ed on bas A oplex A MG ¥ * < A CLASSIFICATION OF THE LEPIDOPTERA BASED ON CHARACTERS OF THE PUPA EDNA MOSHER B.S. A. Cornell University, 1908. M. S. University of Illinois, 1913. THESIS Submitted in Partial Fulfillment of the Requirements for the Degree of DOCTOR OF PHILOSOPHY IN ENTOMOLOGY THE GRADUATE SCHOOL UNIVERSITY OF ILLINOIS 1915 ae Se rel ema ene} , - ba 7 ee ~ ‘ ‘. ‘ + 2 am < a .) pa oe —— = AA » bibwANY ee (vers ah THE i gate le Pan OP ie tite tli Late oe eS ae F i Pie BS Bes | p>? eee a “ - > + * Pia | bey | ‘ e Ps 7 ¢") % 7 } : "yt we ia ‘ ba ; 7 7 J I “ij e ¥ 48 MeV | UNIVERSITY OF ILLINOIS BaueO May 15, THIS IS TO CERTIFY THAT THE THESIS PREPARED UNDER MY SUPERVISION BY Bement MOOT. Bepighg 5 O65 et ENTITLED A Classification of the Lepidoptera based on. mepmarseners of the: pupa IS APPROVED BY ME AS FULFILLING THIS PART OF THE REQUIREMENTS FOR THE SeeeeeOr Doctor of Philosophy. Instrucfor in Charge HEAD OF DEPARTMENT OF... C@@¢t¥-e 11.0 Ca Fag SS + * e . : ee IS a mmm gram : oz \ 4 y , \ ie YW or f a Bey Cibgane Rmancioe i pelt THe re U ‘ ‘ i a a ve i OF aaawas VALS AI Bes IE $e Introduction 4 “ Changes Preceding Pupation | III. External Morphology Betv. classification Analytical table of superfamilies A. Pupae with functional mandibles Superfamily Micropterygoidea AA. Pupae without functional mandibles B. Generalized pupae without maxillary palpi Superfamily Hepialoidea Superfamily Cossoidea Superfamily Eucleoidea BB. Generalized pupae with maxillary palpi Superfamily weneol cca). Superfamily Aegerioidea Superfamily Tortricoidea . Superfamily Gracilarioidea C. Specialized pupae with piljifers Superfamily Pyralidoidea Superfamily Papilionioidea CC. Specialized pupae without piljifers Superfamily Yponomeutoidea Superfamily Gelechioidea Superfamily Noctuoidea Superfamily Bombycoidea Ke bea ticroaanh ; ae ae co Leta 2 2088 Sb Qe ie As | ide 4 ts i wis oe ae cee so) aire OF, 20 ae Se ae 2 ny At 3 rity. Ou Eee ae Sis aes = (2 bie ts ear oa F e a od " a 2h2'C Oeias v4 ie 9 if on LSaw- sae, Dees Takes EtAtT ¥..o6 TAG afc THT SEB : lt 2 Seok Se Se Cee .. POSte2 papere th eka dl | # Oi ae F 29 dee bite y 1 ieee pan - ae =i ke. ae koh oe ia pala » haf SOgySOG | FPR Aaa Superfamily Notodontoidea . Superfamily Sphingoidea . Superfamily Saturnioidea . ', Phylogeny .. I. Bibliography I. Acknowledgments emeptenetion of Plates... . +... edie Pec e cece cc ec eee eeee 186 Oia. Ey Deu Oe 2 tub Peat i Be erg kee) ee hae wa o>) MA see es is | I. INTRODUCTION It is within comparatively recent times that the immature stages of insects have been considered of any taxonomic value. The economic entomologist early realized the value of being able to recognize the immature stages, for in many orders of insects the larval stages alone were responsible for many ravages upon crops and orchards. Still the matter was not taken up by the systematists and the workers in the field of economic entomology contented themselves by rearing the adult to determine the species, and then describing, perhaps all the stages, or, more probably the larval and adult stages as being those of econ- omic importance. Now-a-days we are beginning to see that it is impossible to construct an adequate classification of any group of insects unless we use every bit of information obtainable on the life history and habits of the group in question. It is possible to multiply instances of the value of the larval stages in classification, so that one scarcely needs to cite examples; but the pupae have been less frequently used. There are, however, instances of insects in their pupal stage possessing the only good taxonomic characters available. Such instances are found among the nematocerous Diptera, particularly in the family Chironomidae. Scudder '89 was the first to attempt a classification of lepidopterous pupae, but his keys to the chrysalids were based, not on structural characters, out on the various projections from the body, the cuticular appendages, the coloration and the mode of suspension. i) ih ALY 7 dt | Tg } IO Yi a Y ead De ~ e ‘ba - = * aff OL e e ay ig a at hk cae if om oo ae i=» =~ ye 4 ft SS . j a Payot t “e ‘ bean ' rs} a - .~? EEX) iy oe D r 4 et yas to hetabLenoo meed evel etovent ite el ai _ a? ‘od es gepete tlote bast Leave rei vd witha gamit nos vy! Lev LhemBgmiod ated by eh Ht ! 