8 The Hemipteroid Orders 1 . Intr oduc t ion T he four orders (Psocoptera, Phthiraptera, Hemiptera, and Th y sanoptera) that constitut e t he hemipteroid g roup are united b y the followin g features: specialized, usuall y sucto- rial, mouthparts; small anal lobe in hind win g ; win g venation reduced; cerci absent; fe w M a l p i g hi an tu b u l es; an d ventra l nerve cor d w i t hf ew di screte gang li a. On t h ew h o l e, t he h em i ptero id group i s more h omogeneous t h an t h e ort h optero id group, a l t h oug h two evo l u- ti onar yli nes h ave d eve l ope d , l ea di n g to t h e Psocoptera-P h t hi raptera, on t h e one h an d ,an d t he Hemiptera-Th y sanoptera, on the other . 2 . Psocoptera S ynonyms: C orrodentia, Copeognatha C ommon names: barklice, booklice, p socids S mall or minute soft-bodied insects; mobile head with long filiform antennae and specialized c h ew i n g mout h parts, compoun d e y es usua lly prom i nent b ut re d uce di n some spec i es; prot h orax s mall, win g s present or absent, le g s with two- or three-se g mented tarsi; external g enitalia of both s exes concealed , cerci absent. This order, containin g about 3200 described species, has a worldwide, thou g h predom- i nant l y trop i ca l , di str ib ut i on. Some 290 spec i es occur i n Nort h Amer i ca, a b out 80 i nBr i ta i n, an d 300 i n Austra li a. S tructure Psocoptera are stoc ky ,so f t- b o di e di nsects w h ose l en g t hi s usua lly l ess t h an 10 mm . T he lar g e, mobile head bears a swollen postcl y peus, lon g filiform antennae, and, usuall y , prominent compound e y es, thou g h the latter are reduced in some win g less species. Three oce lli are usua ll y present i nw i nge df orms b ut a b sent i n apterous spec i es. T h eY-s h ape d ep i cran i a l suture i s prom i nent. T h e mout h parts, t h oug h reta i n i ngac h ew i ng f unct i on, are spec i a li ze d .T h e man dibl es are di ss i m il ar, t h ou gh eac hh as b ot hg r i n di n g an d cutt i n g e dg es. I n the maxillae the cardo and stipes are not alwa y s distinct. The g alea is a lar g e, flesh y lobe , 1 99 200 CHAPTER 8 w hereas the lacinia is a narrow, sclerotized rod (the pick), which ma y be used to scrape food f rom the substrate. The h y pophar y nx, which is able to take up water from the atmospher e ( Rudolph, 1982), has a characteristic structure. The lin g ua bears a pair of ventral sclerite s t h at are connecte d to t h eme di an s i top h ore sc l er i te b y fi ve li gaments. T h es i top h ore sc l er i t e i ss i tuate d on t h e ventra l sur f ace o f t h e b ase o f t h ec ib ar i um. Oppos i te to i t, on t h e d orsa l sur f ace o f t h ec ib ar i um wa ll , i sa k no blik e process t h at i s b e li eve d to move a g a i nst t he sclerite in the manner of a mortar and pestle and facilitate the g rindin g up of food . In win g ed forms the small prothorax is lar g el y concealed b y the pterothorax. The win g s are mem b ranous an dh ave a prom i nent b ut re d uce d venat i on. T h e anter i or pa i r i s l arger t h a n t h e hi n d pa i r. At rest t h ey are h e ld roo flik e over t h e b o d y. T h e f ore an dhi n d w i ngs are c oup l e db ot hd ur i ng fli g h tan d at rest. Vary i ng d egrees o fb rac h yptery occur, even w i t hin the same species, and apter y is common, especiall y in females. The le g s are usuall y slender and similar; in some species the hind coxae carr y what is believed to be a stridulator y o rgan. The abdomen is l0-segmented and terminates in a dorsal epiproct and a pair o f l atera l paraprocts t h at may represent t h e 11t h segment. T h e externa l gen i ta li ao f ma l es are wea kl y d eve l ope d ,an d t h e i r h omo l og i es are uncerta i n. A sma ll ov i pos i tor i s usua lly present in females, thou g h it is much reduced or absent in some forms. Cerci are never p resen t. F our Malpighian tubules originate at the posterior end of the midgut, which is long an d convo l ute d .T h e nervous system i s hi g hl ymo difi e d an d compr i ses on l y fi ve gang li on i c c enters: b ra i n, su b esop h agea l gang li on, prot h orac i c gang li on, a compos i te pterot h orac ic g an g lion, and a composite abdominal g an g lion. The tracheal s y stem usuall y opens to th e e xterior b y means of two pairs of thoracic and ei g ht pairs of abdominal spiracles. Each ov ar y contains three to five pol y trophic ovarioles. The lateral oviducts are short and open i nto a l arger me di an d uct. A spermat h eca i s present. T h e testes are roun di s h or t h ree- l o b e d . T h e vasa d e f erent i a l ea di nto l arge sem i na l ves i c l es t h at appear to pro d uce t h e mater i a l o f t h e spermatop h ore. L ife Histor y and Habit s M ost Psocoptera are f oun d on vegetat i on or un d er b ar k ,t