Entomology 3rd edition - C.Gillott - Chapter 7 pps

7 The Plecopteroid, Blattoid, and O rthopteroid Order s 1 . Intr oduc ti on Thi sc h apter d ea l sw i t h t h e f o ll ow i ng 10 or d ers: P l ecoptera, Em bi optera, D i ctyoptera, I soptera, Gr y lloblattodea, Dermaptera, Phasmida, Orthoptera, Zoraptera, and the recentl y established Mantophasmatodea. Members of these orders can be distin g uished from other exopterygotes [the hemipteroid orders (Chapter 8)] by the following features: generalized bi t i ng mout h parts, w i ng venat i on usua ll ywe ll d eve l ope d w i t h numerous crossve i ns (t h oug h l ess net lik et h an t h at o f Pa l eoptera), cerc i present, term i na li ao f ma l e may b e asymmetr i ca l and reduced, man y Malpi g hian tubules, and g eneralized nervous s y stem with several dis - crete abdominal g an g lia. However, as discussed in Chapter 2, the existence of these commo n features should not be taken as confirmation that these orders constitute a monophyleti c group . 2. Pleco p ter a S ynonyms: P erlaria, Perlida C ommon name: s toneflies M o d erate-s i ze d to f a i r ly l ar g eso f t- b o di e di nsects; h ea d w i t hl on g setaceous antennae, wea k mandibulate mouthparts, well-developed compound e y es and two or three ocelli; thorax almost always with two pairs of membranous wings (sometimes reduced), hind pair in most species wi t h a l ar g e ana ll o b e, venat i on f requent ly spec i a li ze d , l e g s id ent i ca l an d w i t h at h ree-se g mente d tarsus; abdomen of most species terminated by long multiannulate cerci, females lacking a true o v i pos i tor, ma l es w i t h out g onost yl es an d p h a lli cor g ans on a bd om i na l se g ment 9 . L arvae aquatic, generally resembling adults except for presence of a varied number of tracheal gill s . More t h an 2000 spec i es o f t hi s very anc i ent or d er h ave b een d escr ib e d , i nc l u di ng j ust over 6 00 from North America, about 30 from Britain, and 200 from Australia. Thou g hth e order has re p resentatives on all continents exce p t Antarctica, most families have a rather restricted distribution . 14 7 148 C H A PTER 7 Structur e Adult . Th ep l ecopteran h ea di s prognat h ous an db ears a pa i ro f e l ongate, mu l t i an - n ulate antennae, well-developed compound e y es, three (rarel y two) ocelli, and weak, often n on-functional bitin g -t y pe mouthparts. Usuall y all the mouthparts are present, but in mem- bers of a few families the mandibles are vesti g ial. The thorax is primitive. Its se g ments are f ree an d t h e prot h orax i s l arge. Two pa i rs o f mem b ranous w i ngs are near l ya l ways present, t h oug hb rac h ypterous an d apterous spec i es occur at hi g h a l t i tu d es an dl at i tu d es. T h e hi n d wi n g t y p i ca lly h as a l ar g e ana lf an, b ut t hi s i sre d uce di nt h e more a d vance df am ili es. T h e w in g venation is g enerall y primitive, but considerable variation is seen within the order. In members of primitive families a t y pical archedict y on is developed to a g reater or lesser d egree; i nt h ose o f a d vance d groups t h e num b er o fb ranc h es o f t h e l ong i tu di na l ve i ns an d t h e num b er o f crossve i ns are great l yre d uce d .T h ea bd omen conta i ns 10 comp l ete segments, wi t h t h e 11t h represente dby t h eep i proct, paraprocts, an dl on g cerc i . In Nemour id ae, h ow - ev e r, the latter are reduced to an unse g mented structure used in copulation. T he esopha g us is ver y lon g , the g izzard rudimentar y , and mid g ut and hind g ut short. T h ere are b etween 20 an d 100 Ma l p i g hi an tu b u l es. In pr i m i t i ve f am ili es t h e centra l nervous system i nc l u d es t h ree t h orac i can d e i g h ta bd om i na l gang li a, b ut i na d vance d groups t he s i xt h to e igh t h a bd om i na lg an gli a f use. T h e trac h ea l s y stem opens to t h e exter i or v i atw o thoracic and ei g ht abdominal spiracles. In males the testes meet in the midline, but their products are carried b y separate vasa deferentia to a pair of seminal vesicles. Usuall y ther e i s a median ejaculatory duct, but in some species the vasa deferentia remain separate unti l t h ey reac h t h eme di an gonopore l ocate db e hi n d t h en i nt h a bd om i na l segment. In f ema l es t h e pano