrubybsdnuowtcilfnistoggam;loowdeliosrosnoiselnikshserfnosggeyal)iniilicuL,iniimrohP,iniiymosyrhC(smrow-wercSxarovinimohaiymoilhcoC dna anaizzebaymosyrhC ;)a7891llewehS(sdnuowlatromotniseirujnironimgninrutnetfo,slaminacitsemoddnasnamuhkcatta

;)1791weksA(ailartsuAniylralucitrap,melborpevisnepxenasipeehsfosisaiym agahpoporhtnaaibolydroC dnaliobekil-ylftobamrofeavral)ylFubmuT(emos;)1791weksA(gnitapupnehtdna)snamuhgnidulcni(lammamtsohgnivaelerofebsyadthgietuobanihtiwpoleved ilicuL suoenatucbusralimisesuaca

;)7891rarreF(snaibihpmanisisaiym arohpillacotorP .)a7891llewehS(doolbkcusdnasdribgniltsenfoniksothcattaeavral

eadinraC suretpamehsunraC .)a7891yksorbaS(snoitercesniksnodeefyamrognikcusdoolbebyam

)seilfssarg(eadiporolhC nailartsuA aiymohcartaB emos;)b7891yksorbaS;7891rarreF(egremeseilfehtnehweidsgorffo%01tuoba;sgorffoskcabehtnoniksehtrednuevileavral.)1791weksA(sisaiymyrotagilbonidevlovnieraseicepsdiporolhcrehto

)seilftiurfllams(eadilihposorD ateahcodalC foyltsom(sgubelttipsfosetisaraplanretxeera aretpotsalC .)9991neyugN&idlamirG()

)seilfesuol(eadicsoboppiH cimonoceelbaredisnocfoeradna,peehsylralucitrap,slaminacitsemodkcattaemos;slammamdnasdribfodoolbnognideef,setisaraplanretxellaerastludA.)7891nosretePdnaaaM(ecnatropmi

)seilfdicsum(eadicsuMaiymoressaP dna sinrolihP emos;)7891htorekcoV&ttekcuH;7891rarreF(sdribeniressapgniltsenfodoolbkcuseavral aibotameaH dna acsobotameaH

eromdikS(gnudhserfnotisopivootylfeirbgnivaelselamefhtiw,doolbnognideefstsohetalugnuriehtnoseviltludariehtfotsomevil)eaniyxomotS(seiceps.)5891

)seilftab(eadiibiretcyN .)7891lezneWdnanosreteP(stabfosetisaraplanretxegnikcus-doolberastludA

)seilfelbrawdnatob(eadirtseOyratnemilaehtsahcussnoitacolgnideeffoyteiravahtiwdezilaicepsyreverayeht;)a7891dooW(eavralsa)sdribylerardna(slammamfosetisaraperallA

-eanilihporetsaG,eanirberetuC(seussitsuoenatucbus,)eanilihporetsaG(tcart arberetucoeN dna ainettuR snoigerlaegnyrahp-osandna),eanitamredopyH,.)1791weksA()eanirtseO,eanilihporetsaG(

)seilfreppiks(eadilihpoiP mulihpoittoeN .)a7891eniplAcM(sdribeniressapgniltsenfodoolbkcuseavral

)seilfhself(eadigahpocraSligivaitrhaflhoW lavralrofgnortsootsemocebstnafniredlofonikseht(egaforaeyenootpusnamuhgnidulcni,slammamfognuoyyhtlaehnisisaiymsesuac

rehto;)noitartenep aitrhaflhoW ;slammamrehtonisisaiymesuacseiceps aiymonidutsiC sesiotrotdnalfosisaiymdnuowesuac misilonA; esuacaiy;snoelemahcfosisaiymsuoenatucbus aihcynorcamuE ;sgnilhctahdnasggeelitpermorfderaerneebevah ateahcotoN ;7891rarreF(snaibihpmakcatta

.)b7891llewehS

)seilftab(eadilbertS foselameftpecxesetisaraplanretxeeralla;stabfosetisarapgnideef-doolberallA noretpidocsA ,suoenatucbusotnimrofsnartdnasgeldnasgniwriehtesoltaht.)7891nosretePdnalezneW;7891rarreF(stsohgnidnifretfaserutaercekil-cas

Table 3.Families of obligate

parasitic Diptera,including notes on

their ecological roles.Note that facultativeparasites, including

species thatsometimes cause

myiasis (the inflictionof wounds by

burrowing in healthytissue), are excluded.

B I O D I V E R S I T Y 3 ( 4 ) 13

(Chironomidae), march flies (Bibionidae), gall midges(Cecidomyiidae), and crane flies (Tipulidae) all com-monly found feeding on decomposing leaves. These,along with Faniidae (Figure 7), a few muscids (Mus-cidae), minute scavenger flies (Scatopsidae), lesser dungflies (Sphaeroceridae), and soldier flies (Stratiomyidae),are often part of the fauna actively working on the deg-radation of an ordinary backyard compost pile.Although best known as pests of lawns and pastures, the lar-vae of crane flies, often called leatherjackets, are especiallyimportant in forest and stream ecosystems. Many species feedexclusively on leaf litter. The experimental removal of Tipulapeliostigma from birch litter-fall was shown to greatly reducethe rate of breakdown of the substrate (Perel et al 1971). Strongmandibles allow the shredding of large amounts of dead leaves,and even enable some crane flies (Ctenophora, Epiphragma)to actively bore into rotting wood. Well sclerotized (hardened)mouthparts are present in other primitive Diptera, and spe-cies of axymiids (Axymiidae) and moth flies (Psychodidae)among others are also found tunnelling through decaying wood(Teskey 1976).Most saprophagous (detritivorous) flies have larvalmouthparts designed to feed on moist or semi-liquid food,and the immatures are normally found in rotting mate-rial with at least some degree of moisture. Rotting fruitsare attractive to many groups, including the highly di-verse small fruit flies (Drosophilidae) (Figure 12). Atthe family level, drosophilids can certainly be classifiedas general opportunists, with a strong preference for anytype of fermenting vegetal substance. Breeding mediarange from damaged cactus tissues to rotting bananas,and the small red-eyed adults are common pests of res-taurants, market places, and household garbage bins.In contrast to the versatile Drosophilidae, other familieshave specialized on a single type of decaying vegetal sub-strate. Many, and possibly all, species of the small fam-ily Periscelididae breed exclusively in fermenting sapruns (Teskey 1976), while all known species of seaweedflies (Coelopidae) feed on rotting seaweed (Ferrar 1987).DUNG AND URINEThe breakdown of vertebrate excrement is of obvious eco-nomic relevance to human societies. A single adult dairycow can produce on average nine tonnes of manure in ayear. Along with many beetles, flies must be acknowledgedfor the quick disposal of this unwieldy material. Amongthe most notorious dung and manure feeders are the lar-vae of many calyptrate Diptera, such as dung flies of thegenus Scathophaga (Scathophagidae), blow flies (Calli-p h o r i d a e ) ,

flesh flies of the tribe Raviniini (Sarcophagidae), and manymuscids, including species whose adults are often of greateconomic importance (for example, the House Fly and theStable Fly, Stomoxys calcitrans). Small acalyptrate flies,like many black scavenger flies (Sepsidae) and lesser dungflies (Sphaeroceridae), are also dependant on dung for theirdevelopment.

Ivermectin, a commonly used antiparasitic drug admin-istered to cattle, seriously interferes with the primarydipteran decomposing fauna (Madsen et al 1990). Mostof the drug is eventually egested by cattle, along withthe faeces, and acts as an efficient insecticide on mostdung-breeding flies. While the reduction of adult fliesmay be seen as a beneficial side effect of the treat-ment, the decomposition of dung pats is greatly delayedin the absence of dung-feeding larvae and could lead tothe quick fouling of pasture land.Extreme specialization is seen in the familiesMormotomyiidae and Mystacinobiidae, who share morepeculiarities than just odd-sounding names. These twofamilies are each known from a single species, and whilenot closely related, both breed in bat guano. Mormotomyiahirsuta is known from a single cave in Kenya, and boththe larvae and the spider-like wingless adults feed ex-clusively on bat dung. Mystacinobia zelandica occurs onlyin New Zealand in association with bats living in hollowtrunks of the giant Kauri Trees.