h oug h some li ve among l ea fli tter, un d er stones, or i n caves. A f ew spec i es are assoc i ate d w i t hh umans an d may be encountered in houses or buildin g s in which food materials are stored. Thou g h the y m a y occur in vast numbers, the y are seldom of economic importance. The y are primaril y phytophagous, feeding on algae, lichens, fungi, pollen, and decaying fragments of highe r p l ants; occas i ona ll yt h ey eat d ea d an i ma l matter. Spec i es assoc i ate d w i t hh umans li ve on c erea l pro d ucts or, i nt h e case o f t h e common b oo kli ce, mo ld st h at d eve l op on o ld b oo k s . Man y species are g re g arious, with individuals of all a g es livin g to g ether, often beneath a silken web produced from secretions of the modified labial g lands. Most outdoor species are fully winged, though brachyptery and aptery are common under certain environmenta l c on di t i ons. M a l es are un k nown i n some spec i es, w hil e i not h ers f acu l tat i ve part h enogenes i sma y o ccur. In di oec i ous spec i esama l et y p i ca lly courts a f ema l epr i or to mat i n g .E gg s, b etween 2 0 and 100 at a time, are laid sin g l y or in g roups, usuall y on ve g etation or under bark, and the y ma y be covered with silk, particles of debris, or fecal material. The y have a thin ch or i on an dl ac k m i cropy l es an d aeropy l es. A f ew spec i es are v i v i parous. Larvae usua lly pass t h roug h s i x i nstars pr i or to metamorp h os i s, b ut t hi s fi gure i so f ten re d uce di npo l ymor- p hi c spec i es. T y p i ca lly t h e apterous morp h s h ave one f ewer i nstar t h an t h e f u lly w i n g e d 201 THE HEMIPTER O ID ORDER S forms. Where climatic conditions permit, Psocoptera ma y have several g enerations per y ear ; elsewhere, species are t y picall y univoltine and ma y enter diapause to overcome adverse conditions . Phylogeny and C lass ifi cat i on Modern Psocoptera are but the remnants of an order that had alread y under g one an extensive evolution b y the end of the Permian period. Of the four hemipteroid orders, th e Psocoptera are genera ll y cons id ere d to b ec l osest to t h e ancestra l stoc k .T h e ear li est f oss il Psocoptera ( f rom t h e Lower Perm i an o f Kansas [U.S.A.] an d Morav i a [Czec h Repu bli c]) diff er f rom mo d ern spec i es w i t h re g ar d to w i n g venat i on an d mout h part c h aracters an d ar e placed in a distinct suborder Permopsocina. However, man y of the numerous Oli g ocene fossils, and even some Cretaceous species, can be assi g ned to extant families (some eve n t o extant genera) indicating that the order has undergone relatively little change since the M esozo i c. L i v i ng Psocoptera are di v i s ibl e i nto t h ree we ll d e fi ne d su b or d ers: Trog i omorp h a , T roctomorp h a, an d Psocomorp h a (Eupsoc id a). T h e Trog i omorp h a conta i ns t h e most pr i m - itive and the Psocomorpha the most advanced Psocoptera. The Psocomorpha has been the d ominant suborder since the Late Cretaceous. Unfortunatel y , insufficient s y stematic work h as been done to p ermit firm conclusions to be reached with res p ect to the relationshi p so f t he many families. This is especially true for the Psocomorpha, to which at least 7 5 %ofthe recent species belong. The classification of Badonnel (19 5 1), which is used here, continues t o be accepted as the one that most accuratel y reflects ph y lo g enetic relationships within th e order ( Smithers, 1991; Mockford, 1993 ) . S uborder Trogiomorph a D i st i n g u i s hi n g c h aracters o f Tro gi omorp h a i nc l u d e antennae w i t h more t h an 20 se g - ments, never secondaril y annulated; tarsi three-se g mented; labial palps two-se g mented ; pterosti g ma not thickened, or absent; and paraprocts with stron g posterior spine. I nt hi ssu b or d er t h ema j or f am ili es are t h e LEPIDOPSOCIDAE, TROGIIDAE, PSO - QUILLIDAE, and PSYLLIPSOCIDAE. Lepidopsocidae (16 5 species) form a primarily t rop i ca lg roup o fb ar k -an dl ea fli tter- i n h a bi t i n gf orms reco g n i ze dby t h e i r somew h at mot h - like appearance produced b y the scales on their bod y and win g s. Thou g h a small famil y (about 20 species), the Tro g iidae is a cosmopolitan g roup that includes several species found in buildings. Examples ar e Tr o g ium pulsatoriu m ( Figure 8.1A), a common book - l ouse t h at f ee d s on paper, vegeta bl e matter, an d cerea l pro d ucts, an d L epinotus inqui l inus th at i s f oun d espec i a ll y i n granar i es an d ware h ouses. Psoqu illid ae an d Psy lli psoc id ae are b oth small (about 20 species in each), widel y distributed families. Psoquillids are found o n b ark, in bird nests, and litter. Ps y llipsocids occur in caves and termite nests. Both families include some s p ecies found indoors. S uborder Troctomorph a Features of Troctomorpha are 12- to 17-se g mented antennae, with some fla g ellar se g - ments secondaril y annulated; 2- or 3-se g mented tarsi; 2-se g mented labial palps; pterosti g ma not t hi c k ene d ;an d paraprocts w i t h out a strong poster i or sp i ne . T h e l argest f am ili es i nt hi ssu b or d er are t h e LIPOSCELIDAE (140 spec i es) , AMPHIENTOMIDAE (40 species), and PACHYTROCTIDAE ( 6 0 species). Th e 202 CHAPTER 8 F IGURE 8.1 . P socoptera. ( A ) Tro g ium pu l satoriu m ( Trog iid ae) ( di sta l antenna l segments om i tte d ); (B) L i- posce l is sp .