9 The Panorpoid Orders 1 . Intr oduc t ion I n this and the followin g chapter we shall deal with the endopter yg ote insects—those tha t h ave a distinct pupal instar in which the insect under g oes a drastic metamorphosis fro m t he larval to the adult form. As noted in Chapter 2, Section 3.2, considerable difficult y h as ar i sen i n d ec idi ng w h et h er t h een d opterygote or d ers h ave a common or i g i norare po l yp h y l et i c. T h e fi ve or d ers cons id ere di nt hi sc h apter, Mecoptera, D i ptera, S i p h onaptera , T r i c h optera, an d Lep id optera, s h ow c l ear a ffi n i t i es t h at ena bl et h em to b e g roupe d to g et h e r as the p anor p oid com p lex. Within the com p lex there are two sister lines of evolution: th e Antlio p hora (first three orders) and the Am p hiesmeno p tera (Tricho p tera and Le p ido p tera). Th e rema i n i ng or d ers, Mega l optera, Rap hidi optera, Neuroptera, Co l eoptera, Hymenoptera , an d Streps i ptera, d ea l tw i t hi nC h apter 10, s h ow f ew a ffi n i t i es w i t h t h e panorpo id g roup . 2 . Mecoptera S ynonyms: P anor p atae, Panor p ina, Panor p id a C ommon name: s cor p ionflies S lender medium-sized insects; head usually prolonged ventrally into a broad rostrum with long filif orm antennae, we ll - d eve l ope d compoun d eyes, an dbi t i ng mout h parts; usua ll yw i t h two pa i rs o f identical membranous win g s with primitive venation and carried horizontall y at rest; abdomen w ith short cerci and, in males, prominent genitalia. L arvae usually eruciform with simple eyes, biting mouthparts, and thoracic legs; abdominal legs present or a b sent. Pupae d ect i cous an d exarate . This is a small order containin g about 500 known species, about 90% of which belon g t o t wo families, Panorpidae and Bittacidae. The order is particularly common in the Northern Hemis p here and includes about 7 5 North American and 4 British s p ecies. About 30 s p ecies occur i n Austra li a. 23 9 240 CHAPTER 9 Structur e Adult . A ch aracter i st i c f eature o f most Mecoptera i st h e ventra l pro l ongat i on o f t he head into a broad rostrum. Incorporated into this structure are the cl y peus, labrum, and m axillae. Compound e y es are well developed, and in most species there are three ocelli. The antennae are multise g mented and filiform. The mouthparts are mandibulate, except in N annoc h orista,w h ere t h ey are spec i a li ze d an d may b e i nterprete d as f ores h a d ow i ng t h e s uctor i a l type seen i n l ower D i ptera. T h e prot h orax i s sma ll ,t h e pterot h orax we ll d eve l ope d. T h e l e g s are l on g an d t hi nan d a d apte df or wa lki n g .T h e yh ave a fi ve-se g mente d tarsus. In Bittacidae the fifth tarsal se g ment folds back on the fourth and is used for catchin g pre y. Two pairs of full y developed, identical, membranous win g s are present in most species; the v enat i on i spr i m i t i ve. In Bore id ae t h ew i ngs o ff ema l es are sma ll sc l erot i ze d pa d sw hil e t h ose o f ma l es are h oo klik ean d use d to grasp t h e f ema l e d ur i ng mat i ng. W i ngs are re d uce d i n some f ema l e Panorpo did ae an d B i ttac id ae, an d a b sent i n f ema l e Apteropanorp id ae. T h e abdomen of females is 11-se g mented and usuall y carries 2-se g mented cerci (unse g mented i n Bittacidae and Boreidae). In female Boreidae the 10th ter g um is prolon g ed and to g ethe r wi t h t h epo i nte d ,sc l erot i ze d cerc if orms a f unct i ona l ov i pos i tor.Inma l es segment 9 i s bif urcate an db ears a pa i ro fb u lb ous c l aspers. Segment 10 i s i nconsp i cuous an db ears unse g mente d cerc i .T h eae d ea g us li es at t h e b ase o f t h ec l aspers. In Panorp id ae t h e term i na l s e g ments are turned upward and resemble somewhat a scorpion’s stin g , hence the common n ame for the order . T he foregut has two interesting features. The esophagus contains two dilations that appear to f orm a suc ki ng apparatus, an d t h e crop i s prov id e d w i t hl ong setae (acant h ae ) t h at may act as a fil ter. S i xMa l p i g hi an tu b u l es occur. T h e nervous system i s genera li ze d , w ith three thoracic and between five and ei g ht abdominal g an g lia (males usuall y with on e m ore than females). Each testis com p rises three or four follicles. The p aired vasa defer- e ntia open separately into a median