CARRIONWhen an animal dies, its carcass will be visited by asuccession of various insects, with flies often being themost diverse and abundant taxa. Blow flies, with generasuch as Calliphora and Lucilia, will be the first to arriveon a fresh carcass, soon followed by flesh fl ies(Sarcophagidae) and muscids. Blow flies are recognizedfor their ability to locate fresh carrion, and will oftenarrive on a body within minutes following death. Earlyinvaders, particularly blow fly larvae, will usually befound in large numbers, consuming most of the decom-posing flesh. In an experiment done on mouse carcasses,the larvae of a single blow fly species consumed over75% of all material decomposed (Putman 1978).

Once the body reaches a more advanced stage of decom-position, it will attract a fauna mostly feeding on putridexudates. Small fruit flies (Drosophilidae), lesser dungflies (Sphaeroceridae), black scavenger flies (Sepsidae),shore flies (Ephydridae), and flower flies (Syrphidae) (spe-cifically rat-tailed maggots: Eristalis) will often be partof that next wave. Species of skipper flies, including thesynanthropic Cheese Skipper, Piophila casei, will also be

found feeding on dead tissues, often in the later stages ofdecomposition.

The taxa mentioned above will generally be foundon an exposed carcass. When the carrion is

buried, a different fauna will invade it, andblow flies may be completely excluded byburial under a few centimeters of soil.

Figure 7.While occasionallyinvolved in cases ofhuman myiasis, thelarvae of the LesserHouse Fly, Fanniacanicularis(Faniidae), shouldalso be acknowl-edged for its abilityto complete itsdevelopment byfeeding on a largevariety of decayingvegetal or animalmatter. (Drawingcourtesy of AAFC[from McAlpine et al1987, p. 1129]).

ABOUT THE AUTHORJade Savage is a PhDstudent at McGillUniversity. She iscurrently working onthe systematics of thetribe Azeliini of thefamily Muscidae.Most azeliine larvaeare saprophagous, atleast in the earlyinstars, and arecommonly found indung, humus, orcarrion.You can reachJade at LymanEntomologicalMuseum, Departmentof Natural ResourceSciences, McGillUniversity,Macdonald Campus,Ste-Anne-de-Bellevue,QC H9X 3V9,Canada;jsavag1@hotmail.com

T R O P I C A L C O N S E R V A N C Y14

Conicera tibialis (Phoridae), also known as the Coffin Fly,will often be found in coffins or on buried bodies that wereunderground for about a year. The adult female CoffinFly, as well as other scuttle fly species, will burrow intothe soil and oviposit directly on the carcass. It is believedthat the Coffin Fly mates within the same coffin and canproduce a number of generations, without the need to goto the surface for copulation (Smith 1986).Dead invertebrates, although much smaller than theirvertebrate counterparts, are also fed upon by flies. Deadsnails are especially attractive to many Diptera, and anumber of lesser dung flies and a few muscids, shoreflies (Ephydridae), flesh flies (Sarcophagidae), and oth-ers have been reared from them.

SEILIMAF

gniyaceDrettamlategev rettamlaminagniyaceD

eadidoposinA

eadiiymohtnA

eadiietsA

eadirtsagicaluA

eadiimyxA

eadinoibiB

eadirohpillaC

eadinraC

eadiiymodiceC

eadinogopotareC

eadimonorihC

eadiporolhC

eadiisulC

eadipoleoC

eaditonotruC

eaditamosolespyC

eadispoiD

eadilihposorD)21erugiF(

eadizymoyrD

eadirdyhpE

eadiinaF )7erugiF(

eadizymoeleH

eadizymoicsoleH

eadiiymonorI

eadiinaxuaL

eadieahcnoL

eadiretpohcnoL

eadizeporciM

eadiihciliM

eadiiymotomroM

adicsuM e

eadilihpotecyM

eadiibonicatsyM

eadiireN

eadiinidO

SEILIMAF

gniyaceDrettamlategev rettamlaminagniyaceD

eadititO

eadiruenyhcaP

eadidilecsireP

eadirohP

eadilihpoiP

eaditamotsytalP

eadidohcysP

eadiretpohcytP

eadiidrahciR

eadiremolapoR

eadigahpocraS

eadigahpohtacS

eadispotacS

eadiraicS

eadizymoicS

eadispeS

eadiilumiS

eadirecoreahpS

eadiymoitartS

eadihpryS

eadizepynaT

eadinihteT

eadilupiT

eadirecohcirT

Litter, humus, com

post

Litter, humus, com

post

Plants, fruits, vegetables

Plants, fruits, vegetables

Fermenting sap

Fermenting sap

Fungi

Fungi

Decaying wood incl. tree holes

Decaying wood incl. tree holes

Seaweed

Seaweed

Fine organic particles(aquatic)

Fine organic particles(aquatic)

Sewage/organic sludge

Sewage/organic sludge

Vertebrate excrement

(fresh dung, urine)

Manure

Vertebrate excrement

(fresh dung, urine)

Manure

Bird / bat guano

Bird / bat guano

Nest material (vert. & invertebrates)

Nest material (vert. & invertebrates)

Frass

Frass

Proteinaceous matter

(milk, etc.)

Vertebrate carrion

Proteinaceous matter

(milk, etc.)

Invertebrate carrion

Vertebrate carrion

Invertebrate carrion

B I O D I V E R S I T Y 3 ( 4 ) 15

Scavenging larvae also eat insect remains. Scuttle fly(Phoridae) larvae, scavenging on miscellaneous debris andinsect parts, occasionally occupy the nests of ants and ter-mites. All known species of the North American flesh flygenus Fletcherimyia are obligate breeders in Sarraceniapitcher plants. A single larva can be found per pitcher,feeding voraciously on drowned insects (Ferrar 1987).CONCLUSIONThe grim thought that we may eventually end up as mag-got food does nothing to increase the popularity of flies.However, life itself is wholly dependant on the properrecycling of organic matter, and with the ever-increas-ing amount of waste products generated by our societ-ies, filth eaters are more important than ever. Whiletheir contribution to the cycle of life often goes unrec-ognized, saprophagous larvae can comfort themselveswith the fact that there will always be enough for dinner.

GOING VEGETARIAN:PLANT AND FUNGUS FEEDINGStéphanie Boucher and Terry A. Wheeler

Although many people associate flies with dead anddecaying material, several thousand species of flies pre-fer fresher food. For many, living plants or fungi are afood source, a shelter, and/or a mating site. Almost 40families of Diptera contain herbivorous species (Brown2001), which feed on living plants or algae (phytopha-gous flies) or on fungi (mycophagous flies). Familieslike crane flies (Tipulidae), gall midges (Cecidomyiidae),fungus gnats (Mycetophilidae), dark-winged fungus gnats(Sciaridae), scuttle fl ies (Phoridae), fruit f l ies(Tephritidae), leaf-miner flies (Agromyzidae), grass flies(Chloropidae), shore flies (Ephydridae) and root-mag-got flies (Anthomyiidae) all contain herbivorous specieswith a wide range of feeding habits.

PHYTOPHAGOUS HABITS: THE PLANT FEEDERSIn contrast to the familiar caterpillars and beetle larvae thatare exposed on their food plants while feeding, herbivorousDiptera larvae are often small and dehydrate easily. Thishas led to a variety of strategies for remaining concealedand protected in or on the host plant while feeding.Root feeders: Some fly larvae feed underground on roots.For example, some march fly (Bibionidae) larvae causedamage to the roots of vegetable crops, cereals, and grasses(Oldroyd 1964). Other root-feeding larvae, such as theCabbage Root Fly, Delia radicum (Anthomyiidae), or theCarrot Rust Fly, Psila rosae (Psilidae), can also damagemany types of cultivated plants (Ferrar 1987).Stem borers: Other larvae are stem borers, like the FritFly, Oscinella frit, the Wheat Stem Maggot, Meromyzaamericana (both Chloropidae), and the Hessian Fly,Mayetiola destructor (Cecidomyiidae). All three are con-sidered pests of cereal crops (Ferrar 1987; Gagné 1989).Another stem borer, the Bean Fly, Ophiomyia phaseoli,lives in the Old World tropics and is one of the most seri-ous pests in the family Agromyzidae (Spencer 1973). In