3 Ls w Ay i¢ fecifaer vyizae te Psofomosne 0 LnosOge ait efoteor am aay. aro fe gerate Lewret edt trom edt tied. .ebmuiMone Bae sel Ny tag ml), tot? ot of evestiow say eee ote vemeteys (iy ‘ruts odt aoiiveen ¢e Ser leemetd betadtgag: | Me - i d+ (fle doedtec. , patdtxobeR Aaa nee ete C ae ‘ OF stniitasd 6te 68 9y2h-8 Wot | e0me noltteolttiesals riéeuogor as rovtdecte a do noitaerotal, To el¢ ynowen~ nest, ov se plane sitcoup of geots say To etidad Sma costes seodstent viel inom of eldigsod. se Fah | fadt Oe y~ockheottdgsele as soll oval soud aves paged sat tid :se Logie 7 rieds al efo@esmt io pocead wet tevewort oo sitave wretoexato olmoxoxad Boog Vlag eat ant stedg lt Bt cotensx edd gnome Davot ote ae 1} gale 6S! tenbves Smbhmongm ko cial d tod ,saqe¢ aporetgobiqel to noiteott ae etoerado Lesvtourte no fon voer botebtfevat Bete 2 son emtt- tony eas LE » 4 i. + ’ ‘ + . & ma, rs 2 a aati {xttosen 6s “et as. aaa ee? abate emoe words: bs bis. . T.CHANGES PRECEDING PUPATION The person who begins the study of pupae with the pre- conceived notion that the pupal stage is an interpolated one in the insect's life and that a pupa bears little or no resemblance to either larva or adult, will probably find abundant cause for a change of mind before his study is completed. In the case of Lepidoptera one is apt to think that no similarities could possi- bly exist between any of the three stages of the insect's develop- ment after it leaves the egg. After careful study, however, one is surprised with the resemblance between the stages, for it is of the highest importance in the study of any group to be able to homologize larval, pupal and imaginal characters. This has been done to some extent in certain orders of insects, particularly in those groups where the resemblance between the larva and adult is more striking than in the case cf the Lepidoptera. Attempts have been made, however, even in this order, to homologize the mouth- parts of the larva and adult, and some of the larval structures have been homologized with certain structures in the pupa, but apparently the idea that all three stages should be studied has been left for other minds to entertain. The first striking difference between larva and pupa is that of size. This difference is easily explained by the great difference in the size of the alimentary canal. Another striking difference is the apparent absence of legs and prolegs. As will be shown later the legs are always present, but folded and not in use, while the scars of all the prolegs remain to show their location and are very easily identified in the majority of cases. i, yao ' +0 ie, ‘at j HOTTAIUA ONTARIANS saoell } | ap -aiq sit dtiv eaquy wh vouta ad? antged, ody doareg oat oe > hetelogzustat as ef epets) Lequq eee tadt gotten | | [deseet of t@ oltekt exaed aqud 2 aay bat ettt. a't t saved teebauds Boks ¢igads aty (Liw ,- vy =f e Sse wv “42 a a a ba a choptera and Coleoptera, the appendages, as well as the body, are encased in a pupal skin, are free from each other and the body, and together with the body segments possess considerable freedom of motion. This does not mean that they have any power of loco- motion; on the contrary they are quite helpless and for this rea- son are frequently, in common with the great majority of pupae, protected by some sort of a cocoon, or earthen cell. The lepi- dopterous genus Micropteryx, which is supposed by many to be the most generalized of its order, retains freedom of motion of all the appendages and in all but the fixed caudal segments of the abdomen. This freedom of motion is gradually lost in lepidopterous pupae as specialization advances, and the adult appendages are not as fully developed when the pupal stage is assumed, although the cases of the appendages of the pupa are fully formed. Specializa- tion in the pupa consists also in the hardening of the exposed parts of the cuticle, through the deposition of chitin, and the soldering of the appendages to each other and to the body of the pupa. In the generalized families the appendages are soldered to each other but often remain free from the body surface; later the wings become attached to the body surface, but any parts of the antennae, legs or maxillae extending beyond their caudal margins remain free. The tips of these appendages are provided for in various ways in the higher families, but are always found soldered firmly to the surface of the body of the pupa. Proceeding hand in hand with the soldering down of the appendages is the loss of motion in the abdominal segments. Among certain families there is motion between all of the adjacent segments. There is, however, he ty lion ie oy aL Pepe yt eit 4 ao bet. es oa a4; . rang .wbhod edt bas tsdto sose mors earty oi vers fogia a t sobeett efdetebleasd enpeand etitsagee Yous see mele todd sf to tevoc yre evad yea? Jemp cise sen eaeb oidT «© otét tot bas seelehad ot inp ete COUR) Buaveaee edt ao a ° i bee aa Pr a Me 6ygsaug To