seminal vesicle, which also receives paired accessor y gl an d s. In f ema l es eac h ovary conta i ns 7–19 po l ytrop hi covar i o l es (pano i st i covar i o l es in Nannoc h or i st id ae an d Bore id ae). T h epa i re d ov id ucts un i te b e f ore enter i ng a gen i ta l pouc h . T h e d ucts f rom t h e spermat h eca an d accessor ygl an d sa l so l ea di nto t h e pouc h. L arva an dP upa . Larvae are typically caterpillarlike, with a distinct head capsule tha t b ears s i mp l e eyes. Pro l egs occur on t h e fi rst e i g h ta bd om i na l segments, an d t h e apex o f t h e a bd omen b ears e i t h er a suct i on di sc or a pa i ro fh oo k s. In Bore id ae an d Panorpo did ae l arvae are g rublike, lackin g prole g s and a terminal suction disc. Larvae of Nannochoristidae are v er y elon g ate, lack prole g s, but have a pair of apical hooks. Pupae are decticous and exarate . Lif eH i story and Hab i t s Adult scorpionflies are most frequentl y encountered in cool, shaded locations, espe- c ially among low vegetation, though a few species occur in semidesert habitats. They ca n fl y act i ve l yw h en di stur b e d ,t h oug h t h ey norma ll y rest on grass, un d er l eaves, etc. A d u l t P anorp id ae f ee d most l yon d ea d so f t- b o di e d art h ropo d s( i nc l u di ng i nsects caug h t i nsp i - ders’ webs); the y also eat nectar, pollen, and fruit j uices. Bittacidae, b y contrast, are insect p redators, catchin g their pre y either in fli g ht or b y han g in g under ve g etation till it comes within range. In members of both of these families much of the food of females is pro - vid e di nt h e f orm o f a nupt i a l g if t b yama l e d ur i ng courts hi p (see C h apter 19, Sect i on 4.2) . T h e f oo di tem may b eanart h ropo d recent l yo b ta i ne db yt h ema l e or a mass o f sa li va s ecreted from the male’s g reatl y enlar g ed salivar yg lands. Adult Boreidae feed on mosses, 241 T HE P A N O RP O ID ORDER S and members of other families ma y be herbivorous, sapropha g ous, or carrion feeders. To attract females, male panorpids and bittacids secrete pheromones from g lands on the pos- t erior abdominal se g ments. Visual si g nals (win g movements and abdominal vibrations) are a l so i mportant i nc l ose-range courts hi p i nteract i ons i nt h ese an d ot h er (non-p h eromone- pro d uc i ng) mecopteran f am ili es. Eggs o fbi ttac id s are d roppe d ran d om l y; t h ose o f panorp ids an d c h or i st id s are l a id i n b atc h es i nmo i st d epress i ons i nt h e g roun d . Bore id ae d epos i te gg s sin g l y or in small batches in soil ad j acent to moss rhizoids. The e gg sta g ema y last from as little as a week up to several months in species where there is an e gg diapause. Larvae ar e saprop h agous, carn i vorous, or moss- f ee d ers an d pass t h roug hf our i nstars b e f ore enter i ng aqu i escent prepupa l p h ase, usua ll y i n an eart h en ce ll .T h e l engt h o f t h e prepupa l p h as e is varied and ma y include a diapause. The pupal stadium usuall y lasts 14– 5 0da y s. Mos t species are univoltine; some are bivoltine; and boreids take 2 y ears to complete a g eneration. Phylogeny and Classification Ar i c h array o f mecoptera lik e f oss il s i s k nown f rom t h e Lower Perm i an, an d t h emo d- ern consensus is that this includes a few g enuine Mecoptera. B y the Upper Permian, and extendin g throu g h the Jurassic, the order was abundant and diverse. Some Upper Permian scorpionflies from deposits in Australia and Siberia are assi g nable to the extant famil y Nan - noc h or i st id ae, w hil ere p resentat i ves o f some ot h er mo d ern f am ili es a pp ear i nt h eLowe r Jurass i c. Recent Mecoptera are arrange db yW ill mann (1987) i ntwosu b or d ers, conta i n i ng n i ne f am ili es w h ose poss ibl eevo l ut i onar y re l at i ons hi ps are s h own i nF ig ure 9.1 . F I GU RE 9.1. Proposed phylogeny of the extant Mecoptera. [After R. Willmann, 1987, The phylogenetic syste m o f t h e Mecoptera , S yst. Entomo l. 12 :519–524. By permission of Blackwell Scientific Publications Ltd.] 242 CHAPTER 9 Suborder Nannomeco p tera Thi ssu b or d er conta i ns t h es i ng l e, sma ll ,pr i m i t i ve Sout h ern Hem i sp h ere f am ily NANNOCHORISTIDAE. The ei g ht species in this g roup differ from other mecopterans in l ackin g a pump for transferrin g sperm from male to female, and