the predominantly predacious family Dolichopodidae, thegenus Thrypticus represents a major evolutionary changewith a switch to a phytophagous lifestyle. All known lar-vae are stem miners in aquatic monocots. Females havedeveloped a hard ovipositor for piercing plants to deposittheir eggs. Members of this genus have been consideredas potential biological control agents for water hyacinth,one of the world’s most invasive aquatic weeds.Flower head feeders: Larvae of many gall midges(Cecidomyiidae), fruit flies (Tephritidae), grass flies(Chloropidae), and root-maggot flies (Anthomyiidae) feedin flower heads. Females often lay their eggs in the flower-buds, which can affect the development of the flower(Ferrar 1987). In addition, many larvae associated withflower heads feed on developing seeds, which can causeproblems in crops like legumes. This is the case with theagromyzid Melanagromyza obtusa, a major pest of PigeonPeas in Asia. The female lays eggs inside the pod and thenewly hatched larvae start feeding on the surface of thepeas and eventually bore right inside (Spencer 1973). OtherDiptera larvae, such as those of Earomyia (Lonchaeidae)will attack conifer cones and seeds.Fruit feeders: Other fly larvae are fruit feeders and ca-pable of causing serious damage to fruit crops. The best-known fruit feeders are in the family Tephritidae, com-monly known as fruit flies. Females of fruit flies lay theireggs under the skin of fresh fruits, where the larvae willdevelop by feeding on the fleshy tissue. The larval feed-ing tracks provide entry points for bacteria and fungi whichwill cause the fruits to rot. The Mediterranean Fruit Fly(or Medfly), Ceratitis capitata, is one of the most seriousof all fruit pests (Figure 8). A well-known North Ameri-can fruit fly is the Apple Maggot, Rhagoletis pomonella,that attacks several fruits including apples, peaches, plums,pears, and cherries (Foote et al 1993).Leaf miners: Leaf mining is another specializedlifestyle in plant-eating Diptera. Although this habit isseen in a few families (e.g. , root maggot fl ies[Anthomyiidae], shore flies [Ephydridae], rust flies[Psilidae], dung flies [Scathophagidae], and fruit flies[Tephritidae]), the leaf-miner flies (Agromyzidae),with probably more than 2,000 species, is by far thedominant group. The larva usually feeds on the upperlayer of cells of a leaf just beneath the epidermis, form-ing a clearly visible mine. The mine can be linear, ser-pentine, or blotch-like; its shape and position on theleaf are often characteristic of the mining species, andcan be used to identify the fly. Some Agromyzidae,especially in the genus Liriomyza, are serious pests ofcrops and ornamental plants (Spencer 1973).Gall dwellers: Some Diptera larvae live concealed in agall, an abnormal tissue growth produced by the plant inresponse to the presence of the gall-inducer (Jolivet 1992).Although much remains unknown about their initiation,galls themselves are complex structures that can developon any part of the plant – leaves, stems, roots, fruits, orflowers. Galls are advantageous to the flies because they

Table 4.(opposite page)Feeding habits oflarvae of sapropha-gous Diptera.Compiled mainlyfrom Ferrar (1987),McAlpine et al (1981,1987), Teskey (1976),and Smith (1986).

ABOUT THE AUTHORSStéphanie Boucher isthe Curator of theLyman EntomologicalMuseum (McGillUniversity). Her mainresearch interest is thebiodiversity andsystematics of leaf-miner flies (familyAgromyzidae). Sheworks mainly onAgromyzidae living indry habitats of theYukon Territory butalso has an interest inthe NeotropicalAgromyzidae.Terry Wheeler is anAssociate Professor ofEntomology at McGillUniversity and theDirector of the LymanEntomologicalMuseum. His researchinterest is systematicsof plant-feeding flies,especially the grassflies (Chloropidae). Hehas published paperson Agromyzidae,,Carnidae, Chloropi-dae, Chyromyidae,Lauxaniidae,Mallochianamyia(Diptera, Schizophora,unplaced to family),Milichiidae,Opomyzidae, andlesser dungflies(Sphaeroceridae).You can reach them atLyman EntomologicalMuseum, Departmentof Natural ResourceSciences,McGillUniversity, MacdonaldCampus, Ste-Anne-de-Bellevue, QC H9X3V9, Canada;sboucher@nrs.mcgill.caandwheeler@nrs.mcgill.ca

T R O P I C A L C O N S E R V A N C Y16

provide food and protection to the larvae; theplant may also benefit because the gall re-

stricts damage to one part of the plant.Fly larvae of the pre-dominantly Australiangenus Fergusonina

(Fergusoninidae)have a remarkablesymbiotic associa-

tion with nematodes,which allows them by

their combined ac-tion to form galls

on myrtaceousplants (particu-

larly Eucalyptus trees). Thegall midges (Cecidomyiidae) and fruit flies (Tephritidae)have many gall-inducing species, and although the mostserious pests in these families do not induce galls, fly spe-cies that cause abnormal growths on ornamental plants areoften considered harmful (Gagné 1989).Microbial ingestion: Green plants do not have a mo-nopoly on photosynthesis and some fly larvae have takenadvantage of other organisms that photosynthesize. Lar-vae of some species of shore flies (Ephydridae) are spe-cialists on blue-green algae (cyanobacteria) – photosyn-thetic bacteria that filled the role of plants long beforeeither plants or flies evolved (Foote 1995).MYCOPHAGOUS HABITS: THE FUNGUS FEEDERS.Fungus gnats (Mycetophilidae) and dark-winged fungusgnats (Sciaridae) are examples of the many flies that eatfungi. These two families often specialize on particulargroups of fungi, especially in forests. Fungus gnats areone of the most diverse and abundant families of flies inmature forests largely because of the great diversity offungi. Dark-winged fungus gnats are also dominant in for-ests but are more familiar as the tiny black flies that some-times emerge from the soil of houseplants, where theyhave developed on soil fungi (also detritivores). Thereare often close relationships between mycophagous andapparently phytophagous habits; for example, some spe-cies of gall midges live in plant galls, but feed on a cultureof fungi apparently deposited by the female during ovipo-sition (Gagné 1989).

HOST SPECIFICITYLarvae of phytophagous Diptera often show a high de-gree of host specificity. A large number of species aremonophagous, feeding on plants of a single genus or evena single species. For example, the fruit fly Bactroceraoleae feeds only on olive fruits and the leaf-miner flyOphiomyia simplex is known only from asparagus. Atthe other extreme, some flies are polyphagous, feedingon a number of unrelated families. The Medfly (Figure8) infests over 260 different fruits, flowers, vegetables,and nuts (Foote et al 1993).Because of their close associations, the geographic distri-bution of plant-eating Diptera often follows the distribu-

tion of their host plant. As a result, many pest species feed-ing on crops have been accidentally introduced and estab-lished in new regions by humans; many North Americancrop pests are actually of European or Asian origin.ECONOMIC IMPORTANCEMany plant-feeding flies are important pests of crops. Phy-tophagous species like those mentioned previously costmillions of dollars annually in control efforts and croplosses in fruits, vegetables, and cereals. Even mycopha-gous species can be costly pests in mushroom houses.Not all plant feeding is negative. Many plant species areweeds and flies that feed on such plants and reduce theirgrowth and survival are considered beneficial, and are some-times used as biological control agents. A successful ex-ample involves species of fruit flies (Tephritidae) that havebeen widely introduced to control weeds like Canada thistleand knapweeds (Ferrar 1987). There are also aquatic her-bivores like shore flies (Ephydridae) that mine the leaves ofinvasive wetland plants (Foote 1995).Most often it is the larvae that cause damage to the hostplant. Like many flies, adults of herbivorous species usu-ally feed on pollen and nectar. These adults are also ben-eficial because they are major plant pollinators, espe-cially in northern ecosystems.

FLOWERS, POLLINATION,AND THE ASSOCIATEDDIVERSITY OF FLIES

Peter G. Kevan

Flies must eat, and their names often suggest theirdiverse sources of food: fungus gnats (Mycetophilidae),fruit flies (Drosophilidae and Tephritidae), flower flies(Syrphidae), blow flies (Calliphoridae), dung flies(Scathophagidae), horse flies (Tabanidae), flesh flies(Sarcophagidae), and so on. One of the most importantfood sources is from flowers, especially with respect tothe adult energetic requirements for flight in dispersing,finding mates, mating, and searching out sites foroviposition (Hocking 1953, Larson et al 2001). Flowersoffer an open banquet, well advertised by their colours,sizes, shapes, and scents (Kevan and Baker 1983; Proctoret al. 1996). Aside from the menu of sugar-rich nectar andprotein-rich pollen, flowers sometimes give other rewardssuch as protection, places to find mates, and ovipositionsites (Kevan and Baker 1983; Kevan 2001).Nectar, the main staple for fuelling flies’ activities, ismore nutritious than sugar water. It contains varioussugars in various ratios, small amounts of amino acids,sometimes oils, vitamins, minerals, phenolics, and othercompounds, some of which reflect the nature of theflowers’ pollinators (Baker and Baker 1990). Flowersused, and sometimes pollinated by, short-tongued fliessometimes present nectar with sugars so highlyconcentrated that the flies must spit onto the crystals todissolve them for ingestion. Long-tongued flies, such astangled-veined flies (Nemestrinidae) and some horse flies

Figure 8.All fruit flies

(Tephritidae) feedupon living planttissues and many

species are economi-cally important aspests or as control

agents for weeds. TheMediterranean Fruit

Fly (or Medfly),Ceratitis capitata, is

one of the mostserious of all fruit

pests. (Illustrated byRoelof Idema).