i st i covar i o l es ar i se f rom a common d uct t h at j o i ns t h eov id ucts o f eac h s id e. A spermatheca is usuall y present . L a r va . I n general form larvae resemble adults, except for the absence of wings an d t h e presence, i n most spec i es, o f severa l pa i rs o f g ill s. Pr i m i t i ve l yt h ere are fi ve or s i xpa i r s of a bd om i na l g ill s, b ut i n mem b ers o f more a d vance d groups t h ese are re d uce di n num b er and secondar yg ill structures ma y appear on more anterior re g ions of the bod y (mentum, submentum, neck, thorax, and coxae) or ma y encircle the anus. In addition to g as exchan g e , the gills are important osmoregulators, equipped with chloride-uptake cells, as is also seen in l arva l Ep h emeroptera. In many spec i es t h e l egs are f r i nge d w i t hh a i rs t h at ass i st sw i mm i ng. L ife Histor y and Habit s A d u l t stone fli es are wea kfl yers an d se ld om f oun df ar f rom t h e b an k so f streams or e d ge s of l a k es w h ere t h e y rest, o f ten we ll camou fl a g e d ,onve g etat i on, roc k s, l o g s, etc. Nocturna l species usuall y hide in crevices or amon g ve g etation durin g the da y . Man y stoneflies do no t f eed as adults. Others feed on lichens, acellular al g ae, pollen, bark, and rotten wood . P rior to mating, many Arctoperlaria tap the substrate with the tip of the abdome n (d rumm i ng). Ma l es i n i t i ate t h e d rumm i ng an d v i rg i n f ema l es respon d .T h e d rumm i ng i s spec i es-spec ifi can d serves to b r i ng t h e partners toget h er (Stewart an d Ma k eton, 1990). Matin g usuall y occurs in da y li g ht, on the g round, thou g h a few species are nocturnal. Lar g e n umbers of e gg s are laid, sin g l y or, more often, in batches of 100 or more. In fl y in g specie s f emales hover over the water and dip the abdomen beneath the surface. Brachypterous an d apterous f orms craw l to t h e water’s e d ge, or b e l ow t h e water sur f ace, i nor d er to ov i pos i t. Eggs o f many spec i es d eve l op a dh es i ve propert i es on contact w i t h water. Em b ryon i c d eve l - o pment is usuall y direct, thou g he gg s of some species ma y survive drou g ht conditions i n 149 T HE PLECOPTEROID , BLATTOID, AND O RTH O PTER O ID ORDERS d iapause. A few species are ovoviviparous. Larvae are t y picall y found in streams or lake s w hose bottom is covered with stones under which the y can hide. Development is slow, frequentl y takin g more than a y ear in the lar g er species. Man y molts occur, 33 havin g been recor d e d over a per i o d o f 3 years f or one spec i es. Most stone fl y l arvae are p h ytop h agous , f ee di ng on li c h ens, a l gae, moss, an ddi atoms. Typ i ca ll yt h ese are t h e spec i es t h at a l so f ee d i nt h ea d u l t sta g e. Juven il es o f ot h er spec i es are carn i vorous, li v i n g on ot h er i nsects. T h es e species do not feed as adults. Like that of Odonata and ma y flies, emer g ence of stoneflies is frequentl y hi g hl y s y nchronized. Phylogeny and C lass ifi cat i on Plecoptera, ver y primitive insects sometimes described as “fl y in g Th y sanura,” probabl y h ad their ori g ins in the Lower Permian period from a stem g roup, the plecopteroid assem- b lage, that included the extinct Paraplecoptera and Protoperlaria (Illies, 1965). Some pale - oentomo l og i sts ass i gne d some o f t h e Perm i an f oss il P l ecoptera to recent f am ili es, t h oug h Z w i c k (1981) cons id ere d t hi s i ncorrect, representat i ves o f t h e l atter not appear i n gi nt h e fossil record until the Eocene (or possibl y the Cretaceous). S toneflies traditionall y were placed in two suborders, Filipalpia (Holo g natha) and Seti- palpia (Systellognatha). Illies (1965), however, considered the extremely primitive Souther n Hem i sp h ere f am ili es Eust h en iid ae an d D i amp hi pno id ae su ffi c i ent l y di st i nct f rom t h e re- ma i n i ng F ili pa l p i at h at t h ey s h ou ld b e groupe di n a separate su b or d er, t h e Arc hi per l ar i a. B oth Henni g (1981) and Zwick (1981) ar g ued that Illies’ arran g ement was not soundl y b ased, and Zwick (1980, 1981, 2000), whose classification is followed here, p ro p osed that t he stoneflies could be divided into an exclusively Southern Hemisphere group (suborder Antarctoper l