in details of their external g enitalia, win g venation, and mandibular structure. Adults are small and live in the vicinit y of streams or l a k es. T h e i r l arvae are aquat i can d carn i vorous, f ee di ng on l arva l D i ptera, e spec i a ll yc hi ronom id s. Su b or d er Pisti ll i f era T he suborder Pistillifera (literally “piston-bearers” in reference to the sperm pump o f ma l es) i s di v id e db yW ill mann (1987) i nto two i n f raor d ers, Rapt i pe di aan d O pisthogonopora. The Raptipedia includes the large (14 5 species) and cosmopolitan fam - i l y BITTACIDAE (Fi g ure 9.2C) whose members have raptorial tarsi with which the yg ras p their pre y , often han g in g under ve g etation b y their forele g s. Amon g the Opistho g ono- pora, the BOREIDAE (Figure 9.2A) constitute the most distinct family; indeed, Hinto n ( 19 5 8) placed this holarctic group of about 2 5 species in a separate order, Neomecoptera . F I GU RE 9.2. Meco p tera. (A) B oreus b ruma l is ( Bore id ae ) ; ( B ) Panorpa h e l en a (Panor pid ae); an d (C ) B i ttacus p ilicornis ( Bittacidae). [A, B, from D. J. Borror, D. M. Delong, and C.A. Triplehorn, 1976 , An Intro d uction to th e S tu d y of Insects , 4t h e d . By perm i ss i on o f Broo k s/Co l e, a di v i s i on o f T h omson Learn i ng. C, f rom D. W. We bb ,N. D . Penn y , and J. C. Marlin, 197 5 . The Mecoptera, or scorpionflies, of Illinois . B u ll .I ll . Nat. Hist. S ur v. 31 :2 5 1–316. By permission of the Illinois Natural History Survey.] 243 T HE P A N O RP O ID ORDER S B oreidss are small, with hooklike win g s in males used to g rasp the female durin g matin g, and scalelike reduced win g s in females. The y are sometimes called “snow fleas” as the y ma y be found walkin g or j umpin g on snow patches. Onl y two, widel y dis j unct species, A ustromerope pou l toni ( Western Austra li a) an d M erope tu b e r ( eastern Un i te d States), are p l ace di nt h e MEROPEIDAE. T h ese are coc k roac hlik e i n appearance as t h e i r h ea d is lar g el y hidden beneath the enlar g ed pronotum. Males have elon g ate g enitalia possi - b l y used in fi g htin g for females. N otiothauma reed i , found in central Chile, is the onl y member of the NOTIOTHAUMIDAE. Similarl y , A pteropanorpa tasmanica f r o mT as m a- n i a, w hi c h resem bl es b ore id s i n genera lf orm an dh a bi ts, i st h eso l e representat i ve o f th e APTEROPANORPIDAE. CHORISTIDAE, w i t h on l ye i g h t spec i es, are restr i cte d to Australia. The two remainin g families, PANORPODIDAE and PANORPIDAE (Fi g ur e 9.2B), are the most specialized mecopteran g roups. The former includes onl y two g enera, Brachypanorp a ( four s p ecies in the United States) and P anorpode s (five s p ecies in Ja p an an d Korea), w hil et h e l atter i st h e l argest mecopteran f am il y (300 spec i es) w i t h an essent i a ll y h o l arct i c di str ib ut i on, t h oug hi t h as representat i ves i nAs i a. Literatur e Because o f t h e centra l pos i t i on t h at t hi sor d er occup i es i nany di scuss i on o f t h eevo l ut i on o f most en d opterygote or d ers, i nterest i nt h e Mecoptera h as b een out o f proport i on to t h e number of extant species. Information on the biolo gy of Mecoptera is provided b y Webb et al. (1975), Kaltenbach (1978), and B y ers and Thornhill (1983). The evolution of th e order and its relationship to the other panorpoid groups are discussed by Hennig (1981), Kr i stensen (1981), W ill mann (1987), an d W hi t i ng (2002). Keys to f am ili es o f Mecoptera in N ort h Amer i ca an d Austra li a are prov id e db y Arnett (2000) an d Byers (1991), respect i ve l y . Th eBr i t i s h spec i es are id ent ifi a bl e f rom P l ant (1997) . A rnett, R. H. Jr., 2000, American Insects: A h an db oo k of t h e Insects of America Nort h of Mexico , 2n d e d ., CRC P ress , Boca Raton , FL. B yers, G. W., 1991, Mecoptera, in : T he Insects o f Australia , 2nd ed., Vol. 2 (CSIRO, ed.), Melbourne Universit y P ress, Car l ton, V i ctor i a . By ers, G. W., an d T h orn hill , R., 1983, B i o l o gy o f t h e Mecoptera , A nnu. Re v . Entomo l. 28 : 203–228 . H ennig, W., 1981, Insect Ph y lo g en y , W iley, New York. WW H inton, H. E., 1958, The phylogeny of the panorpoid orders , Annu. Rev. Entomo l . 3 :181–20 6 . Ka l ten b ac h , A., 1978, Mecoptera (Sc h na b e lh a f te, Sc h na b e lfli e g en), i n: Han db uc hd er Zoo l ogie ,V ol. IV, Insecta V V L fg . 