B I O D I V E R S I T Y 3 ( 4 ) 17

(Tabanidae) of South Africa (Goldblatt and Manning2000), have no other source of sugar for energy thandilute protected nectar provided by flowers with deeptubes. A wide variety of fl ies feed upon pollen,notoriously the flower flies (Syrphidae). It is presumablythe source of protein for general nutrition and maturationof the ovaries and testes (Kevan and Baker 1983). Someother flies appear to have switched from feeding on bloodto feeding on pollen, as in some biting midges(Ceratopogonidae) that bite pollen to remove thenutritious protoplast from within (Downes 1958).Some flowers entice flies to visit them, but may not offerany reward except shelter. The aroids (Araceae) andAristolochia species are infamous for temporarily trappinginsects (mostly flies and beetles) (Proctor et al 1996). Theyretain the insects for about a day in the younginflorescences that have receptive stigmata, but allow themto escape as the stigmata age and the anthers split openand dust the captives with pollen. Cross-pollination is thuseffected. Some flowers, such as Stapelia species(Asclepiadaceae) of South Africa and the floral giants ofthe genus Rafflesia (Rafflesiaceae) of South East Asia,dupe saprophagous flies into visiting the flowers and layingeggs. In most instances, the maggots starve and die. Thescents of these flowers are of carrion, dung, or animalmusk, and the colours tend to be dull and also attract bymimicking oviposition sites (Proctor et al 1996).Although there are huge numbers of records of flies as flowervisitors, proof of their importance as pollinators is oftenwanting. To be a pollinator, a fly must carry pollen in sucha way that it is transferred from the anthers to the stigmata.This may be simply accomplished within the same floweror on the same plant. Self-pollination may be achieved, butif the plant is self-incompatible, it is of no consequence.Fertilization must occur if pollination is to be consideredsuccessful from the plant’s perspective (Free 1993, Roubik[ed] 1995). From the fly’s perspective, a visit to a flower isa success if a reward is obtained, regardless of whether ornot pollination or fertilization results.The process of fertilization in flowering plants (Angio-spermae) sets them aside from other plants, and basedon fossil and molecular evidence, it is suspected thatthis system of fertilization (aka pollination) arosemore than 135 million years ago (Sun et al 1998;Sanderson and Doyle 2001). It is also suspectedthat insects have been involved since the verybeginning. Pollination by insects (entomo-phily), and particularly Diptera (myio-phily), can be considered to be basicto angiosperm evolution (Labandeira1998; Grimaldi 1999).The long-horned f l ies(suborder Nematocera)mostly have short suctorial and lappingmouthparts that restrict them to feeding onexposed nectar at open flowers, such asroses, euphorbs, saxifrages, and carrots

(Larson et al 2001). Some may be more important aspollinators than usually considered. In particular, thereare orchids (e.g. Platanthera (Habenaria) spp.) that arepollinated by mosquitoes (Kevan et al 1993). Cacoa(Theobroma cacao), from which chocolate is made, ispollinated by several kinds of midges (Ceratopogonidaeand Cecidomyiidae) (Free 1993; Roubik [ed] 1995).Moth flies (Psychodidae) are temporarily trapped in andpollinate Lords and Ladies (Arum maculatum) in Europeand North America (Proctor et al 1996). It is a commonmyth that black flies (Simuliidae) pollinate blueberryflowers (Hunter et al 2000).Among the short-horned flies (suborder Brachycera),there are many records of flower visiting (Larson et al2001). The bee flies (Bombyliidae), with their long out-stretched proboscides, are often seen sucking the nectarfrom flowers and form one of the most recognizable anddiverse families at flowers (Figure 9). It is interestingthat the relationship of the flower-loving fl ies(Apioceridae) and flowers has been rarely documented,even for the somewhat misnamed and endangered DelhiSands Flower-Loving Fly (Rhaphiomidas terminatusabdominalis) that actually belongs in the family of Mydasflies (Mydidae). Some small-headed flies (Acroceridae),with special hairs, seem especially adapted to carryingand feeding on pollen. The Brachycera contains some ofthe most highly adapted flower-visiting and pollinatingflies as exemplified by the South African tangle-veinedflies (Nemestrinidae) and some horse flies (Tabanidae),which have extremely long tubular mouthparts that mayexceed the remaining body length (Goldblatt and Manning2000). A number of dance fl ies (Empididae) areimplicated as significant pollinators because of their longmouthparts, and some genera, such as Anthalia,Anthepiscopus, and Iteaphila, are known to be obligatepollen feeders (Grimaldi 1999). The flower fl ies(Syrphidae) are perhaps the Diptera most well knownas flower feeders (Figure 9a). They feed on nectar andon pollen. Sometimes pollen feeding by female flowerflies has been correlated with their ovarian maturation

(Schneider 1969). The scuttle flies (Phoridae) areprobably under-appreciated as flower visitors andas possible pollinators, and most records come fromthe tropics (Larson et al 2001).

Flower feeding is recorded in about twodozen families among

the house fl ies andtheir re la t ives ( the

Schizophora) (Larsonet al 2001). However, for most, therecords a re sparse and mos t ly

descriptive. Among the small fruitf l i e s (Drosophi l idae)are species that feed on

nec ta r tha t suppor t s a microf lo ra o f yeas t s(Lachance et al 2001). Some of these species are

impor tan t in c rop po l l ina t ion , such as fo rmangoes, and for horticultural seed production in

Figure 9.The bee flies(Bombyliidae), withtheir long out-stretched proboscides,are often seen suckingthe nectar fromflowers and form oneof the mostrecognizable anddiverse families atflowers. Bombyliuspygmaeus is typical ofthese spectacular andoften hairy flies.(Drawing courtesy ofAAFC [fromMcAlpine et al 1981,p. 589]).

ABOUT THE AUTHORPeter Kevan is aProfessor ofEntomology at theUniversity of Guelph.His research isprimarily onpollination but he hasalso published paperson other aspects ofecology such asthermoregulation.Much of his researchhas been conducted inthe CanadianArctic.You can reachPeter at Departmentof EnvironmentalBiologyUniversity of Guelph,Guelph, ON N1G 2W1,Canada;pkevan@uoguelph.ca

T R O P I C A L C O N S E R V A N C Y18

enclosures (Free 1993). The blow flies (Calliphoridae)are well known as visi tors to dung- and carrion-mimicking blooms. They often effect pollination, buttheir egg-laying act ivi t ies serve no reproduct ivepurpose for the flies (Proctor et al 1996). The maggotsd ie on the vege tab le subs t ra te , excep t in someexcept ional cases in Aristolochia f lowers . Manyspec ies o f roo t maggot f l i e s (Anthomyi idae :etymologically from the Greek for flower-fly) andtachinid flies (Tachinidae) feed on nectar of openbowl-shaped flowers. The special relations of seedparas i t i c Anthomyidae and Trol l ius europaeus(Ranunculaceae) involves mutual is t ic pol l inat ionrelationships (Pellmyr 1992). Among the flesh flies(Sarcophagidae) Blaesoxipha f letcheri may be animportant pollinator of pitcher plants (Sarraceniaspp.). Some Diptera, such as the Common Tiger Fly,Coenosia tigrina (Muscidae), and the Arctic Dung Fly,Scathophaga apicalis (Scathophagidae), hunt prey atflowers (Larson et al 2001).Pollination by flies seems to be particularly important in

Arctic and alpine areas (Kevan 1972; Levesqueand Burger 1982; Primack 1983; Pont 1993),South Africa (Vogel 1954; Goldblatt and Manning2000), and New Zealand (Primack 1978). In theArctic, for example, several species of Dipterafly between the flowers of adjacent plants and feedon nectar in such a way as to transfer pollen ofthe Arctic Aven, Dryas integrifolia. They assumetwo stances: 1) perched on the central styles and

stigmata while dipping between the styles and stamens sothat their dorsal surfaces collect pollen from the anthers,2) perched on the anthers and dipping for nectar so thattheir dorsal surfaces rub the stigmata (Kevan 1972).Even though there are many records of flies visitingflowers, there are many taxa that seem to avoid flowers.The robber flies (Asilidae), most stiletto flies (Therevidae)and long-legged flies (Dolichopodidae) serve as examples.That said, there is much to be learned about theimportance of flower visiting to flies, and about theimportance of that habit to plants and pollination.Al though there a re many records o f l a rge andconspicuous flies as flower visitors, there are few thatdemonstrate their importance as pollinators. In theblossoms that entrap and detain flies, the pollination

relationship (sapromyiophily) is clear (Proctor et al1996), but in most more colourful and showy flowers,pollination is less well documented. Another relativelyunstudied area of myiophily involves small flies, suchas Phoridae, Sciaridae, Mycetophilidae, Piophilidae,and so on (Larson et al 2001). As Carol Kearns (2001)so ably discusses , new discoveries about flower relationsand Diptera are waiting to be found within the broadaspects of biogeography, systematics, behaviour, andphysiology of this diverse and multi-faceted order.