(L ip osce lid ae); an d (C ) E ctopsocus ca l ifornicus ( Ecto p socidae). [A, from P P. Grass´e (ed.), 19 5 1 , Tr ait r r ede ´ Z oolo g ie, V ol. X. By permission of Masson, Paris. B, C, from L. A. Swan and C. S. Papp, 1972, V V Th e C ommon Insects of Nort h America. C opyr i g h t 1972 b yL.A.Swanan d C. S. Papp. Repr i nte db y perm i ss i on o f Har p er&Row,Pu bli s h ers, Inc.] L i posce lid ae i s a cosmopo li tan group w h ose mem b ers are recogn i ze db yt h e i r great l y en- l ar g ed hind femora. The famil y includes a number of common booklice ( L i p osceli s s pp. ) ( Fi g ure 8.1B) found in houses, warehouses, and ship holds. Outdoor species t y picall y occu r i n litter and under bark. However, occasionally they have been taken in the nests of ver- te b rates, i nt h e f ur o f mamma l s, an d on bi r d s’ f eat h ers, t h ese assoc i at i ons poss ibl ya idi ng di spersa l o f t h e psocopterans. Amp hi entom id s are pre d om i nant l y f oun di n trop i ca l reg i ons o f the Old World. The y occur under bark and in litter. Pach y troctidae are also mainl y trop - i cal, occurrin g in both the Old and the New World. T y picall y , the y are found under bark o r i n litter, occasionall y in buildin g s. Suborder Psocomor p ha F eatures o f Psocomorp h a i nc l u d e antennae a l most a l wa y s 13-se g mente d ,w i t hfl a g e ll ar se g ments not secondaril y annulated; tarsi 2- or 3-se g mented; labial palps unse g mented; pterosti g ma not thickened; and paraprocts with stron g posterior spine. Ju d g i ng b yt h e i r common f eatures, t hi s very l arge su b or d er, conta i n i ng more t h an 20 f am ili es, i s pro b a bl y d er i ve df rom or h a d a common ancestry w i t h t h e Troctomorp h a. Mos t m em b ers o f t h esu b or d er are f oun d out d oors, on g row i n g ve g etat i on, i n l ea fli tter, or on b ar k . The AMPHIPSOCIDAE (140 species) is a cosmopolitan famil y that includes some of the l ar g est Psocoptera. The y are g enerall y found on broad-leaved folia g e. ARCHIPSOCIDAE ( 60 species) form a tropical family, largely from South America and Africa, some species of whi c hli ve i n mass i ve aggregat i ons un d er s h eets o f we bbi ng t h at may cover an ent i re tree . T h e CAECILIIDAE i saver yl ar g e f am ily (370 spec i es most ly i nt h e g enu s C aeci l iu s ) o f w orldwide distribution. Most species are folia g e dwellers. ECTOPSOCIDAE (120 species) , w hich form a cosmopolitan g roup, are t y picall y found in dr y folia g e and leaf litter thou gh af ew ff sp ecies are found elsewhere; for exam p le, E ctopsocus cali f ornicu s ( Figure 8.1C) is a co mm o n booklouse a n d E. pumi l is i s a cosmopo li tan spec i es f oun di n granar i es an d ware- h ouses. PERIPSOCIDAE (120 spec i es) are b ar kd we ll ers w i t h a cosmopo li tan di str ib ut i on . Members of the ELIPSOCIDAE and EPIPSOCIDAE (each with about 100 species) are 203 THE HEMIPTER O ID ORDER S b ark and litter inhabitants, with worldwide (especiall y Southern Hemisphere) and primar - il y tropical distributions, respectivel y . The cosmopolitan LACHESILLIDAE (250 species ) are primaril y found on dr y folia g e or in leaf litter with a few widespread species (e. g ., Lac h esi ll ape d icu l ari a ) occurr i ng i n granar i es, b arns, an d ware h ouses. MYOPSOCIDAE (12 5 species) are generally large, tropical Psocoptera found on bark. PHILOTARSIDAE (1 5 0 species) live on bark or low ve g etation. The famil y is widel y distributed but especiall y common in the southwest Pacific re g ion. The PSEUDOCAECILIIDAE (120 species) is a mainl y tropical g roup of folia g e- and bark-dwellin g species. The PSOCIDAE, a cosmopoli- t an group of about 5 20 species, is the largest family in the order. Its members are darkly co l ore d an dli ve on b ar k or, occas i ona ll y, on t h e groun d . Literatur e A d eta il e d account o f t h e bi o l o gy o f t h e Psocoptera i s gi ven by New (1987) an d M ockford (1993). Thornton (1985) discusses the zoo g eo g raph y and ecolo gy of the arborea l forms. The ph y lo g en y and classification of the order are considered b y Smithers (1972) . N ew (1987) an d Sm i t h ers (1990) prov id e k eystot h ewor ld f am ili es an d genera, respect i ve l y . N ew (1974) h as a k ey to t h eBr i t i s h spec i es, an d Sm i t h ers (1991) an d Moc kf or d (1993) k eystot h e Austra li an f am ili es an d Nort h Amer i can spec i es, respect i ve l y. B adonne1, A., 1951, Ordre des Psocopt`eres, in: Tr ait r r ede ´ Z oo l ogie (P P. Grass´e, e d .), Vo l . X, Masson, Par i s . M oc kf or d, E. L. , 1993 , Nort h American Psocoptera (Insecta ) , San dhill Crane Press , Ga i nesv ill e , FL. N ew, T. R., 1974, Psocoptera, Han d b. I d ent. Br. In s ect s 1 (7):1–102. N ew, T. R., 1987, B i o l ogy o f t h e Psocoptera, Or. Insects 21 :1 – 109. R u d o l p h , D., 1982, Occurrence, propert i es an dbi o l o gi ca li mp li cat i ons o f t h e act i ve upta k eo f water vapour f rom t he atmosphere in Psocoptera , J . I nsect Ph y siol . 2 8 : 111–121 . S m i t h ers, C. N., 1972, T h ec l ass ifi cat i on an d p h y l ogeny o f t h e Psocoptera , M em. Aust. Mus . 14 :1 – 349. S m i t h ers, C. N., 1990, Ke y stot h e f am ili es an dg enera o f Psocoptera , T ec h. Rep. Aust. Mus . 2 : 1– 8 2 . S mithers, C. N., 1991, Psocoptera, in : T he Insects o f Australia , 2nd ed., Vol. 1 (CSIRO, ed.), Melbourne Universit y P ress, Car l ton, V i ctor i a . T hornton, I. W. B., 198 5 , The g eo g raphical and ecolo g ical distribution of arboreal Psocoptera, A nnu. Re v . Entomo l . 3 0:175–196. 3 . Phth i ra p tera S ynonyms: Pseu d or hy nc h ota, Ma ll op h a ga C ommon names: li ce, suc ki n gli ce ( i n part), (in part), Lipoptera (in part) , chewing lice or bird lice Si p h uncu l ata ( i n part), ( i n part) Ano p lura (in p art) Minute to small, apterous, dorsoventrally flattened ectoparasites of birds or mammals; head prognat h ous or h ypognat h ous, compoun d eyes re d uce d or a b sent, oce lli a b sent, antennae 3 - to 5 -se g mented, mouthparts of chewin g or piercin g -suckin g t y pe with palps of maxillae and l abium reduced or absent; prothorax free or fused with pterothorax, legs with unsegmented or 2 -se g mente d tars i , more or l ess mo difi e df or c li n gi n g to h a i ror f eat h ers, one, two, or no tarsa l claws; abdomen 7- to 10-segmented, cerci absent. T h eor d er, w hi c hh as a wor ld w id e di str ib ut i on, i nc l u d es a b out 3100 d escr ib e d spec i es . T he suckin g lice (suborder Anoplura) are exclusivel y parasites of placental mammals, w hile the remainder are chewin g lice (“Mallopha g a”) comprisin g the suborders Ambl y cera , 204 CHAPTER 8 Ischnocera, and Rh y nchophthirina. About 8 5 % of the species of chewin g lice are parasites o f birds. Man y Phthiraptera are important pests, in their role as disease vectors, of domesticated animals and humans . S tructur e T he minute to small (0.35–10 mm lon g ) bod y is dorsoventrall y flattened and shows av a ried de g ree of sclerotization. In chewin g lice the head is relativel y lar g e and bear s ch ew i ng mout h parts t h at s h ow certa i n resem bl ances to t h ose o f Psocoptera. In suc ki ng li c e t h ere l at i ve l y sma ll h ea dh as part i a ll y retracte d , suctor i a l mout h parts. T h e l a b rum f orms a s h ort evers ibl e pro b osc i s. T h ree st yl ets are conta i ne d w i t hi n a pouc h t h at runs ventra lly o ff the cibarium. The ventral st y let represents the modified labium, the middle st y let probabl y i s an extension of the openin g from the salivar y duct, and the dorsal st y let is either th e m odified maxillae or the hypopharynx. The mandibles disappear during embryogenesis . T h e h ea db ears t h ree- to fi ve-segmente d antennae t h at may b e filif orm ( i n Anop l ura), cap- i tate an di n grooves (Am bl ycera), or mo difi e df or grasp i ng (Isc h nocera). Compoun d eyes are reduced or absent and ocelli are never present. In Anoplura the thoracic se g ments ar e f used, but in other lice the p rothorax is distinct from the p terothroax. The well-develo p ed l egs include a two-segmented or unsegmented tarsus and usually one or two tarsal claws. E i g h t to ten v i s ibl ea bd om i na l segments occur. Ma l e gen i ta li a i nc l u d e a permanent l yev- e rte d en d op h a ll us. T h ere i s no true ov i pos i tor t h oug hi na ll suc ki ng an d some c h ew i n g l ice the g onapoph y ses on se g ment 8 are used to attach e gg s to the host’s hair. Cerci ar e absent . In sucking lice the cibarium and pharynx form a strong sucking pump, but the crop an d g i zzar d are poor l y diff erent i ate d .Inc h ew i ng li ce t h e crop i s l arge. In a ll P h t hi rapter a t h em id gut i s l arge an dh as two mesenter i c ceca. Four Ma l p i g hi an tu b u l es enter t h em id gu t poster i or ly .T h e nervous s y stem i s highly mo difi e d an di nc l u d es a compos i te metat h ora - c oabdominal or