28 :1–111, de Gruyter, Berlin. Kr i stensen, N. P., 1981, P h y l ogeny o fi nsect or d ers , A nnu. Rev. Entomo l . 2 6 : 135–157 . Pl ant, C. W., 1997, A k e y to a d u l ts o f Br i t i s hl acew i n g san d t h e i ra lli es (Neuroptera, Me g a l optera, Rap hidi opter a and Mecoptera) , Field S tud. 9 :179–269 . W ebb, D. W., Penny, N. D., and Marlin, J. C., 1975. The Mecoptera, or scorpionflies, of Illinois, Bu ll .I ll . Nat . H ist. S ur v. 31 ( 7 ) :2 5 1–316 . W hiting, M. F., 2002, Mecoptera is paraphyletic: Multiple genes and phylogeny of Mecoptera and Siphonaptera , Zoo l . Scrip. 3 1: 93 – 104 . W illmann, R., 1987, The phylogenetic system of the Mecoptera, WW Syst. Entomo l . 12 : 5 1 9 – 5 24 . 3 . Diptera S ynonyms: n one C ommon names : true flies; includes mosquitoes, mid g es, black flies, dee r flies , horse flies , house flie s 244 CHAPTER 9 Generally minute to small soft-bodied insects; head highly mobile with large compound eyes, a ntennae o f var i e d s i ze an d structure, an d suctor i a l mout h parts; prot h orax an d metat h orax sma ll a n df use d w i t hl ar g e mesot h orax, w i n g s present on ly on mesot h orax, h a l teres present on metat h o- r ax; legs with five-segmented tarsi; abdomen with varied number of visible segments, femal e g en i ta li as i mp l e i n most spec i es, ma l e g en i ta li a comp l ex, cerc i present. Larvae eruc if orm an di n most spec i es apo d ous; h ea di n many spec i es re d uce d an d retracte d . P u p ae a d ect i cous an d o b tect or exarate, t h e l atter enc l ose di na p u p ar i um. T he more than 120,000 s p ecies of Di p tera described to date re p resent p erha p s two-thirds o f the world total. The order has a truly worldwide distribution, representatives occurring ev e n in the Antarctic. More than 19, 5 00 species have been described from North America, a l most 8000 f rom Austra li a , an d c l ose to 7000 f rom Br i ta i n. Some o f t h ewor ld ’s commonest i nsects and a lar g e number of species of veterinar y and medical importance belon g to this o rder . S tructur e T he g reat structural diversit y found in the Diptera reflects the variet y of niches that th e true flies have ex p loited. A du lt . Adults range in size from about 0. 5 mm to several centimeters and the y are g enera lly so f t- b o di e d .T h e h ea di sre l at i ve ly l ar g ean d highly mo bil e. It carr i es we ll- developed compound e y es, which in males are frequentl y holoptic. The antennae are o f v aried size and structure and are important taxonomicall y . In Muscomorpha-Schizophor a t h e r e i sa ∩ - shaped ptilinal (frontal) suture that runs transversely above the antennae and e xten d s d ownwar d on eac h s id eo f t h em. T hi s suture i n di cates t h e pos i t i on o f t h ept ili num, a m em b ranous sac t h at i s exserte d an ddi sten d e d at ec l os i on i nor d er to rupture t h e pupar i u m and assist a fl y in tunnelin g throu g h soil, etc. The mouthparts are adapted for suckin g an d are described in Chapter 3 (see Section 3.2.2 and Fi g ures 3.14–3.16). The prothoracic and m etathoracic segments are narrow and fused intimately with the very large mesothorax, whi c hb ears t h es i ng l epa i ro f mem b ranous w i ngs. T h e hi n d w i ngs are extreme l ymo difi e d , f orm i ng h a l teres, sma ll ,c l u blik e structures i mportant as organs o fb a l ance (see C h apter 14 , S ect i on 3.3.4). In a f ew spec i es h a l teres an d w i n g s are re d uce d or a b sent. T h e l e g sa l most alwa y s have five-se g mented tarsi, and in some species one or more pairs are modified fo r g raspin g pre y . Primitivel y , there are 11 abdominal se g ments, but in most Diptera this number i s reduced and rarely more than 4 or 5 are readily visible. Frequently, the more posterior segments (posta bd omen) are te l escope di nto t h e anter i or part o f t h ea bd omen (prea bd omen) ( see F ig ure 3.29). T h e posta bd omen t h us f orme di s use d as an extens ibl eov i pos i tor. T h e m ale g enitalia are complex and their homolo g ies uncertain because of the rotation of the abdomen and as y mmetric g rowth of the individual components durin g the pupal sta g e . In most D i ptera t h ec ib ar i um i s strong l y muscu l ar an d serves as a pump f or suc ki ng u p li qu id s i nto t h e gut. In t h