FLIES AS VECTORS OF DISEASEDesmond H. Foley

Fever, grossly swollen limbs, diarrhoea, internal bleed-ing, brain damage … death! These are just some of thesymptoms millions of people endure due to fly-bornediseases. Malaria alone infects 300-500 million peopleeach year and causes an estimated annual loss of 35 mil-lion future life-years due to premature mortality and dis-ability (World Bank 1993).Since time immemorial flies have been the constant com-panions of humans and, due to the tendency of some spe-cies to feed on humans and pass on pathogens that causedisease, they have helped to shape human history. His-torically, such “vectors” or transmitters of disease ren-dered large areas of the world out-of-bounds to would-be conquerors, colonizers, and nation builders (Harrison1978, Bruce-Chwatt 1988). Human diseases caused byflies have their largest impact in tropical areas althoughthey were more widespread in the past (de Zuluetta 1994).Vector-borne disease requires an interaction between thefly, pathogen, host (e.g. humans), and the environment,and it may be complicated by reservoir hosts. For example,sylvan yellow fever circulates among forest monkeys inUganda through the mosquito Aedes africanus, but the vi-rus “spills over” to humans when monkeys invade Bananaplantations and are bitten by another mosquito Aedesbromeliae, which then feeds on humans (Haddow 1965).Non-biting flies can transmit fly-borne pathogensmechanically (e.g. “fi l th fl ies” spread intestinalpathogens causing typhoid [Graczyk et al 2001]), or bitingflies can transfer pathogens biologically where the mainnatural route for the pathogen is via the fly (e.g. malariaundergoes part of its life cycle in the fly). Only a limited

Table 6. Diptera of major

medical importancethat contain biological

vectors and the maindiseases they cause in

humans.

YLIMAF EMANNOMMOC)SUNEG( NEGOHTAP ESAESID

eadiciluC(seotiuqsoM selehponA )

seotiuqsoMseotiuqsoM

aozotorPsuriV

sedotameN

airalaMsitilahpecne/eugned/revefwolleY

sisairaliFcitahpmyL

eadidohcysP (seilfdnaS sumotobelhP ) aozotorP sisainamhsieL

eadiilumiS (seilfkcalB muilumiS ) edotameN sisaicrecohcnO

eadinabaT (seilfesroH sposyrhC ) edotameN sisaioL

eadinissolG (seilFestesT anissolG ) emosonapyrT sisaimosonapyrT

Figure 9a.Flower flies

(Syrphidae) are thebest known of the

flower feeding flies.They feed on bothnectar and pollen.

Many flower flyspecies mimic bees or

wasps. Spilomyiafusca (shown here) is

a mimic of thefamiliar Bald-faced

Hornet(Dolichovespula

maculata). Photo byH. Goulet, AAFC.

B I O D I V E R S I T Y 3 ( 4 ) 19

number of dipteran families contain species that requirea blood meal to develop their eggs. Mouthparts in thesefamilies are modified for piercing, blood sucking, andlapping, and vertebrate host location mechanisms arehighly developed (Gibson and Torr 1999).Table 6 illustrates the diversity of Diptera and patho-gens involved in the biological transmission of diseasesto humans. This list does not cover the many dipteranvectors of non-human diseases, for instance biting midges(Ceratopogonidae) that transmit bluetongue virus to sheep(Mellor et al 2000).Flies are both abundant and vagile, all of which add to thedifficulty in their control. Biting flies may be a local prob-lem or may arise far away; Ochlerotatus notoscriptus (Cu-licidae) will range over only a few hundred metres (Watsonet al 2000) but virus-laden biting midges 1-3 mm long canbe carried by the wind for over 700 kilometres (Mellor etal 2000). Biting flies inhabit undisturbed ecosystemsthrough to urban environments. The Yellow Fever or Den-gue Mosquito (Aedes aegypti) is now so dependent onhumans it usually completes its life cycle indoors, feedingon humans and breeding in water stored inside houses.Humans have often unwittingly assisted the spread ofvectors (Lounibos 2002). For example, the Asian TigerMosquito (Aedes albopictus) spread throughout the USAvia imported automobile tires that contained water(Moore 1999). A malaria mosquito Anopheles punctulatusextended its range in Papua New Guinea due to roadbuilding, which increased its preferred habitat of muddytemporary puddles (Ebsworth et al 2001).Humans have responded to vector-borne disease by wagingwar on flies, especially mosquitoes (Harrison 1978). Be-tween 1904 and 1914, discipline and an army of mosquitocontrol workers enabled General William Gorgas to controlyellow fever and malaria long enough to complete thePanama Canal (Powers and Cope 2000). Diseases like ma-laria often account for more casualties among the militarythan the fighting. For example, during WWI over one thirdof the 300,000 British and French and many of the Germantroops along the Macedonian front secumbed to malariathereby halting that campaign (Bruce-Chwatt 1988).The vulnerability of troops to diseases like malaria ex-plains the continued military interest in vector-borne dis-ease research (http://wrbu.si.edu/). In 1955, the WorldHealth Organization (WHO) launched a worldwide ef-fort to eradicate malaria armed with the new weaponschloroquine and DDT (Najera 1989). Although this at-tempt failed partly due to drug and insecticide resistance,the WHO continues vector-borne disease research (http://www.who.int/tdr/index.html).A militaristic approach to disease control has been aban-doned, but biting Diptera, especially mosquitoes, are stillviewed only as an enemy. However, only a minority ofthe over 3,000 species of mosquitoes are important pestsof humans. The rest include the bizarre and the beautiful– as well as the beneficial! Witness Malaya leei as it

steals food from the mouths ofCrematogaster ants (Miyagi 1981), theegg-guarding behaviour of the Green-eyed Mosquito Trichoprosopondigitalum (Lounibos 1991), the spec-tacular metallic blue Sabethes cyaneus(Foster and Taylor 1991), orToxorhynchites species whose adultsfeed on nectar and other plant exudatesbut whose larvae prey on larvae of lessbenign mosquitoes.

One legacy of this war on biting flies isthat research is skewed toward taxa ofmedical and veterinary importance.Closer inspection of mosquitoes and blackflies revealedsimilar-looking species that were previously overlooked.For example, using new techniques, the number of spe-cies recognized within the Anopheles punctulatus groupwas almost doubled (Foley et al 1993). At least 90 of the458 species of Anopheles listed by Harbach (1994) arepart of species complexes. These so-called “sibling” spe-cies display differences in biting behaviour and distribu-tion that helps explain why the apparent presence of vec-tors does not always match the level of disease transmis-sion. For example, the Anopheles farauti complex in theSolomon Islands (Figure 10) was found to include non-human biting species (Foley et al 1994), whose presencemasks the importance of the vector species and can leadto wasted effort in controlling mosquito larvae.

A trade-off often exists between vector control and theenvironment. Some see a worldwide ban on the use ofDDT as a win for the environment but others argue thatthis robs poor countries of an affordable and effective in-secticide for malaria control (Curtis and Lines 2000).Worldwide release of exotic fish species like theMosquitofish, Gambusia affinis, to control mosquito lar-vae (Rupp 1996), use of pesticides like DDT, or clearingof riparian vegetation for mosquito control has not alwaysbeen evaluated for efficacy or weighed against possibleadverse effects on native fish species or the environment.

Lately there has been renewed interest in environmentalmanagement (Utzinger et al 2001) and environmentallyfriendly and appropriate technology for vector control. Forinstance, dengue has been controlled in parts of Vietnamby applying mosquito-larvicidal copepods to water tanksand limiting breeding habitats by encouraging a recyclingindustry based on plastic containers (Vu et al 1998). Flytraps to reduce trachoma have been designed from recycledplastic bottles (Dobson 2000), and good control of tsetseflies has been achieved using traps (Jordan 1995).Biopesticides such as Bacillus thuringiensis israelensis,environmental management plans (http://www.lgaq.asn.au/),and drainage techniques, such as runnelling (Dale et al1993), are now used for mosquito control.