ar i a) an d a pre d om i nant l y Nort h ern Hem i sp h ere group (su b or d er Arctoper - l ar i a), t h e separat i on an d su b sequent evo l ut i on o f t h e two groups resu l t i ng f rom t h e b rea k u p of the Pan g ean landmass (into Laurasia and Gondwanaland) durin g the Jurassic period. A few Arctoperlaria occur in the Southern Hemisphere, presumabl y as a result of secondar y invasions. Figure 7.1 provides a suggested phylogeny for the order . S uborder Antarctoperlari a I n Zwick’s classification this suborder includes the su p erfamilies Eusthenioidea (fam - ilies EUSTHENIIDAE and DIAMPHIPNOIDAE) and Gripopter yg oidea (AUSTROPER - L IDAE and GRIPOPTERYGIDAE). Illies (1965) considered members of the small family E ust h en iid ae, w hi c hi s restr i cte d to eastern Austra li a, New Zea l an d ,an d C hil e, to represent th e protot y pe o f p l ecopteran or g an i zat i on. T h e y are l ar g e, co l or f u li nsects h av i n g w i n gs w ith numerous crossveins and an anal fan in the hind win g with ei g ht or nine anal veins. L arvae are carnivorous and have four to six pairs of abdominal g ills. The Gripopter yg ida e is a large family (about 150 species) mostly found in Australia, with a few species in New Z ea l an d an d Sout h Amer i ca. T h ea d u l ts are most l y d u ll i nco l or; t h e l arvae, w hi c h ar e s l u ggi s h an d t y p i ca lly f oun d un d er roc k san dd e b r i s i n f ast-mov i n g water, h aveatu f to f g ills around the anus. Larvae of a few species are terrestrial and lack g ills (Zwick, 2000). S uborder Arcto p erlari a Z w i c k (2000) di v id e d t hi ssu b or d er i nto t h e i n f raor d ers Syste ll ognat h aan d Eu- h olo g natha. The former contains the superfamilies Perloidea (families PERLODIDAE, 150 C H A PTER 7 F I GU RE 7.1. P roposed ph y lo g en y of the Plecoptera. [Modified from P. Zwick, 1980, Plecoptera (Sternflie g en) , i n : H andbuch der Zoolo g i e , V ol. IV, Insecta Lfg. VV 26 :1–115. By permission of Walter de Gruyter and Co. ] P ERLIDAE, and CHLOROPERLIDAE) and Pteronarcyoidea (families PTERONARCYI - DAE an d PELTOPERLIDAE). Inc l u d e di nt h eEu h o l ognat h a i sas i ng l e super f am il y Nemouro id ea (f am ili es TAENIOPTERYGIDAE, NOTONEMOURIDAE, NEMOURI - D AE, CAPNIIDAE, and LEUCTRIDAE) and the ver y small famil y SCOPURIDAE. The S copuridae forms the sister g roup to the nemuroids . T he Pteronarc y idae (Fi g ure 7.2A) is a small, primitive famil y whose members includ e t h e l argest stone fli es an dh ave w i ngs w i t h numerous crossve i ns. It i spr i mar il y a Nort h A mer i can group t h at h as i nva d e d eastern As i a i nre l at i ve l y recent t i mes. T h e h er bi vorous o r d etr i t i vorous l arvae are f oun di nme di um- to l ar g e-s i ze d r i vers. Anot h er sma ll f am ily , the Peltoperlidae, has a similar distribution to the Pteronarc y idae, and the larvae, which are somewhat cockroachlike in a pp earance, also feed on p lant material or detritus. The P er l o did ae (F i gure 7.2B,C) i sa l arge h o l arct i c group ( > 200 s p ec i es) o f me di um-s i ze d stone fli es w h ose l arvae are carn i vorous, l ac k g ill s, an d are typ i ca ll y f oun di ns l ow l y fl ow i n g rivers. The Perlidae is the lar g est famil y in the order with some 3 5 0 species. Thou gh primaril y a holarctic-oriental g roup, the famil y has representatives in South America and 151 THE PLE CO PTER O ID , BLATTOID, AND O RTH O PTER O ID ORDER S F IGURE 7.2. P l ecoptera. (A) Pteronarcys ca l ifornic a (Pteronarcy id ae) a d u l t; (B ) Isoper l a confus a (Per l o did ae) a d u l t; an d( C ) I. con f usa l arva. [A, f rom A. R. Gau fi n, W. E. R i c k er,M.M i ner,P.M il am, an d R. A. Ha y es, 1972 , T he stoneflies (Plecoptera) of Montana , Tr ans. Am. Entomol. Soc. rr 98 :1–161. By permission of the American E ntomological Society. B, C, from T. H. Frison, 1935, The stoneflies, or Plecoptera, of Illinois, B u ll .I ll . Nat. Hist . S ur v . 