e bl oo d suc ki ng Ta b an id ae an d Cu li c id ae a l arge p h aryngea l pump i sa l so present. T h ea li mentar y cana li s, i n most pr i m i t i ve f orms, re l at i ve ly unconvo l ute d . In Muscomorpha, however, it is much more coiled because of the increase in len g th of the m id g ut. The esopha g us divides posteriorl y into the g izzard and, usuall y , one diverticulum, the food reservoir (misleadingly called the “crop”). In Culicidae three diverticula are found . In Nematocera t h em id gut i sas h ort, sac lik e structure; i n Muscomorp h a i t i s l ong an d c onvo l ute d . Genera ll yt h ere are f our Ma l p i g hi an tu b u l es t h at ar i se i npa i rs f rom a common duct on either side of the g ut. In the nervous s y stem a complete ran g e of specialization 245 T HE P A N O RP O ID ORDER S is seen. In primitive Nematocera three thoracic and seven abdominal g an g lia occur, bu t all intermediate conditions are found between this arran g ement and the situation in the more advanced Muscomorpha where a composite thoracoabdominal g an g lion exists. I n f ema l es t h epa i re d ovar i es compr i seavar i e d num b er o f po l ytrop hi covar i o l es. In v i v i parous spec i es t h ere may b eon l y one or two, b ut i nt h ema j or i ty o f ov i parous fli es t h ere may b e more t h an 100. In v i v i parous f orms t h e common ov id uct i s dil ate d to f orm a uterus, an d t he accessor y (“milk”) g lands produce a nutritive secretion. One to four spermathecae ar e alwa y s present. In males the testes are g enerall y small, ovoid, and pi g mented. The short , pa i re d vasa d e f erent i a l ea di nto a muscu l ar e j acu l atory sac. Pa i re d accessory g l an d s may occur. L arva an dP u p a . Larvae are usuall y elon g ate and c y lindrical. Bod y se g mentatio n is usually distinct, but in a few groups the true number of segments is masked as a re - su l to f secon d ary di v i s i on or f us i on o f t h eor i g i na l segments. True t h orac i c l egs are a l- w ays a b sent, t h oug h pro l egs may occas i ona ll y b e present on t h et h orax an d /or a bd omen . I n primitive Diptera the head capsule is distinct and sclerotized (the eucephalous condi- t ion). In most species, however, it is much reduced (hemicephalous) or entirel y vesti g ia l (acephalous) (Fi g ure 3.13). The antennae and chewin g mouthparts, includin g horizontall y mov i ng man dibl es, are we ll d eve l ope di n Nematocera. In ort h orr h ap h ous spec i es anten - nae an d max ill ae may b ewe ll d eve l ope d , b ut t h e man dibl es are s i c kl e-s h ape d an d mov e i navert i ca l p l ane. In Muscomorp h at h e antennae are i nt h e f orm o f m i nute pap ill ae an d t he mouthparts are reduced to a pair of curved hooks (the ori g inal mandibles). The internal structure of larvae g enerall y resembles that of adults. Dipteran pupae are alwa y s adecticous . Pu p ae o f nematocerous an d ort h orr h a ph ous s p ec i es are o b tect, w h ereas t h ose o f Musco- morp h a are secon d ar il y exarate an d coarctate, b e i ng enc l ose di n a pupar i um, t h e h ar d ene d cut i c l eo f t h et hi r dl arva li nstar. L if eH i story and Hab i ts Adult Diptera are active, mostly free-living insects that are found in all major habitats. Th ey are pre d om i nant l y di urna l an d usua ll y assoc i ate d w i t hfl owers or w i t hd ecay i ng organ ic matter. W i t h t h e except i on o f a f ew spec i es t h at d o not f ee d as a d u l ts, fli es f ee d on li qu id s (rarel y , also pollen), a habit that ma y reflect the use of hone y dew from homopterans as t heir ancestral ener gy source (Downes and Dahlem, 1987). Most Diptera feed on nectar or the j uices from deca y in g or g anic matter, but a few g roups are adapted for feedin g o n th et i ssue fl u id so f ot h er an i ma l s espec i a ll y art h ropo d san d verte b rates. T hi s i sac hi eve din some spec i es b ys i mp l y cutt i ng t h es ki n or squeez i ng prey w i t h t h e l a b e ll aan d suc ki ng up th eexu d e dfl u id .Int h ema j or i t y o fb o dy - fl u id f ee d ers, h owever, a fi ne pro b osc i s i s use d t o pierce the skin and penetrate directl y to the fluid, usuall y blood. The habit is usuall y confined to females. It is throu g h the bloodsuckin g habit and the subsequent importanc e o f t h ese i nsects as vectors o fdi sease-caus i ng m i croorgan