Increasing international movement and developmentssuch as dams, roads, logging, and irrigation, have opened

Figure 10.A bloodfed Anophelesfarauti mosquito, theprincipal coastalmalaria vector in theSW Pacific. (Photo byDes Foley, TropicalHealth Program,University ofQueensland).

ABOUT THE AUTHORDes Foley has anabiding interest in allthings culicid. Usingbiochemical andmolecular techniques,he has helped tounravel the speciescomposition andphylogeneticrelationships of theprincipal anophelinedisease vectors of theSW Pacific. Des hasalso published onanopheline ecology andbehaviour, especiallyas related to diseasetransmission. Inaddition, he hasauthored a series ofcomputer-driven keysfor morphologicalidentification ofAustralasianmosquitoes. You canreach Des at TropicalHealth Program andDepartment of Zoologyand Entomology,University ofQueensland, Brisbane,Queensland, 4072Australia;D.Foley@sph.uq.edu.a

T R O P I C A L C O N S E R V A N C Y20

up new areas and opportunities for human/vector con-tact (Gratz 1999). Anthropogenic changes in climate may(Epstein 2001) or may not (Reiter 2001) increase thiscontact, and resistance is developing to the drugs andinsecticides used to control vector-borne diseases(Hemingway and Ranson 2000). In the past, reliance onnew insecticides and drugs heralded a decline in basicresearch on vectors. As genetically engineered, patho-gen-resistant vector species develop into the next pos-sible “magic bullet” in the war on vector-borne disease(Ito et al 2002), will the research on medically impor-tant Diptera again suffer “collateral” damage?

WHEN BEING A MAGGOTIS A GOOD THING:

THE ROLE OF DIPTERAIN FORENSIC SCIENCE

Andrew McDowell

Blow flies and their maggots are generally considered tobe nuisance animals. They aren’t the prettiest insects to lookat and they tend to loiter around filth such as garbage androtting material. However, it is this very behaviour that makesblow flies potentially useful in forensic science.Forensic entomology is a general term for the applicationof the study of insects and other arthropods to legal is-sues, especially in courts of law (Catts and Goff 1992). Itcan be further divided into three broad categories – storedproduct entomology, urban entomology, and medico-crimi-nal or medico-legal entomology. The first two of thesecategories deal primarily with civil legal action, such aslitigation over issues like arthropod contamination of com-mercial products, while the last category deals with in-sects associated with criminal activities, such as murder,suicide, physical abuse, and neglect. For the remainder ofthis review the term forensic entomology will refer onlyto medico-criminal entomology.The use of arthropods in forensic science is not a newconcept. The thirteenth century writings of the Chinesemagistrate Sung Tzu detailed the identification of a mur-derer based on entomological evidence (McKnight trans-lated 1981), and 600 years later, during the nineteenthcentury, blow flies and cast puparia were used as indica-tors of post-mortem interval (Bergeret 1855; Megnin1894). However, until the 1980s, the use of entomologi-cal evidence in modern forensic science was largely ig-nored. Various case studies document how insects, pri-marily blow flies, have been used to estimate post-mortem intervals (PMI) (Goff 1991; Goff 1992; Lord etal 1992; Lord et al 1994; Benecke 1998; Introna et al1998). Insects have also proven useful in demonstratingthat corpses have been moved (Benecke 1998) or inter-fered with (Anderson 1997) and in showing that toxinswere present in a corpse when there was no other appar-ent cause of death (Beyer et al 1980; Introna et al 1990;Miller et al 1994). Entomological evidence has also beenused to link suspects to a scene of crime (Prichard et al1986). Not all forensic cases involve blow flies – the

advanced developmental stage of black fly pupae on asubmerged car (Merritt 1994) and the presence of bumblebee hairs on stolen banknotes (Howden 1964) both re-sulted in murder convictions.Insects can be particularly useful in forensic sciencebecause they invade carrion in a predictable pattern overtime (Swift et al 1979; Goff 1991; Shean et al 1993;Richards and Goff 1997), with blow flies (Calliphoridae)(Figure 11) and flesh flies (Sarcophagidae) typically be-ing the first to arrive. Other groups of flies (e.g. muscidflies [Muscidae] and scuttle flies [Phoridae]) and beetles(e.g. carrion beetles [Silphidae], hister beetles[Histeridae], and rove beetles [Staphylinidae]) are alsoattracted to decomposing corpses and arrive at differenttimes depending on factors such as state of decay andseasonal conditions. By examining species successionpatterns and determining the age of maggots collectedfrom a corpse, it is possible to make an accurate estima-tion of time passed since death. Blow flies are particu-larly useful for estimating post-mortem interval becausethey are usually the most abundant arthropods found inassociation with decaying flesh. Forensic entomologistscollect flies, eggs, maggots, and pupae from crime scenesand return them to the laboratory for identification. Adultflies are usually killed and preserved for identification,but because immature stages are often difficult to iden-tify to the species level, some are reared to adulthoodfor confirmation of species identity. Others are preservedfor age determination and tentative species identifica-tion. Estimates of post-mortem interval based on ento-mological evidence also rely on an accurate interpreta-tion of environmental conditions at the crime scene. Fac-tors including ambient and corpse temperatures, humid-ity, and the position of the corpse may affect the devel-opment rate of maggots and thus estimations of their age(Catts 1992). Other factors associated with estimates oftime of death include presence or absence of narcoticsin the corpse, and also whether the corpse is wrapped inmaterial (e.g. blankets) (Catts and Goff 1992) or fullyclothed or naked (Mann et al 1990).To be able to accurately estimate times of death, forensic en-tomologists must be familiar with both the species present inthe area in which a corpse is found and the biology of carrion-breeding insects. Succession patterns and overall decomposi-tion times have been shown to vary with location, seasonalconditions, and habitat type (Early and Goff 1986; Tullis andGoff 1987; Goff 1991; Shean et al 1993; Tantawi et al 1996;Richards and Goff 1997; Avila and Goff 1998; Davis and Goff2000). Thus, to become familiar with local species, forensicentomologists conduct field trials under various conditions.Typically such experiments are conducted using pig carcassesas models for humans since pigs are similar to humans intheir physiology, gut fauna, and hair coverage. Most researchin the area of forensic entomology has been conducted in NorthAmerica and Europe, but more recently forensic entomologyhas become an area of increasing focus and research is nowbeing conducted throughout South America, Southeast Asia,and Australia. Australia is of particular significance given its

ABOUT THE AUTHOR Andrew McDowell

is a Doctor ofPhilosophy student at

the University ofQueensland,

Australia. With abackground in insectecology, he has beeninvestigating the use

of blow flies inforensic science forthe past three years

and specifically, howblow fly successionpatterns vary with

habitat and season.You can reach Andrew

at Department ofZoology and

Entomology, School ofLife Sciences,University ofQueensland,

Brisbane, QLD 4072,Australia;

email: a.mcdowell@mailbox.uq.edu.au

B I O D I V E R S I T Y 3 ( 4 ) 21

great diversity of carrion-breeding blow flies from two fo-rensically significant genera, Chrysomya (Figure 11) and Cal-liphora. In Australia, these two genera may co-exist on car-rion, but dominate the carrion system at different times of theyear. For example, in sub-tropical southeast Queensland, Chry-somya is more abundant during warmer months from Sep-tember to May while Calliphora dominates the cooler monthsof June through to August (McDowell, unpublished data).

The applications of insects, particularly Diptera, in foren-sic science are many and varied. However, in order to useentomological evidence efficiently, a thorough understand-ing of the ecology of carrion systems in a given area isrequired. With the recent resurgence of interest in the useof arthropods as a form of evidence in criminal situations,and with new and innovative research being conducted inthis field, the use of entomological evidence is becomingmore widely accepted throughout the forensic community.

MOLECULAR SYSTEMATICSOF FLIES (DIPTERA)

Shaun L. Winterton

Genetic and molecular methodologies are now widelyused in the study of dipteran biology, ecology, specia-tion, biogeography, population genetics, systematics, anddevelopmental biology. To justly treat the many facetsof fly research using molecular techniques is beyond thisbrief summary (e.g. black fly systematics based on poly-tene chromosomes). In this section I will highlight re-search into the evolutionary relationships of Diptera thathas used molecular sequencing methods.