thoracoabdominal g an g lion. The internal reproductive or g ans g enerall y resemble those of Psocoptera except that female Phthiraptera ma y have accessor yg land s an dl ac k as p ermat h eca . Lif eH i story and Hab i t s Generall y , lice are considered to be hi g hl y host-specific bein g restricted to a sin g le o r af ew ff closely related species of host, often occupying specific regions on the host’s body. H owever, Mars h a ll (1981) quest i one d w h et h er taxonom i c pro bl ems w i t h t hi s group may b eo b scur i ng t h e rea l s i tuat i on, c i t i ng t h e examp l eo f Menacant h us eur y sternu s , w hi c h, o n the basis of a recent revision, now counts 118 passerine species (in 20 families) and 5 w oodpecker species amon g its hosts! B y contrast, a g iven bird or mammal ma y pla y host t o several species of lice, up to 15 being recorded on a South American tinamou. In contrast to fl eas, li ce spen d t h e i r ent i re lif eont h e h ost. T h ey are hi g hl y sens i t i ve to c h anges in temperature an dh um idi ty an d surv i ve f or on l ya f ew d ays s h ou ld t h e h ost di e. M ost species of chewin g lice live amon g the feathers of birds where the y feed on f ra g ments of feathers and skin. A few species, for example, the chicken bod y louse , Me n- acanthus stramineu s , feed on blood in addition to e p idermal p roducts and are able to p ierc e t h es ki nor d eve l op i ng qu ill s. T h e mem b ers o f two sma ll f am ili es o f c h ew i ng li ce are para- s i t i c on mamma l s, t h oug h t h e i r genera lh a bi ts are lik et h ose o f spec i es f oun d on bi r d s. Som e ch ew i n gli ce, lik e Psocoptera, h ave t h ea bili t y to ta k eupmo i sture f rom t h e atmosp h ere v i a 205 THE HEMIPTER O ID ORDER S t heir h y pophar y nx, presumabl y an adaptation to prevent desiccation in the dr y air amon g feathers or fur of their host (Rudol p h, 1983). The suckin g lice feed exclusivel y on the blood of the host, alwa y s a placental mammal. I t h as b een suggeste d t h at a poss ibl e reason f or t h e hi g hh ost spec ifi c i ty o f suc ki ng li ce i s th e l et h a l e ff ect an unsu i ta bl e h ost’s bl oo d m i g h t h aveont h e sym bi ot i c b acter i a present in certa i n g ut ce ll sor i nt h em y cetome, a structure c l ose ly assoc i ate d w i t h t h e g ut (see C h apte r 16, Section 5.1.2 ) . H eav y infestations of lice ma y render a host more susceptible to disease and cause econom i c l oss re l ate d to re d uct i on i n qua li ty.Ina ddi t i on, some suc ki ng li ce are i mportant di sease vectors (see Su b or d er Anop l ura). I n some Anop l ura, at l east, mat i ng occurs f requent l y, presuma bl y b ecause f ema l es l ac k a spermatheca. In man y species of chewin g lice, males are less common than females, and in some species, are rare or unknown so that partheno g enesis occurs. E gg s are usuall y cemented to hairs or feathers by means of a secretion from the female’s accessory gland . Postem b ryon i c d eve l opment i s rap id , j uven il es pass i ng t h roug h t h ree mo l ts, an d a d u l ts b ecome sexua ll y mature w i t hi na f ew d ays o f t h e fi na l mo l t. Trans f er to new h osts i s b y ph y sical contact and occurs durin g matin g , communal roostin g , and broodin g and feedin g t he y oun g . Some bloodsuckin g Diptera ma y carr y lice from host to host . Phylogeny and Classification I tis g enerall y accepted that the Phthiraptera are derived from a free-livin g Psocopter - alike ancestor, based on a number of s y napomorphies (Chapter 2, Section 3.2). There i s virtuall y no fossil record: the Earl y Cretaceous chewin g louse S aurodectes vrsanskyi may h ave paras i t i ze d pterosaur rept il es, an d a suc ki ng l ouse, N eo h aematopinus re l ictu s has been f oun d onaro d ent , C ite ll us, f rom t h eP l e i stocene o f S ib er i a. However, t hi s h as not prevente d some aut h ors f rom specu l at i n g t h at t h eor d er ma yb equ i te anc i ent, poss ibly or igi nat i n g as earl y as the Upper Carboniferous/Lower Permian. Kim and Ludwi g (1982) proposed thi s v e r y earl y ori g in for the order to be consistent with their h y pothesis that the g roup aros e f rom an ancestor w i t hi nt h e psocopteran su b or d er Permopsoc id a, an id ea re j ecte db yLya l (198 5 ). Rather, Lyal (198 5 ) proposed, the lice may have evolved from liposcelidlike Pso - coptera t h at, as note di nt h e prev i ous sect i on, h ave occas i ona ll y b een f oun di nt h e nests an d on the bod y of birds and mammals. Initiall y , this association perhaps aided dispersal of the insects, but in time the y ma y have be g un to feed on flakes of dead skin, bits of feathers, etc. At this stage, the association between the ancestral louse and its host presumably woul d h ave b