20 (4). B y perm i ss i on o f t h eI lli no i s Natura l H i stor y Surve y . ] A f r i ca. T h at t hi s i s a rat h er a d vance d group i s suggeste db yt h ere d uce d g l ossae, re d uce d fi rst abdominal sternite, the fusion of the first two abdominal g an g lia with that of th e metathorax, and the absence of abdominal g ills in larvae that are g enerall y carnivorous. Containing more than 110 species, the holarctic family Chloroperlidae is considered to be th e most spec i a li ze d o f t h esu b or d er b yv i rtue o f t h ere d uce db o d ys i ze an d w i ng venat i o n (espec i a ll yt h ea b sence o f t h e ana lf an i nt h e hi n d w i ng) an d t h e comp l ex ma l e repro d uct i ve s y stem. A d u l ts are o f ten g reen ( h ence t h e f am ily name); l arvae o f most spec i es are pre d ator s t hou g h a few are detritivores or herbivores, the y lack g ills, and ma y show adaptations for b urrowin g in the substrate of the streams and small rivers where the y are found. Taen i opteryg id ae const i tute t h e most pr i m i t i ve f am il yo f Nemouro id ea as i s i n di cate d b yt h e comparat i ve l yr i c h w i ng venat i on, l arge ana ll o b e i nt h e hi n d w i ng, an dfi ve-ors i x - part cerc i .A d u l ts o f t hi s h o l arct i c g roup, compr i s i n g a b out 70 spec i es, are common ly k nown as winter stoneflies because of their habit of emer g in g between Januar y and April. Som e 152 C H A PTER 7 F I GU RE 7.3 . Pl eco p tera. (A) C apnia nan a (Ca p n iid ae) a d u l t; (B) Nemoura fl exura ( Nemour id ae ) a d u l t; an d( C ) N . fl exura larva. [From A. R. Gaufin, W. E. Ricker, M. Miner, P. Milam, and R. A. Hayes, 1972, The stoneflies (Ple- c optera) o f Montana , Tr ans. Am. Entomol. Soc. r r 98 : 1–1 6 1. By permission of the American Entomological Society.] a d u l ts f ee d on po ll en. Larvae, common l y f oun di n l arge streams an d r i vers, are h er bi vores o r detritivores. In Capniidae (Fi g ure 7.3A), a holarctic famil y of about 200 species, adults are g enerall y small, their win g s have few cross veins, and the size of the anal fan in the hind w ing is reduced. The cerci, however, are long. Like Taeniopterygidae, capniids may emerge d ur i ng t h ew i nter. T h e genera ll y d etr i t i vorous l arvae are most l y f oun di n sma ll r i vers an d streams, t h oug h a f ew spec i es occur i na l p i ne l a k es. Leuctr id ae, w hi c h compr i se a h o l arct ic f am ily o f a b out 170 spec i es, are reco g n i ze dby t h e i ra bili t y to ro ll t h e i rw i n g s aroun d t he abdomen. The small anal area of the hind win g s, the undivided cerci, and the specialized m ale g enitalia su gg est that this is an advanced famil y .T y picall y , the larvae are found in small m ounta i n streams w h ere t h ey f ee d on d etr i tus. W i t h a b out 340 spec i es, t h e h o l arct i c f am il y Nemour id ae (F i gure 7.3B,C) ran k snexttot h e Per lid ae i n terms o f s i ze. T h oug h t h ew i ng v enat i on i spr i m i t i ve, t h e g enera lly sma ll s i ze o f t h ea d u l ts, t h e highly mo difi e d cerc i an d g enitalia of the male, and the nerve cord with onl y five abdominal g an g lia (due to fusion o f 153 THE PLE CO PTER O ID , BLATTOID, AND O RTH O PTER O ID ORDERS posterior ones) make this perhaps the most advanced famil y in this g roup. Larvae are found in fast-movin g streams, often with rock y substrates, where the y feed on detritus or, rarel y, g rowin g plants and al g ae. The famil y Notonemouridae (about 60 species in Mada g ascar , Sout h A f r i ca, Sout h Amer i ca, Austra li a, an d New Zea l an d ) i s lik e l y a parap h y l et i c group . Z w i c k (1981) suggeste d ,ont h e b as i so f diff erences i n gen i ta li aan di nterna l structure, t hat the g roup arose as a result of several independent invasions from ori g inall y Norther n Hemisphere stock. Larvae are found in a variet y of habitats and are detritivores . L i t e r a t u r e H itchcock (1974), Hynes (1976), and Harper and Stewart (1984) provide much infor- mat i on on t h e genera lbi o l ogy o f stone fli es. T h ep h y l ogeny o f t h eor d er i s di scusse dby I llies (196 5 ) and Zwick (1980, 1981, 2000). Keys for identification are provided by Hynes (19 6 7) [British species], Harper and Stewart (1984) and Stewart and Stark (1988) [Nort h American g enera], and Theischin g er (1991) [Australian families]. Har p er, P. P., and Stewart, K. W., 1984, Pleco p tera, in : A n Introduction to the A q uatic Insects of North America , 2nd ed. (R. W. Merritt and K. W. Cummins, eds.), Kendall/Hunt, Dubuque, IA. Henn ig , W., 1981, I nsect P h y l ogen y ,W iley, New York. WW Hitchcock, S. W., 1974, Guide to the insects of Connecticut. Part VII. The Pleco p tera or stoneflies of Connecticut , Conn. State Geol. Nat. Hist. Surv. Bull . 