i sms t h at t h eor d er i s genera lly cons id ere d t h e most i mportant o f t h e ent i re c l ass f rom t h eme di ca l an d veter i nary po i nt o f v i ew . A l t h oug h part h enogenes i s i s k nown to occur i na f ew spec i es, most D i ptera repro d uce b isexuall y . Copulation is preceded in some species b y an elaborate courtship. Usuall y females activel y search for, and la y e gg s directl y on, the larval food source. Members of af ew f f g roups are ovoviviparous or viviparous. Egg development is normally rapid and h atc hi ng occurs i na f ew d ays. Larvae are usua ll y f oun di nmo i st l ocat i ons suc h as so il ,mu d , d ecay i ng organ i c matter , and plant or animal tissues, thou g h a few are trul y aquatic. The ma j orit y are liquid feeders o r 2 4 6 CHAPTER 9 m icropha g ous. Some aquatic forms trap their food in speciall y developed mouth brushes . Larvae of man y species are of a g ricultural or medical importance. Usuall y four larval i nstars occur, but up to ei g ht are found in some species, and in Muscomorpha the fourth i s suppresse d .Pr i or to pupat i on, l arvae genera ll y craw l to a d r i er l ocat i on. Pupae may b ena k e d, b u t th ose o f many nematocerous an d ort h orr h ap h ous spec i es are enc l ose di n a cocoon, an d t h ose o f Muscomorp h a are ens h eat h e dby t h e pupar i um. T h ep h arate a d u l tswa ll ows a i r to facilitate emer g ence from the pupal skin or puparium, the latter splittin g len g thwise in o btect p u p ae; in coarctate forms, however, the adult p rotrudes the p tilinum to p ush off the anter i or en d o f t h e pupar i um . P hylo g eny and C lass ifi cat i o n F rom a structural comparison of primitive livin g Mecoptera and Diptera and from the re l at i ve l y scarce f oss il ev id ence, i t seems pro b a bl et h at D i ptera evo l ve df rom mecoptera lik e i nsects i nt h e Perm i an per i o d. P ermoti p u l a p atrici a , f rom t h e Upper Perm i an o f Austra li a, w as o r igi na lly t h ou gh tto b et h eo ld est di pteran. However, i t i s now cons id ere d an ear ly M ecopteran. Similarl y , other four-win g ed forms (the Protodiptera P ermotanytarsus a n d C horistotanyderus) from the same strata, which were ori g inall y considered to be Diptera ( R i e k , 1977), h ave b een reexam i ne d an d , as a resu l t, move df rom t h e di pteran stem group ( W ill mann, 1989; Wootton an d Ennos, 1989). T h us, t h e ear li est re li a bl e recor d so ff os - s il D i ptera come f rom t h eM iddl eTr i ass i c, t h ou gh t h ese are qu i te mea g er. B y t h e l at e Triassic, the more primitive suborder, Nematocera, had under g one a considerable radia- tion, possibl y makin g use of the hone y dew produced b y the alread y abundant homopteran s ( Downes and Dahlem, 1987). By the Lower Jurassic, well-developed Nematocera (som e ass i gna bl e to recent f am ili es) an d pr i m i t i ve ort h orr h ap h ous f orms were present. T h e Mus - c omorp h a (Cyc l orr h ap h a) evo l ve df rom ort h orr h ap h an stoc k , pro b a bl y i nt h e Late Jurass i c . The g reat radiation of the order, and the establishment of man y of the structures and habit s o f modern flies, took p lace in the Cretaceous p eriod. This was, of course, correlated wit h the evolution of the flowering plants and mammals. By the Eocene period, the diptera n f auna was s i m il ar i n many respects to t h at w hi c h surv i ves to d ay. In d ee d , many Eocen e f oss il s are ass i gne d to mo d ern genera. A poss ibl ep h y l ogeny o f t h eor d er i ss h own in F i g ure 9.3. Classification of the modern Diptera continues to present problems, particularl y the rank assi g nable to different g roups. The difficult y arises in part because the order is ex- treme l yo ld ; i t conta i ns many ext i nct groups an d ot h ers t h at are i n d ec li ne; an d yet t h ere ar e a l so groups t h at are st ill evo l v i ng at a rap id rate. T h us, w hil e anc i ent f am ili es h ave we ll - e sta bli s h e d diff erences an d are eas ily separate d , more recent g roups, nota bly t h e f am ili e s and su p erfamilies of Muscomor p ha, are sometimes little more than convenient divisions because of the vast number of s p ecies that must be considered. The differences between t h ese groups, t h ere f ore, are re l at i ve l ym i nor. T h ec l ass ifi cat i on use dh ere i sa d apte df ro m