een f acu l tat i ve l y paras i t i can d muc hl ess h ost-spec ifi ct h an i st h e case w i t h mo d er n P h t hi raptera. T h ev i rtua l a b sence o ff oss il s h as meant t h at proposa l sw i t h respect to t h eor i - g in and ph y lo g en y of the order have come lar g el y from examination of the host association s of lice, and the ph y lo g en y and zoo g eo g raph y of their mammalian and avian hosts. Suc h studies su gg est that ancestral lice did not arise until the Cretaceous, with either birds o r mamma l sas h osts. T h e ear li est li ce were c h ew i ng f orms, an df rom t h ese arose t h emo d ern Am bl ycera on t h e one h an d ,an d a li ne l ea di ng to t h e rema i n i ng groups on t h eot h er. T h e l atter i tse lf sp li t, gi v i n g r i se to t h e Isc h nocera, w h ose ancestor reta i ne d c h ew i n g mout h part s and had either a bird or a mammal as host, and the common ancestor of the Rh y ncoph - t hirina and Anoplura which, like all modern members of these two g roups, presumabl y f e d on mamma l s. W hil et h eR h yncop h t hi r i na reta i ne d c h ew i ng mout h parts, t h e Anop l ur a e v ol ve d suctor i a l mout h parts i n con j unct i on w i t h t h e i r bl oo d - f ee di ng h a bi t. T h oug h t hi s sc h eme ma yb e a reasona bl eexp l anat i on o f t h eevo l ut i on o f P h t hi raptera, Bar k er (1994 ) 206 CHAPTER 8 F IGURE 8.2 . A s uggeste d p h y l ogeny o f t h eP h t hi raptera. T h ere l at i ons hi po f t h eTr i menopo did ae to ot h e r A m bly ceran f am ili es i s uncerta i n. has stressed that the coevolution of lice and their vertebrate hosts has not alwa y s occurred, and that host-switching by lice is fairly common. Thus, a phylogeny based on lice character s rat h er t h an c h aracters o f t h e h ost i s pre f era bl e. Some aut h ors p l ace t h ec h ew i ng li ce an d suc ki ng li ce i n separate or d ers, t h eMa ll op h ag a and Anoplura, respectivel y , the g reat differences in mouthpart structure and feedin g habit s bein g taken as sufficient j ustification for this separation. Generall y , however, all lice ar e i ncluded in the order, Phthiraptera, divisible into four suborders, Amblycera, Ischnocera , R h yncop h t hi r i na, an d Anop l ura. T h us, t h eMa ll op h aga, w hi c h compr i ses t h e fi rst t h re e su b or d ers, i s a parap h y l et i c group i nt hi ssc h eme, l ea di ng some aut h or i t i es (e.g., Bar k e r et al. , 2003) to ur g e that the term be abandoned. A su gg ested ph y lo g en y of the Phthirapter a i s presented in Fi g ure 8.2 . Su b or d er Am bly cer a M embers of the suborder Ambl y cera have the followin g characteristics: capitate, four- segmente d antennae, l y i ng i n grooves; man dibl es h or i zonta l , max ill ary pa l ps present; an d m esot h orax an d metat h orax usua ll y separate . Generall y six families are reco g nized in this suborder, a g roup of about 8 5 0 species that is considered to contain the more primitive lice. Of the six families, three are restricte d to avian hosts, two to marsu p ials, and one to p lacental mammals. The MENOPONIDAE f orm the largest family (650 species) and have a cosmopolitan distribution. Its members i n f est bi r d s, an d severa l spec i es are i mportant pests o f pou l try, f or examp l e, Menacant h u s s tramineu s (t h ec hi c k en b o d y l ouse) an d M enopon g a ll inae (t h es h a f t l ouse) (F i gure 8.3A). The LAEMOBOTHRIIDAE are a small famil y containin g the sin g le g enu s Lae m obot hr io n , 207 THE HEMIPTER O ID ORDER S F IGURE 8.3. P hthiraptera. (A) The shaft louse, Menopon g allinae ( Menoponidae); (B) the large turkey louse, C h e l opistes me l eagri d is ( Gon i o did ae ) ; ( C ) t h ere d catt l e l ouse , Dama l inia b ovi s ( Tr i c h o d ect id ae); (D) t h e l on g - n osed cattle louse , L ino g nathus vitul i (Lino g nathidae); (E) the short-nosed cattle louse, H aematop i nus eur y sternus ( Haematopinidae); (F) the human body louse , Pe d iculu s humanu s ( Pediculidae); and (G) the crab louse , P hthiru s p u b i s (P h t hi r id ae). [D, E, f rom L. A. Swan an d C. S. Pa pp 1972, Th e Common Insects of Nort h America . C op y r igh t 1972 b y L. A. Swan and C. S. Papp. Reprinted b y permission of Harper & Row, Publishers, Inc.] th e 14 spec i es o f w hi c h are paras i tes o f water bi r d san dh aw k s. T h e RICINIDAE, conta i n i ng 6 5 species in two g enera, are found on hummin g birds and several families of passerines. Be- cause of the sister- g roup relationship of their hosts, the BOOPIIDAE (35 species), found o n Australian marsu p ials, and the TRIMENOPONIDAE (10 s p ecies), found on South Amer- i can marsup i a l san dhi str i comorp h ro d ents (porcup i nes, gu i nea p i gs, etc.), were f ormer l y th oug h tto h ave h a d a common ancestor. T hi sv i ew no l onger appears tena bl e; rat h er, t he B oopiidae appear to be the sister g roup of the GYROPIDAE ( 6 0 species). This famil y i s endemic to South and Central America, thou g h two species , G yropus ovalis and G liricol a p orcelli, which are found on g uinea pi g s, have been spread b y commerce to other parts of th ewor ld . Most spec i es o f Gyrop id ae li ve on hi str i comorp h ro d ents. S u b or d er Isc h nocera Ch aracter i st i cs o f Isc h nocera are expose d ,t h ree- to fi ve-segmente d filif orm anten- nae; man dibl es vert i ca l , max ill ar y pa l ps a b sent; an d mesot h orax an d metat h orax usua lly fused . 