1 0 7 : 262 pp . H y nes, H. B. N., 19 6 7,Ake y to the adults and n y mphs of British stoneflies (Plecoptera) (2nd ed.) , F . W . B io l . A ssoc. S ci. Publ. 17 : 86 pp . Hynes, H. B. N., 1976, Biology of Plecoptera , A nnu. Rev. Entomol . 21 : 1 35 –1 5 4 . Illies, J., 196 5 ,Ph y lo g en y and zoo g eo g raph y of the Plecoptera , A nnu. Rev. Entomo l. 10 :11 7– 140 . S tewart, K. W., and Maketon, M., 1990, Intraspecific variation and information content of drummin g in thre e P lecoptera species, in : Mayflies and Stoneflies: Li f e Histories and Biolog y (I. C. Campbell, ed.), Kluwer , D or d rec h t. S tewart , K. W. , and Stark , B. P. , 1988 , N ymphs o f North American Stonefly Genera (Plecoptera), Thomas Sa y F oundation, Lanham, MD. T h e i sc hi n g er, G., 1991, P l ecoptera, i n : Th e Insects of Austra l i a ,2n d e d ., Vo l . I (CSIRO, e d .), Me lb ourne Un i vers i t y P ress , Carlton , Victoria . Z wick, P., 1980, Plecoptera (Sternfliegen), in : Handbuch der Zoolo g i e , V ol. IV, Insecta Lfg. V V 2 6 : 1–115, de Gruyter, B er li n. Z wick, P., 1981, Plecoptera, revisionar y notes, in : I nsect Ph y lo g en y ( W. Henni g ), Wile y , New York . Z wick, P., 2000, Phylogenetic system and zoogeography of the Plecoptera , A nnu. Rev. Entomol . 4 5 : 709–746. 3 . Emb i o p ter a S ynonyms: Em bi o d ea, Em biidi na, Em biida C ommon names: w e b s pi nners, em biid s, f oots p inner s E longate, small, or moderately sized insects that live gregariously in silk tunnels; head wit h filif orm antennae, compoun d e y es, an d man dib u l ate mout h parts b ut l ac ki n g oce lli ;ma l es o f almost all species with two pairs of nearl y identical win g s in which radial vein is thickened, f emales apterous, tarsi three-segmented and basal segment of fore tarsus greatly enlarged; cerci t wo-se g mente d an d usua lly as y mmetr i ca li nma l es. T h eEm bi optera (F i gure 7.4) are most l y con fi ne d to t h e l arger l an d masses i n trop i ca l or subtropical areas of the world, thou g h a few have found their wa y even to oceanic islands. Althou g h fewer than 200 species have been described, includin g 13 species from Nort h 154 C H A PTER 7 F IGURE 7.4. E m b ia major ( Em bi optera). (A) Ma l e; an d (B) f ema l e. [From A. D. Imms, 1913, O n E m b ia major n . sp. From t h eH i ma l a y as , Tr ans. Linn. Soc. Zool r r . 11 : 167–19 5 .B y permission of Blackwell Publishin g Ltd. ] A merica and 65 from Australia, Ross (1970, 1991) suggested that this figure may represent o n l y 10% o f t h ewor ld tota l . S tructur e As a g roup, the Embioptera are of remarkabl y uniform structure, a feature related to the widespread similarit y of the tunnels in which the y live. Webspinners are soft-bodied i nsects t h at fl yon l y wea kl y or not at a ll .T h e prognat h ous h ea db ears filif orm antennae, c ompoun d eyes (o f ten l arge an d kid ney-s h ape di nma l es, sma ll i n f ema l es), an d man dib u l ate m out h parts. In ma l es t h e man dibl es are usua lly fl attene d an d e l on g ate. Oce lli are a b sent . No trace of win g s can be seen in females; males ma y be apterous, brach y pterous, or full y w in g ed. In the latter the fore and hind win g s are ver y similar. The radius is thickened ; t h eot h er ve i ns are re d uce d .T h ew i ngs are fl ex ibl ean d a bl eto f o ld at any po i nt. T his f ac ili tates b ac k war d movement a l ong t h e tunne l s. For fli g h tt h ew i ngs are ma d e more r i g id b y pump i ng bl oo di nto t h era di us. T h e f ore l egs are stout, an d t h e b asa l tarsa l segment is swollen to accommodate the silk g lands, which number about 200. Ducts from the g lands c arr y the product to the exterior via hairlike e j ectors. The hind femur is also enlar g ed t o c ontain a large tibial depressor muscle. This is correlated with the ability to run backward wi t h great spee d .T h ere are 10 o b v i ous a bd om i na l segments. T h e cerc i are two-segmente d an d tact il e, serv i ng as cau d a l “eyes” w h en t h e i nsect i s runn i ng b ac k war d .Inma l es t h e cerc i are usuall y as y mmetrical. . T he internal structure is g eneralized. The g ut is strai g ht, and 20–30 Malpi g hian tubule s op en into it. The ventral nerve cord is p aired and includes three thoracic and seven abdominal gang li a. Eac h ovary cons i sts o ffi ve pano i st i covar i o l es t h at are connecte d at i nterva l sw i t h t h eov id uct. A spermat h eca i s present. T h e fi ve test i s f o lli c l es on eac h s id e are a l so arrange d ser i a lly a l on g t h evas d e f erens, w hi c h swe ll s prox i ma lly i nto a sem i na l ves i c l e. Two pa i rs o f accessor yg lands occur in males. 