t h ev i ews presente di nt h e Manua l o f Nearctic Diptera (McA l p i ne et a l ., 1981–1989). In t hi s arrangement, t h eD i ptera are arrange di ntwosu b or d ers, Nematocera an d Brac h ycera . Within the Nematocera, seven infraorders are reco g nized. Fi g ure 9.3 indicates that Nema - tocera are monoph y letic. However, it should be noted that some authors believe that this is a paraphyletic group, with the Bibionomorpha or Psychodomorpha being the sister group t o t h e Brac h ycera (Yeates an d W i egmann, 1999). In t h eun d ou b te dl y monop h y l et i c Brac h yc - e ra t h ere are t h ree i n f raor d ers; t h e fi rst two, As il omorp h aan d Ta b anomorp h a, correspon d to the “Orthorrhapha” while the third, Muscomorpha, is equivalent to the “C y clorrhapha” o f 247 T HE P A N O RP O ID ORDER S F I G URE 9.3. A s u gg este d p hyl o g en y o f t h eD i ptera. [A f ter J. F. McA l p i ne an d D. M. Woo d , coor di nators , 1989 , M anual o f Nearctic Dipter a ,V ol. 3, Agriculture Canada Monograph No. 32. By permission of the Minister V V of Supply and Services Canada.] earlier s y stems. Within the Muscomorpha, there are two easil y defined but ver y unequall y sized sections, the probabl y paraph y letic Aschiza and the monoph y letic Schizophora. Th e latter has two subdivisions, Acalyptratae and Calyptratae. S uborder Nematocera Most Nematocera are small, delicate flies, with 6 - to 14-segmented antennae of simple structure, and 3- to 5 -se g mented maxillar y palps. Larvae have a well-developed head an d chewin g mandibles that move in the horizontal plane. 248 CHAPTER 9 FIGURE 9.4 . T i pu l omorp h a. A crane fl y, Ti p u l a trivittata ( T i pu lid ae). [ From F. R. Cole and E. I. Schlin g er, 19 6 9 . T h eF l ies of Western Nort h A m e r ica. By permission of the Universit y of California Press.] T he suborder contains the oldest families of Di p tera, most of which are now on the decline. Some, however, like the Culicidae, have undergone relatively recent radiations and are among t h e most success f u l mo d ern groups. Infraor d er Tipu l omorp h a M em b ers o f t hi s group are p l ace di nt h es i ng l e f am il y TIPULIDAE (F i gure 9.4) ( crane fli es, d a dd y l ong l egs), w hi c h ,w i t h a b out 14,000 spec i es ( i nc l u di n g > 1 5 00 in North A merica), is the lar g est famil y of Diptera. Representatives of this worldwide famil y are m ostl y associated with moist, temperate habitats, thou g h some occur in open meadows, rangelands, and deserts. Adults range in size from small (wingspan of 2 mm) to very larg e ( w i ngspan up to 8 cm). Larvae are f oun di naw id evar i ety o fh a bi tats f rom str i ct l y aquat i c to d ry so il sw h ere t h ey typ i ca ll y f ee d on p l ant mater i a l or d ecay i ng organ i c matter; occa - sionall y , the y become pests b y feedin g on seedlin g field crops. In f raorder Blephariceromorpha Almost all members of this small g roup are included in the widel y distributed famil y BLEPHARICERIDAE (net-winged midges) (200 species). Adults are slender flies wit h l ong l egs, an di n b ot h sexes t h e eyes are h o l opt i c. In some spec i es b ot h sexes f ee d on nectar ; i not h ers f ema l es catc h sma ll er fli es an df ee d on t h e i r h emo l ymp h .A d u l ts are genera lly f oun d near f ast- fl ow i n g streams; t h e aquat i c l arvae attac h t h emse l vestoroc k sw h ere t h e y f eed on diatoms and al g ae . In f raorder Axymyiomorpha Containin g onl y five species in the one famil y AXYMYIIDAE, the taxonomic position o f this group is controversial, and other authorities have included it in a variety of othe r n ematoceran f am ili es. Spec i es occur i n Nort h Amer i ca, eastern Europe, an d S ib er i a. A d u l ts are stout- b o di e dfli es t h at resem bl eB ibi on id ae. Larvae li ve i ncav i t i es excavate di n rott i ng w o od, perhaps feedin g on fun g i or other microor g anisms . [...]... ( 196 4) and Volumes 1 and 2 of the series edited by McAlpine et al ( 198 1– 198 9) are excellent sources of information on the biology of the group Rohdendorf ( 197 4), authors in Volume 3 of McAlpine et al ( 198 1– 198 9), Michelsen ( 199 6), Friedrich and Tautz ( 199 7), and Yeates and Wiegmann ( 199 9) discuss the phylogeny and classification of the order Griffiths ( 199 4) [Brachycera], Oosterbroek and Courtney ( 199 5)... and sea birds Literature Holland ( 196 4, 198 5), Askew ( 197 1), Rothschild ( 197 5), Traub and Starcke ( 198 0), Marshall ( 198 1), and Traub ( 198 5) include a good deal of general information on fleas, especially concerning host-parasite relationships The phylogeny of fleas is discussed by Hennig 268 CHAPTER 9 ( 198 1), Kristensen ( 198 1), Byers ( 199 6), and Whiting (2002) Lewis ( 199 8) summarizes the world flea fauna... 