208 CHAPTER 8 Included in this probabl y paraph y letic or even pol y ph y letic suborder of about 17 50 species are two lar g e families. The paraph y letic PHILOPTERIDAE, a cosmopolitan g rou p w ith about 1460 species, is the lar g est famil y of the order. Its members are parasitic on bi r d san di nc l u d e a num b er o f pest spec i es f oun d on pou l try, f or examp l e, Cuc l uto g aste r h etero g rap h us (t h ec hi c k en h ea dl ouse). T h e cosmopo li tan f am il y TRICHODECTIDAE ( 290 spec i es), w hi c hi s restr i cte d to p l acenta l mamma l s, conta i ns a num b er o f spec i es f ound on domesticated animals, for exam p le, Damalinia ( Bovicola) bovis (cattle) (Fi g ur e 8 . 3C) , D .ov is (shee p ), D. e q u i ( horses ), F elicola subrostratu s ( cats ) , and Trichodectes canis (d ogs), w hi c h can serve as an i nterme di ate h ost f or t h e d og tapeworm, D ip yl i d ium caninu m . T h e GONIODIDAE, f ormer l y i nc l u d e di nt h eP hil opter id ae, are f oun d on ga llif orm an d c o l um bif orm bi r d s. T h e f am il y i nc l u d es some ma j or pou l try pests, f or examp l e, Ch e l o p iste s meleagridis ( lar g e turke y louse) (Fi g ure 8.3B) , G oniodes gigas (lar g e chicken louse) and G . dissimilis ( brown chicken louse ). Suborder Rhyncophthirina M embers of the suborder Rh y ncophthirina have the followin g characteristics: hea d prolon g ed into a rostrum, mandibles at apex of rostrum, and labium and maxillae vesti g ial . T his suborder contains only two species, Haematomyzus elephantis , ap arasite of bot h In di an an d A f r i can e l ep h ants, an d H. h op k insi , w hi c hi n f ests wart h ogs. It h as b een suggeste d t h at t hi s group may resem bl et h e ancestors o f t h e Anop l ura. Certa i n l y, t h e two groups s h ar e a number of primitive characters (L y al, 198 5 ) . Su b or d er Ano pl ura M em b ers o f t h esu b or d er Anop l ura are recogn i ze db yt h e i rre l at i ve l y sma ll h ea d , sty li- f orm mout h parts, an dl ac k o f man dibl es. T he approximatel y 530 species of Anoplura were arran g ed b y Kim and Ludwi g (1978 ) i n 15 families, of which 8 contain 4 or fewer species, The features of the seven lar g est fam- ili es, p l us t h ePe di cu lid ae an d P h t hi r id ae, w hi c hh ave spec i a l s i gn ifi cance f or h umans, are summar i ze db e l ow , T h e ECHINOPHTHIRIIDAE (12 spec i es) are paras i tes o f P i nn i pe dia ( sea l s, sea li ons, an d wa l ruses ) an d t h er i ver otte r (Lutra cana d ensis) . T h e i r b o d y i s covere d w ith stron g spines or scales that retain a film of air over the bod y when submer g ed. It i s e vident that man y of these lice must be ver y lon g -lived, as their hosts spend most of their l ife at sea, coming ashore to breed (when presumably transfer of parasites occurs) for onl y as h ort t i me eac h year. T h e cosmopo li tan LINOGNATHIDAE (70 spec i es) paras i t i ze d ogs ( Lino g nat h us setosus ) ,h y raxes, an d rum i nants, i nc l u di ng s h eep ( L. ovi ll u s a n d L . p e d a l is) , c attle ( L. vituli ) ( Fi g ure 8.3D), and g oat s (L. steno p sis). T he HAEMATOPINIDAE ( 22 s pecies) form a cosmopolitan g roup whose members parasitize un g ulates, includin g several domesticated forms on which the y ma y become serious pests . Haematop i nus eurysternus ( F i gure 8.3E) occurs on catt l e, H . s u is o np i gs, an d H .a si n i on h orses. T h eH O PL O PLE U RI- DAE (a b out 130 spec i es) are a l so a l arge group w h ose h osts are ma i n l yro d ents an d ra bbi ts , b u t a l so i nc l u d emo l es an d s h rews. T h e l ar g est anop l uran f am ily i st h e POLYPLACIDAE , a cosmopolitan g roup whose 175 species ver y lar g el y parasitize rodents, thou g h some are f ound on primates (lemurs, lorises, and g ala g os), rabbits, moles, and shrews. The ENDER - LEINELLIDAE (50 species) are another rodent-infesting group, specifically parasitizin g S c i ur id ae (squ i rre lf am il y). T h e f am il y i sw id esprea d t h oug h not represente di n Austra li a, Mada g ascar, and southern South America. PEDICINIDAE (1 6 species) infest Old Worl d [...]