155 THE PLE CO PTER O ID , BLATTOID, AND O RTH O PTER O ID ORDER S Life Histor y and Habits Bot h a d u l tan dj uven il eEm bi optera can pro d uce s ilk en tunne l st h at are j ust w id e enoug h t o permit the animals to move forward or backward alon g them. Generall y , man y embiids are found associated to g ether in a “nest” of interconnected tubes. It must be emphasized, h owever, that this g re g arious behavior is in no wa y social; that is, there is no caste s y ste m or di v i s i on o fl a b or. In h um id reg i ons an ent i re nest may b e expose d , b ut i n d r i er parts o f th ewor ld i t i s usua ll y part i a ll ysu b terranean as a protect i on aga i nst d es i ccat i on an dfi re . N ests are constructe di nt h e i mme di ate v i c i n i t y o f a f oo d source, an d tunne l so f ten exten d d irectl y into this. Embiids are ph y topha g ous, with dead g rass and leaves, lichens, moss, an d b ark constitutin g the main food. Earl y workers believed that males mi g ht be carnivorou s b ecause o f t h e rat h er di st i nct man dibl es. It i snow k nown, h owever, t h at t h e structure o f t h e l atter i s corre l ate d w i t h t h e i r use i n grasp i ng t h e f ema l e’s h ea dd ur i ng copu l at i on, an d , i n man y spec i es, mature ma l es d o not f ee d . At y pical nest contains a few mature females and their developin gy oun g . Matur e males are g enerall y short-lived and, in some species, are eaten b y the female after matin g . Part h enogenes i s pro b a bl y occurs i n some spec i es. Eggs are l a id i n a tunne l an d guar d e d b yt h e f ema l e. Parenta l care i s exten d e d to t h e young l arvae, b ut t h ese soon pro d uce t h e ir ow n t unne l s i nw hi c h to d eve l op. New co l on i es are f orme di nt h ev i c i n i t y o f t h eo ld ones, and it is durin g this short mi g ration to new sites that embiids are especiall y vulnerable. The absence of win g s in females has more or less restricted the distribution of the Embioptera to t he major land masses, though some species, perhaps transported by commerce, are found on remote Pac ifi c i s l an d s. Phylogeny and C lass ifi cat i on The fossil record of Embioptera is poor. Some authors (e. g ., Henni g , 1981; Kukalov´a - Peck, 1991) believe that it is a ver y ancient insect order with a fossil record that extends to t he Lower Permian p eriod. It is claimed that these fossil remnants have a combination o f pr i m i t i ve (e.g., w i ngs i n f ema l es, mu l t i segmente d cerc i ,an d s h ort ov i pos i tor) an d a d vance d c h aracters (e.g., asymmetr i c cerc ii nma l es an d re d uce d w i ng venat i on). However, t he “em biid ” nature o f t h ese Perm i an f ra g ments i s di spute dby ot h er wor k ers (e. g ., Carpenter, 1992; Rasnits y n and Quicke, 2002), so that g enuine embiopteran fossils do not appear b efore the Late Cretaceous-Earl y Eocene. The order is clearl y orthopteroid but its stron g l y apomorp hi cc h aracter h as hi n d ere d c l ar ifi cat i on o fi ts pos i t i on w i t hi nt h e l arger group . R e l at i ons hi ps w i t h P l ecoptera, Dermaptera, P h asm id a, an d Zoraptera h ave b een suggeste d by var i ous aut h ors (see Kr i stensen, 1991). Because of the neotenous nature of females, identification and classification usin g morpholo g ical characters can be carried out with certaint y onl y b y examinin g mature males . R oss (1970) recognized eight families of living Embioptera, but it is not yet possible to d raw many conc l us i ons regar di ng t h e i rp h y l ogenet i cre l at i ons hi ps b ecause o f t h e genera l structura l un if orm i ty o f t h eor d er an d t h e amount o f para ll e l evo l ut i on t h at h as ta k en p l ace amon g families. The northern South American and West Indian famil y CLOTHODIDAE is t he most primitive