The development of the polymerase chain reaction (PCR)and automated sequencers to respectively amplify andsequence selected gene regions has greatly eased mo-lecular research. As a result, PCR use has become com-mon in modern phylogenetic analysis, and to create anevolutionary tree, modern systematics now relies not onlyon traditional comparative morphological study but alsoon a range of molecular data sources.

THE CONTRIBUTION BY DROSOPHILAThroughout most of the twentieth century the use of a smallfly of the family Drosophilidae has formed the basis forresearch into eukaryote genetics (genetics of animalswhose cells contain nuclei). In 1910, Nobel Laureate T.H.Morgan chose the Vinegar Fly (Drosophila melanogaster)

(Figure 12) for his studies ofheredity, and with the

help of A.H.Sturtevant

and C.B.B r i d -ges, heformu-l a t e d

the theory ofc h r o m o s o m a lheredity. Since

then, Drosophila has been used to construct the first ge-netic maps and formulate much of our understanding ofsex determination, genetic linkage, and chromosomal me-chanics and behaviour (Kornberg and Krasnow 2000).In early 2000, researchers announced that the sequenc-ing of the Drosophila melanogaster genome was com-plete (Adams et al 2000)—a significant advancement ineukaryotic genomics. The Drosophila genome comprisesapproximately 120 megabases and encodes around 13,600genes. Although previously well over 2,500 genes hadbeen defined genetically, now the positions of all theDrosophila genes can be mapped, enabling researchersto investigate the functions of genes not linked with par-ticular phenotypes (appearances). Through conservationof biological processes, studies of genes in Drosophila(and other organisms for which genomes are sequenced)will also increase our understanding of comparable genesin humans.

Recently, the completion of genome sequencing of themosquito Anopheles gambiae (Culicidae) has given theresearch community a second dipteran genome (http://www.ensembl.org/Anopheles_gambiae/). The results ofthis research are due to be published in the journal Sci-ence in the second half of 2002.

PHYLOGENETIC STUDIES OF DIPTERAAs a whole, our knowledge of dipteran evolutionary rela-tionships is relatively patchy. Phylogenetic studies based onmorphology alone have had their difficulties; despite sig-nificant consensus, there is still strong debate, particularlywith respect to higher-level relationships (reviewed by Yeatesand Wiegmann 1999). A wealth of new data available fromDNA sequences is beginning to be exploited at various taxo-nomic levels from interfamily level relationships to popula-tion genetics. Rates of mutational change and the propor-tions of variable versus conserved regions vary consider-ably across different genes. Variable rates of mutation meansthat individual genes may be suitable only for particular taxo-nomic levels within a given group, and may not be appro-priate for the same level in another group. In general, geneswith slower rates of mutation change are more informativefor higher-level relationships, while genes with high ratesof mutation change are informative for exploring lower levelrelationships such as population studies. This makes geneselection for any phylogenetic study highly idiosyncratic,and the appropriateness of a particular gene for studying aspecific study group can only be determined with certaintythrough preliminary studies. Various genes have been ex-plored for their phylogenetic utility, although compared withvertebrate systematics we still have relatively few genes foruse in insect systematics. There are two sources of sequencedata in the animal cells:Mitochondrion. Individual mitochondria in the eukary-

otic cell each contain a circular, maternally inheritedDNA genome betraying their prokaryotic prehistory.Genes commonly used for examining phylogenetic re-lationships in Diptera include the ribosomal DNA genes12S and 16S, transfer DNA genes and protein encoding

Figure 12.Throughout most ofthe twentieth centurythe use of a small fly(Drosophilamelanogaster) of thefamily Drosophilidaehas formed the basisfor research intoeukaryote genetics(genetics of animalswhose cells containnuclei). This is afamiliar, cosmopoli-tan species that iscommonly encoun-tered around rottingfruit. (Drawingcourtesy of AAFC[from McAlpine et al1987, p. 1011]).

Figure 11.(opposite page)Insects canbe particularly usefulin forensic sciencebecause they invadecarrion in apredictablepattern over time.The blow fly(Calliphoridae)Chrysomya sulcifronsis a typical secondaryinvading speciesin south-eastQueensland; theirlarge hairy larvae aremost common in thewarmer months.(Photo by AnthonyO’Toole, Universityof Queensland)).

T R O P I C A L C O N S E R V A N C Y22

genes such as Cytochrome oxidases (COI, II and III),Cytochrome b and ND1, 2, 3, 5 and 6 (Simon et al 1994).

Nucleus. Nuclear ribosomal DNA (e.g. 5.8S, 18S and 28S)has been the mainstay of many phylogenetic studies ofDiptera owing to their numerous repeats in the genomeand relative ease of amplification and sequencing. As withmitochondrial ribosomal genes, there are regions of con-served and variable regions corresponding to secondarystructure stems and loops. Conserved regions have beenuseful at higher taxonomic levels, while variable regionshave been plagued by difficulties of alignment. Gene ex-ploration and increasing efficiency in amplification andsequencing techniques have led to increased use of nuclearprotein-encoding genes in studies of fly evolution. Manyare single copy genes, but others may form part of multi-gene families that, if not fully concerted, may result insimultaneous amplification of multiple, closely related,but slightly different genes. Other sequencing and align-ment difficulties with using nuclear protein encoding genesmay come from the presence of introns (extra non-cod-ing DNA) in the gene’s sequence in certain taxa, the pres-ence of pseudogenes (non-functional copies of genes) orallelic polymorphisms (different forms of the gene at thesame locus).

The number of genes being utilized for molecular system-atics is increasing and detailed reviews of the use of nucleargenes in insect systematics include Friedlander et al (1992),Brower and DeSalle (1994), and Caterino et al (2000).

The description of the evolutionary relationships of Dipterahas been based largely on morphology over the last cen-tury, and relatively few studies have been undertaken us-ing molecular data. Table 7 presents an overview of theimportant contributions to molecular phylogenetics of flies.Obvious from the examples of higher phylogenies (e.g.above the family level) is the almost exclusive use of alimited number of mitochondrial and nuclear rDNA genesequences. Considering the increasing number of nuclearprotein encoding genes that have been shown to be usefulfor recovering higher-level relationships in other insectgroups (Friedlander et al 1992), their usefulness in higher-level dipteran systematics is clearly worthy of investiga-tion. Contrary to this, studies on lower level relationships(e.g. families, subfamilies) of Diptera have used a farbroader range of genes to investigate the internal hierar-chy of relationships.

With the advent of PCR and the subsequent ease of se-quencing using automated PCR and sequencing machines,the use of large amounts of sequence data is becomingmore and more common. Frequently, multiple genes aresequenced and the level of congruence is determined be-tween individual genes, gene regions, or with morpho-logical data and other non-molecular sources of data.Combining multiple data sets from a variety of molecu-lar and non-molecular sources to generate robust totalevidence phylogenies appears to be the future directionof systematic research on Diptera.

PUORGCIMONOXAT STNEMMOCDNADESUSENEG SROHTUAHIGHER LEVEL PHYLOGENIES (RELATIONSHIPS ABOVE THE FAMILY LEVEL)

)seilfrehgih(arecyhcarBsiydutssiht;enegS82etelpmocehtdesuyehTrebmunregralhcumaesulliwdnassergorpni

.krowsuoiverpynanahtsralpmexefonoitacinummoclanosrep,latennamgeiW

)seilimafdetalerdnaseilfwolb(eatartpylaC ANDrS61dnaS81 1002latealamriN

)sevitalerriehtdnaseotiuqsom(ahpromociluC ANDrS8.5dnaS81 7991laterelliM

)sevitalerriehtdnaseotiuqsom(ahpromociluC ANDrS82 6991lateikswolwaP

)seilf)narecyhcarBrehgih(ahpahrrolcyC ANDrS82 9891namdierFdnakcnirbssoV

)seilfetartpylac(aediocsuM IIOCdnaIOC 0002lateinocsanreB

)seilfrewolfdnaseilfdedaeh-gib(aediohpryS ANDrS61dnaS21 0002setaeYdnanotgnivekS

)seilimafdetalerdnaseilfesroh(ahpromonabaT ANDrS82 0002latennamgeiW

LOWER LEVEL RELATIONSHIPS (FAMILIES AND BELOW)

)seilfdedaeh-llams(eadirecorcA sucolyratnemidurdnaS82,S61 ataddehsilbupnu,notretniW