264 CHAPTER 9 ( 199 6) [orthorrhaphans], and Nirmala et al (2001) [Calyptratae] deal with the major subordinal groups Keys for identification of Diptera are given by Oldroyd ( 194 9) (and other authors in Volumes 9 and 10 of the same series) and Colyer and Hammond ( 195 1) [British species]; Cole and Schlinger ( 196 9) and McAlpine et al ( 198 1– 198 9) [North American genera]; and Colless and McAlpine ( 199 1) [Australian... Insect Mol Biol 10:475–485 Oldroyd, H., 194 9, Diptera: Introduction and key to families, R Entomol Soc Handb Ident Br Insects 9( 1): 49 pp W Oldroyd, H., 196 4, The Natural History of Flies, Weidenfeld and Nicolson, London Oosterbroek, P., and Courtney, G., 199 5, Phylogeny of the nematocerous families of Diptera (Insecta), Zool J Linn Soc 115:267–311 Riek, E F., 197 7, Four-winged Diptera from the Upper Permian... CHAPTER 9 FIGURE 9. 12 Tabanomorpha (A) A horse fly, Tabanus opacus (Tabanidae); and (B) a soldier fly, Odontomyia hoodiana (Stratiomyidae) [A, from J F McAlpine, 196 1, Variation, distribution and evolution of the Tabanus (Hybomitra) frontalis complex of horse flies (Diptera: Tabanidae), Can Entomol 93 : 894 92 4 By permission of the Entomological Society of Canada B, from F R Cole and E I Schlinger, 196 9,... and Mardon ( 199 1) [Australian genera], and Smit ( 195 7) [British species] Arnett, R H., Jr., 2000, American Insects: A Handbook of the Insects of America North of Mexico, 2nd ed., CRC Press, Boca Raton, FL r Askew, R R., 197 1, Parasitic Insects, American Elsevier, New York Byers, G W., 199 6, More on the origin of Siphonaptera, J Kansas Entomol Soc 69: 274–277 Dunnet, G M., and Mardon, D K., 199 1, Siphonaptera,... M., and Tautz, D., 199 7, Evolution and phylogeny of the Diptera: A molecular phylogenetic analysis using 28S rDNA sequences, Syst Biol 46:674– 698 Griffiths, G C D., 199 4, Relationships among the major subgroups of Brachycera (Diptera): A critical review, Can Entomol 126:861–880 McAlpine, J F., Peterson, B V., Shewell, G E., Teskey, H J., Vockeroth, J R., and Wood, D M (eds.), 198 1– 198 9, Manual of Nearctic... FIGURE 9. 8 Culicomorpha Chironomus tentans (Chironomidae) (A) Larva; and (B) pupa [From O A Johannsen, 193 7, Aquatic Diptera Part IV Chironomidae: Subfamily Chironominae, Mem Cornell Univ Agri Exp Stn 210:52 pp By permission of Cornell University Agriculture Experimental Station.] 252 CHAPTER 9 FIGURE 9. 9 Culicomorpha A punkie, Culicoides dovei (Ceratopogonidae) [From F R Cole and E I Schlinger, 196 9,... the cabbage FIGURE 9. 17 Muscoidea (A) The house fly, Musca domestica (Muscidae); (B) the wheat bulb fly, Hylemya coarctata (Anthomyiidae); and (C) a dung fly, Cordilura criddlei (Scathophagidae) (A, from V B Wigglesworth, 195 9, Metamorphosis, polymorphism, differentiation, Scientific American, February 195 9 By permission of Mr Eric Mose, Jr B, C, from F R Cole and E I Schlinger, 196 9, The Flies of Western... N.S.W 101:250–255 Rognes, K., 199 7, The Calliphoridae (blowflies) (Diptera: Oestroidea) are not a monophyletic group, Cladistics 13:27–66 Rohdendorf, B., 197 4, The Historical Development of Diptera, University of Alberta Press, Edmonton Willmann, R., 198 9, Rediscovered: Permotipula patricia, the oldest known fly, Naturwissenschaften 76:375–377 Wootton, R J., and Ennos, A R., 198 9, The implications of function . of S imuliu m a r eo f e x - t reme i m p ortance as vectors f or t h e fil ar i a l nemato d e , Onc h ocerca vo l vu l us , whi c h cause s onc h ocerc i as i s(r i ver bli n d ness) i n trop i ca l A f r i ca, Centra l Amer i ca, nort h ern. Willmann, 198 7, The phylogenetic syste m o f t h e Mecoptera , S yst. Entomo l. 12 :5 19 524. By permission of Blackwell Scienti c Publications Ltd.] 242 CHAPTER 9 Suborder Nannomeco p tera Thi ssu b or d er. s p ecies of which are vectors of various virus- and leishmania-induced diseases . T RICHOCERIDAE (w i nter crane fli es) (110 spec i es, wor ld w id e) are eas il y con f use d w i t h th e true crane fli es.