g roup. In this famil y , to which certain Miocene fossils are assi g ned, the cerci of the male are symmetrical and comprise two smooth segments. The largest family, E MBIIDAE, is a rather heterogeneous group of Old and New World forms. Szumik’s (1996) c l a di st i c ana l ys i ss h owe d t h at, as present l y const i tute d ,t h eEm biid ae i s a parap h y l et ic gro u p. 156 C H A PTER 7 Another lar g e and likel y paraph y letic famil y is the OLIGOTOMIDAE, a rather prim - i tive g roup with representatives in Asia, Australia, southern Europe, and possibl y Eas t A frica. Three s p ecies of O ligotoma have been introduced accidentall y into the United States . O t h er f am ili es are t h e AUSTRALEMBIIDAE (restr i cte d to eastern Austra li aan d Tasman i a) , NOTOLIGOTOMIDAE (Sout h east As i aan d Austra li a), EMBONYCHIDAE (East As i a) , TERATEMBIIDAE ( South America and southern United States ) , and ANISEMBIIDA E ( Central America and southern United States ). L iterature Th e bi o l ogy o f t h eEm bi optera i s d ea l tw i t hb y Ross (1970, 1991) w h oa l so (1984, 1 991) h as d escr ib e d an d prov id e dk eys f or t h e id ent ifi cat i on o f Nort h Amer i can genera and Australian families, respectivel y . Ross (1970) and Szumik (199 6 ) have discussed the ev olution and classification of the order . C ar p enter, F. M., 1992 , Tr eatise on Invertebrate Paleontology. Part R. Arthropoda rr 4 , V o l s. 3 an d 4 ( Superc l ass H exa p oda ) , University of Kansas, Lawrence. H enn i g, W., 1981, Insect P h y l ogeny ,W iley, New York. WW Kr i stensen, N. P., 1991, P hyl o g en y o f extant h exapo d s, i n : T h e Insects o f Austra l ia , 2n d e d ., Vo l .I ( CSIRO, e d . ) , Melbourne University Press, Carlton, Victoria. K u k a l ov´a-Pec k , J., 1991, Foss il hi story an d t h eevo l ut i on o fh exapo d structures, i n : Th e Insects of Austra l ia , 2n d e d ., Vo l . I (CSIRO, e d .), Me lb ourne Un i vers i t y Press, Car l ton, V i ctor i a. Rasnitsyn, A. P., and Quicke, D. L. J. (eds.), 2002 , H istory o f Insects , Kluwer, Dordrecht. Ross, E. S., 1970, B i os y stemat i cs o f t h eEm bi optera, Annu. Rev. Entomo l . 15 : 1 5 7–172 . Ross, E. S., 1984, A s y nops i so f t h eEm biidi na o f t h eUn i te d States, Proc. Entomo l . S oc. W as h. 86 :82– 9 3. Ross, E. S., 1991, Embioptera, in: The Insects o f Australia , 2nd ed., Vol. I (CSIRO, ed.), Melbourne Universit y Press , Car l ton , V i ctor i a. Szumik, C. A., 1996, The hi g her classification of the order Embioptera: A cladistic anal y sis, C ladistic s 12 :41– 6 4. 4 . Dictyopter a S ynonyms : D i ctuoptera, Oot h ecar i a, B l att if orm i a , C ommon names: cock r oaches a n d Blatto p teriformi a m a n t i ds Small to very large terrestrial insects of varied form; head hypognathous with filiform, multiseg - m ente d antennae, man dib u l ate mout h parts an d we ll d eve l ope d compoun d eyes, oce lli present ( Manto d ea) or usua lly a b sent (B l atto d ea); pronotum l ar g ean ddi sc lik e(B l atto d ea) or e l on g ate ( most Mantodea), legs with five-segmented tarsi, fore wings modified as tegmina, brachyptery, a n d apter y common; ov i pos i tor re d uce d an d hidd en, ma l e g en i ta li a comp l ex an d concea l e d , cerc i fairly short but multisegmented. T his mainl y tropical to subtropical order contains some 5500 described species that f all into two clearl y defined suborders, Blattodea (cockroaches), with at least 3500 species (i nc l u di ng a b out 70 i n Nort h Amer i ca, more t h an 400 i n Austra li a, an d 9 i nBr i ta i n) , an d Manto d ea (mant id s), a pre d om i nant l yO ld Wor ld group o f a b out 2000 spec i es ( i n - c ludin g about 900 in Africa and 5 30 in Asia). About 20 mantid species occur in Nort h A merica and 1 6 0 in Australia. Several s p ecies of cockroaches are im p ortant cosmo p olitan p es t s . [...]... comprehensive coverage of the group Other introductions to the biology of the order are given by Wilson (1 971 ), Howse (1 970 ), Harris (1 971 ), and Pearce (19 97) Termite phylogeny is discussed by McKittrick (1964), Krishna (1 970 ), 171 THE PLECOPTEROID, BLATTOID, AND ORTHOPTEROID ORDERS 172 CHAPTER 7 FIGURE 7. 11 Castes of the higher termite, Amitermes hastatus (Termitidae) (A) Worker; (B) soldier; (C) physogastric... By permission of Masson, e r ´ Vol Paris.] 