)seilfottelitsfoylimafbus(eanityhpopagA ahpla-1fEdnaS82 1002latenotretniW

)seotiuqsomfoylimafbus(eanilehponA G,S82,5DN 6 etihwdna,dp 1002lateiksniwyzrK

)seotiuqsom(eadiciluC deyeetihW 7991yehaFdnayksnaseB

)seilfdeye-klats(eadispoiD sselgniw,etihw,ahpla-1fE,S61,S21,IIOC 1002laterekaB

)seilftiurfllams(eadilihposorD S82,hdA,tnemgarfANDtm dnasikadnaleP;5991lateossuR;2991ellaSeD3991cangiloS

)seilfsadyM(eadidyM ANDrS82 1002nnamgeiWdnaniwrI

)seilfkcalb(eadiilumiS IIOC 0002latesseurP

)seilfkcalb(eadiilumiS ahpla-1fEdnaKCPEP,CDD,S82,2DN,S21 0002notluoM

)seilftiurf(eaditirhpeT ANDrS61 0002naH

)seilfottelits(eadiverehT ahpla-1fEdnaS82 0002lategnaY

SIGNIFICANT PHYLOGENIES OF SPECIES AND SPECIES-GROUP RELATIONSHIPSselehponA )seotiuqsom( IIOC 8991lateyeloF

arecortcaB )seilftiurf( ANRt,S61 lav 1002arahakaNdnaijaruM

sumonorihC )segdim( b2bg,061pss 0002latehciverakaM

alihposorD )eadilihposorD(arenegdetalerdna dna,II&IOC,1-STI,hdpG,doSnZ,uC,hdAstnemgarfenegrehtosuoremun

ellaSeDdnarekaB;9991alayAdnaikswotaiwK8991lateydarG'O;7991

azymotyhP )seilfrenim-fael( II&IOC 0002nnamgeiWdnareffehcS

sitelogahR )seilftiurf( S61 7991naHdnanorehPcM

agahpohtacS )seilfgnud( btyC,IOC 1002lateinocsanreB

ABOUT THE AUTHORShaun Winterton is a

postdoctoral fellow inthe Molecular

Systematics Laboratoryat North Carolina StateUniversity. His currentresearch is focused on

the systematics and co-evolution of small-

headed flies(Acroceridae) and theirspider hosts. He is also

interested in thesystematics of nerve-

winged insects(Neuroptera) and has

published papers onthese as well as on

stiletto flies(Therevidae) and on

biological control. Youcan reach Shaun at

Molecular SystematicsLaboratory,

Department ofEntomology

North Carolina StateUniversity, Box 7613,

Raleigh,NC 27695-7613 USA;

email: WintertonShaun@netscape.net

B I O D I V E R S I T Y 3 ( 4 ) 23

DIPTERAN GLOW-WORMS:MARVELLOUS MAGGOTS

WEAVE MAGIC FOR TOURISTSClaire H. Baker

Insects and tourism are not two words commonly or posi-tively put together. In fact, quite often it is insects likemosquitoes (Culicidae), biting midges (Ceratopogonidae)and horse flies (Tabanidae) that make some tourist at-tractions less desirable places to visit. However, aroundthe globe, for an increasing number of ecotourists, a fewspecific insect species have become the main attraction.Butterflies (e.g. Monarch Butterfly reserves in Mexico)and fireflies (e.g. in Great Smoky National Park, USA)spring to mind, but one of the biggest insect drawingcards belongs to the Diptera. A number of species withinKeroplatidae (Orfelia fultoni, Arachnocampa spp., andKeroplatus spp.) emit a bioluminescent blue/green lightas larvae. In some instances these “glow-worms” con-gregate in large numbers and form impressive displays,similar to a star-lit night sky.

The genus name, Arachnocampa, or “spider-like grub,” re-fers to the larval characteristics of glow-worms. The larvaeof all glow-worms construct mucus tubes from which theyhang a snare or web of silk and mucus to capture prey thatis attracted by its bioluminescence (Richards 1960). Thelarvae are voracious predators and will feed on many typesof arthropods attracted to their glow (Broadley 1998, Baker1999). Larvae are generally long-lived while the non-biolu-minescent adults are very short-lived, dying within a fewdays of emergence (Richards 1960, Baker 1999).Glow-worms are found in a variety of habitats – such ascaves, mine tunnels, railway tunnels, rainforest banks, anddamp overhangs – but always within areas of very high hu-midity. Larvae will glow during daytime hours if subjectedto constant darkness (Baker 1999) and therefore, touristscan observe cave populations that glow both day and night.

NEW ZEALANDThe renowned Waitomo caves on the North Island of NewZealand have been a popular tourist attraction since firstbeing discovered in 1887 (http://www.new-zealand.com/waitomocaves/). The endemic New Zealand glow-worm,Arachnocampa luminosa, attracts approximately 300,000tourists annually (Peter Dimond, Manager Waitomo Mu-seum of Caves, personal communication). The glow-wormsin the Te Anau caves of the South Island also attract largenumbers of tourists. Due to the importance of glow-wormsto the New Zealand tourist industry, a number of biologicaland ecological studies have focused on this species (fromNorris [1894] through to the most recent, Broadley [1998]).

AUSTRALIAIn Australia, at Natural Bridge in Springbrook NationalPark, part of the World Heritage-listed Central EasternRainforest Reserves, another large colony of glow-wormsis the subject of increasing tourism interest, with an esti-mated 110,000 night visitors per annum (Mike Hall,

Queensland National Parks Ranger, personal communica-tion). A number of smaller glow-worm tourism operatorshave tours in other rainforest areas of eastern Australia,with glow-worm displays in parts of Tasmania rivalingthose of the New Zealand caves (personal observation).Three endemic species are currently known from Austra-lia (Harrison 1966): A. flava, endemic to rainforest areasof southeast Queensland (Perkins 1935; Harrison 1966);A. richardsae identified from the Newnes railway tunnel,New South Wales; and A. tasmaniensis from Tasmania. Ihave also identified glow-worm species, which I will soonbe describing from isolated caves and rainforest pocketsin parts of Australia.OTHER LOCALESOther bioluminescent keroplatid genera exist in NorthAmerica, Japan, and Europe. For example, tourists canview the “Dismalites” (Orfelia fultoni) in The DismalsCanyon of Alabama (USA); the canyon itself is a spec-tacular backdrop for this amazing insect.OBSTACLES AND OPPORTUNITIESAs the enthusiasm for ecotourism continues to surge, in-sects promise to play an ever-growing role. Most “mainattraction” species are localized, and their habitats and foodresources are often precariously balanced. More researchis required to better understand the requirements of theseinsects and the careful management that some species mayrequire to avoid exploitation. Scientists need to fuel gov-ernment policy makers and business owners with solid in-formation that will ensure the right decisions are made.Despite the concerns, ecotourism offers the promise of con-servation. More people travelling to more wild places willdrive governments and local landowners in the direction ofland preservation, which is so vital to conservation. In turn,by showcasing one of the most diverse groups of animalson the planet, ecotourism operators can generate new inter-est and appreciation for the often- berated “lowly” fly.

ACKNOWLEDGMENTSWe thank Dan Bickel, Jeff Cumming and DavidMcAlpine for reviewing the articles in this series.

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ABOUT THE AUTHORClaire Baker isstudying for a PhD atthe University ofQueensland inAustralia. Her interestsare in insect taxonomyand phylogenetics,molecular biology, andtourism management,and she is currentlyworking on Australianglow-worms (Diptera:Keroplatidae). You canreach Claire atDepartment of Zoologyand Entomology,School of Life Sciences,University ofQueensland, BrisbaneQLD 4072, Australia;email: c.baker1@mailbox.uq.edu.au

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T R O P I C A L C O N S E R V A N C Y44

A N N O U N C E M E N T S

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FOURTH MINISTERIAL CONFERENCEON THE PROTECTION OF FORESTSIN EUROPE28 – 30 April 2003. Vienna, Austria. Contact: MinisterialConference on the Protection of Forests in Europe, LiaisonUnit Vienna. Phone: +43 1 710 77 02; Fax: +43 1 710 02 13;E-Mail: liaison.unit@lu-vienna.at; Web: http://www.-mcpfe.org/

Fly Heads.The right picture

illustrates the head of atypical fly, in this case

the flower fly,Spilomyia fusca

(Syrphidae) [NorthAmerica]. The left

picture is of a tsetsefly, Glossina sp.

(Glossinidae)[Africa].Note the difference in

mouthparts betweenthese two flies.

Spilomyia adults feedon nectar and pollen of

flowers while adulttsetse flies are blood

feeders and carryhuman diseases such as

trypanosomiasis(sleeping sickness).

actual size

actual size