... includes some important pest species, for example, Lygus lineolaris, the FIGURE 8. 18 Tingoidea The sycamore lacebug, Corythuca ciliata (Tingidae) [From R C Froeschner, 1944, Contributions to a synopsis of the Hemiptera of Missouri, Am Midl Nat 31(3):6 38 683 By permission of the American Midland Naturalist.] 2 28 CHAPTER 8 FIGURE 8. 19 Miroidea and Cimicoidea (A) The tarnished plant bug, Lygus lineolaris (Miridae);... Baranowski, R M., 19 78, How to Know the True Bugs (Hemiptera-Heteroptera), William Brown, W Dubuque, Iowa Sorensen, J T., Campbell, B C., Gill, R J., and Steffen-Campbell, J D., 1995, Non-monophyly of Auchenorrhyncha (“Homoptera”), based upon 18S rDNA phylogeny: Eco-evolutionary and cladistic implications within pre-Heteropterodea Hemiptera (s.l.) and a proposal for monophyletic suborders, Pan-Pac Entomol... Heteroptera Phylogeny and classification are discussed by Hennig (1 981 ), Hamilton (1 981 ), Popov (1 981 ), Schuh (1 986 ), Wootton and Betts (1 986 ), Wheeler et al (1993), Campbell et al (1994), Sorensen et al (1995), and von Dohlen and Moran (1995) Keys to the North American families of Hemiptera are given by Arnett (2000) and Slater and Baranowski (19 78) [Heteroptera] Genera of the aquatic and semiaquatic Hemiptera... Hemiptera-Fulgoromorpha and Hemiptera-Cicadomorpha, R Entomol Soc Handb Ident Br Insects 2(3):1– 68; 2(2):1–64, 65–1 48 Macan, T T., 1965, A revised key to the British water bugs (Hemiptera-Heteroptera), 2nd revised ed., Sci Publ F.W Biol Assoc 16: 78 pp Miller, N C E., 1971, The Biology of the Heteroptera, 2nd ed., Hill, London Novotny, V., and Wilson, M R., 1997, Why are there no small species among xylem-sucking insects?... growth They are, however, economically more important through their role 2 18 CHAPTER 8 FIGURE 8. 8 Life cycle of the rosy apple aphid, Dysaphis plantaginea [After D J Borror, D M Delong, and C A Triplehorn, 1976, An Introduction to the Study of Insects, 4th ed By permission of Holt, Rinehart and Winston, Inc.] as vectors of disease-producing viruses Migratory species that are not particularly hostspecific... C R., 1 986 , Homology and function in the wings of Heteroptera, Syst Entomol 11: 389 –400 5 Thysanoptera Synonyms: Physapoda, Thripida Common name: thrips Small to minute slender-bodied insects; head with short four- to nine-segmented antennae, asymmetrical suctorial mouthparts, small but prominent compound eyes, ocelli present or absent; prothorax large and free, legs with unsegmented or two-segmented... pp Kim, K C., 1 985 , Evolution and host associations of Anoplura, in: Coevolution of Parasitic Arthropods and Mammals (K C Kim, ed.), Wiley, New York Kim, K C., and Ludwig, H W., 19 78, The family classification of the Anoplura, Syst Entomol 3:249– 284 Kim, K C., and Ludwig, H W., 1 982 , Parallel evolution, cladistics, and classification of parasitic Psocodea, Ann Entomol Soc Am 75:537–5 48 Kim, K C., Pratt,... persicae, the green peach aphid (Figure 8. 9A), is the classic example, being known as a vector for more than 100 virus diseases PEMPHIGIDAE (ERIOSOMATIDAE) are closely related to the Aphididae The family is small (275 species), widely distributed (often by commerce), and includes both above- and below-ground feeders and many gall-making species Many of the woolly aphids, so-called g , yp , ] because of the... family containing about 380 species Juveniles live beneath bark or in rotting wood More than 1000 species 219 THE HEMIPTEROID ORDERS 220 CHAPTER 8 FIGURE 8. 10 Fulgoroidea (A) Cixius angustatus (Cixiidae); (B) Stenocranus dorsalis (Delphacidae); (C) Scolops perdix (Dictyopharidae); (D) Anormenis septentrionalis (Flatidae); and (E) Fitchiella robertsoni (Issidae) [From H Osborn, 19 38, The Fulgoridae of Ohio,... or other object Eggs are laid in twigs, a process that may cause considerable dieback of the tree FIGURE 8. 12 Cicadoidea Juvenile periodic cicada Magicicada septendecim (Cicadidae) [From C L Marlatt, 1907, The periodical cicada, Bull U.S Dept Agr Bur Ent (N S.) 71: 181 pp.] 222 CHAPTER 8 FIGURE 8. 13 Cicadelloidea (A) The beet leafhopper, Circulifer tenellus (Cicadellidae); (B) the potato leafhopper, . mea l yor fil amen- tous waxy secret i on. Severa l spec i es o f Ps eu d ococcu s ( F i gure 8. 7C) are ma j or pests as t h e y are vectors of disease-causin g viruses. The ERIOCOCCIDAE (felt scales), a cosmopolitan g roup. lac insect, p roduces a secre- tion from which shellac is p re p ared. The PSEUDOCOCCIDAE (2000 s p ecies worldwide) are t h e common mea l y b ugs, so-ca ll e db ecause f ema l es are covere d w i t h a. Aphidoidea an d Cocco id ea occur i nt h eTr i ass i c. T h eCo l eorr h ync h a, w h ose f oss il recor d exten d s b ac k t o th e Lower Jurass i c, s h are c h aracters w i t hb ot h t h e Auc h enorr h ync h aan d t h e