177 THE PLECOPTEROID, BLATTOID, AND ORTHOPTEROID ORDERS 178 CHAPTER 7 includes about 10 species of Hemimerus (Figure 7. 14C), all of which are epizoic on African giant rats of the genus Cricetomys Some authors consider that the features by which Arixenia (Popham, 1965) and Hemimerus (Klass, 2001) differ from free-living forms are simply adaptations to their... The six legs are similar in structure, 173 THE PLECOPTEROID, BLATTOID, AND ORTHOPTEROID ORDERS 174 CHAPTER 7 FIGURE 7. 13 Grylloblatta campodeiformis (Grylloblattodea) [From E M Walker, 1914, A new species of Orthoptera, forming a new genus and family, Can Entomol 46:93–99 By permission of the Entomological Society of Canada.] each with a large coxa and a five-segmented tarsus The abdomen has 10 obvious... (19 37) , Mills and Pepper (19 37) , and various authors in Ando (1982) Kamp (1 979 ) and other authors in Ando (1982) discuss the taxonomy and distribution, and provide a bibliography, of the group Giles (1963), Kamp (1 973 ), and Storozhenko (19 97, 2002) discuss their affinities with other orthopteroids Storozhenko (1988) provides a key to living species Ando, H (ed.), 1982, Biology of the Notoptera, Kashiyo-Insatsu,... three-segmented tarsi; abdomen with unsegmented forcepslike cerci, ovipositor of females reduced or absent 175 THE PLECOPTEROID, BLATTOID, AND ORTHOPTEROID ORDERS 176 CHAPTER 7 Dermaptera are found in all but the polar regions of the world, though they are most common in the warmer parts Of the approximately 1800 described species, only about 25 occur in North America, 45 in Europe (including 7 in... 48:1 87 193 Grass´ , P P., 1982–1986, Termitologia, V e Vols I–III, Masson, Paris Harris, W V., 1969, Termites as Pests of Crops and Trees, Commonwealth Institute of Entomology, London Harris, W V., 1 971 , Termites—Their Recognition and Control (2nd ed.), Longmans-Green, London Helfer, J R., 19 87, How to Know the Grasshoppers, Crickets, Cockroaches and Their Allies, Dover, New York Hickin, N E., 1 971 ,... CHAPTER 7 into many more families (e.g., Bradley and Galil, 1 977 ; Kevan, 1982) Key (1991) supported an earlier suggestion that the aberrant North American genus Timema, species of which have a very short (1–3 cm long) body, three-segmented tarsi, and the metanoturn not fused with the first abdominal segment, be placed in its own family TIMEMATIDAE Literature Phasmid biology is reviewed by Clark (1 974 ),... by Clark (1 974 ), Bedford (1 978 ), Kevan (1982), and Key (1991) The taxonomy and distribution of the order are discussed by G¨ nther (1953) and u Kevan (1982), and phylogeny by Sharov (1968), Hennig (1981), and Mazzini and Scali (19 87) Helfer (19 87) provides a key to the North American forms, while Bradley and Galil (1 977 ) have keys to subfamilies and tribes Bedford, G E., 1 978 , Biology and ecology of... and Galil, B S., 1 977 , The taxonomic arrangement of the Phasmatodea with keys to the subfamilies and tribes, Proc Entomol Soc Wash 79 : 176 –208 Clark, J T., 1 974 , Stick and Leaf Insects, Sherlock, Winchester, Hants ¨ Gunther, K., 1953, Uber die taxonomische G1eiderung und die geographische Verbreitung der Insektenordnung ¨ der Phasmatodea, Beitr Entomol 3:541–563 Helfer, J R., 19 87, How to Know the Grasshoppers,... 90:212–219 Krishna, K., 1 970 , Taxonomy, phylogeny and distribution, in: Biology of Termites, V II (K Krishna and F M Vol Weesner, eds.), Academic Press, New York Krishna, K., and Weesner, F M (eds.), 1969, 1 970 , Biology of Termites, V I and II, Academic Press, New York Vols McKittrick, F A., 1964, Evolutionary studies of cockroaches, Mem Cornell Univ Agri Exp Stn 389: 177 pp Pearce, M J., 19 97, Termites: Biology . insects , C an. J . Earth S ci . 6 : 115 9 –1 177 . Grandcolas, P., 1996, The phylogeny of cockroach families: A cladistic appraisal of morpho-anatomical data , C an. J . Z oo l . 74 : 5 08– 5 27. Grandcolas,. importance of cockroaches, S mithson. M isc.Co ll ect . 134 (10) : 14 7 pp. R oth , L. M. , and Willis , E. R. , 19 6 0 , The biotic associations of cockroaches , S mit h son. Misc. C o ll ect. 141 ( 4422 ): 4 7 0 pp. Sc h a l ,. s p ecies of Periplaneta, P. ameri- can a (the American cockroach) (Figure 7. 6B) , P. au s tra l a s iae (t h e Austra li an coc k roac h ) , P . f u l iginosa ( t h e smo k ey- b rown coc k roac h ),