ytitetae teete ott Beir eomaee al ,yitosupert |, treet scat (Leo ceditee to .néades 8 BO Peoe saoe Ye a % 4 os uP a hescyaaes ef ae.hiiy (3UTOFMOIOIN soneg aM Cie to wolfom to mobeext eaietos) tObte Bei go bestioss “0 ¥ ‘ad¢ to efaeuaes fabteo Bez dt sie sed Sie Bees Seneeee Bgoxstyohizel wk teal yliacketg sf Goitam 30 aobes1t eid? i ‘ l, Poe a ene ssardacgae tube ad? bat ,@eSravhe adtcea tletoogayt a end dauwodtis ,beagyese ni anaeve L8GUT sai? asdv reqolevebst t patie loud -baaiet yilei eae eqs ous zo egyebaeq ie . D> pecoges ad’ Yo’ gr tnobtel eit Rivets “etebenoo ong at & it bate ,aAbtino lo aokhreoqeb ent Gycqtae elottud ot fl gee ; edit be (ete tSaa/ 2 17549 og ao geeks 77s edz 103 J of Bertekioe Sta ebasbhnagge eft ssilingeal bes tisteueg eit. Bh aat seésl ,9o [Ae LOI9 act gort est tt stene ast to ris 4 : ; ‘ a > pi dt Yo etied yaa tod (egsttve hed oat ot Desoatis: Pee 4 7 +) vy) a? ae ie anigzem fabged xheds ace ved BALSA ETA sattasen to oped b, 1s ni to beblvoTg €t2 ebeodggs seont to e@ate oat et =) a “webloe. bavet eyawlé se Tee ,eehe ieee tedgid OH. me i | basi arihseooxt -eqn ails to xbod edt Bo ecataae oa bes - a0 eeol ons ai segabroage ear to mob gatsshloe oat. bat et atiiart cledzeo goons etagugeg Lan teobie ea e1saT .einomgos taosetha eat20 te Pty &@ successive loss of motion between segments, until the conjunc- tiva between all but two of the segments is inflexible in some forms; and even in some of the Lepidoptera, entire freedom of motion has been lost. Among generalized forms where the appendages are soldered together, the cuticle of the exposed parts of the body contains but very little chitin, and is but slightly differentiated in texture from the cuticle of the hidden surfaces. When the imago emerges, or even before that time, if the body is slightly pressed, the appendages separate very readily from each other and are not torn upon the emergence of the insect, so that the pupal skin often remains complete except for the slit on the dorso-meson through which the imago emerged. A very different condition exists, however, among highly specialized forms. Here the exposed por- tions of cuticle become very hard and firm, while those which are not exposed are very thin and delicate and are almost entirely destroyed at the emergence of the imago. The outer covering, of course, being so firmly soldered together, remains in one piece and is apparently complete except for the slit through, the insect emerged. This has led many to think that this outer chitinized portion was the entire pupal skin and that it was a structure, analogous perhaps to an egg sheil, in which the pupa had been enclosed. Another remarkable difference between the generalized and the highly specialized Lepidoptera lies in the fact that in the latter the appendages are not fully formed when pupation takes place, but consist of the transparent cuticular coverings through } ne) Poms } ; rasigen nsowded a0 tfem Ter eR wee . in . Co a1asqobiged ont To ance ‘mt: wove) vrteot nood § ets siete eazcl boc ttateady, axoas 8 txat beeoqke, off 20 slotico ett witdatle td, af Baas ethics ettyrt 1 “tess cebbid est 26 Sldbeud’ ott sot) dd Th ows Vawe exoned avs mort yi thaeedt VTey stataqse 3 Audio nize [aqua edt tedt os ,tooedl eaF 26 RATES ot a oe efs co tile ear. t07 tqeo%e xs noitibaoc @astettiS (Teva hegtede | em x0% bostiete yitigtd hrsd era¥ smgned éfot bose ait ae oak bila > “8 pnd: wal to eorsatems wdt ‘=a b + boreifos ylartt oe acted ; 50x62 Lonroo yiteoreqae au (seu hel esd ett .5egs sive Laqug) et itcs att pen aoLiiog Liete age ate ot a yadtaq suoyo tana : -beeotons Mt agended acne tettss ‘eldaxaeme tedtonk: a f ton? enter eeas sietdobiqed bealleioeqe te . * ' aoliaquq sede bento yllot toa eae wag alnesee ony, a dgvotdt agatreves walyotigo Jue teqeaess ats to we oe Pare ren i ‘att A A bs) j : . cen " 10. which one or more slender tracheae may be seen. The duration of the pupal stage doubtless influences this, there being a stronger tendency for highly specialized forms to hibernate as pupae. During the life of the pupa the adult parts are develop- ing, and before it is time for the imago to emerge, the cuticular parts of the adult are fully formed. In the generalized families previously mentioned and in some specialized forms where the pupal cuticle remains more or less transparent, one is able to see a part of the development taking place, especially in the case of the appendages. The scales appear on the legs and wings and the color pattern may often be easily traced on the latter several days before the emergence of the insect. This stage of the insect, after the cuticular parts are fully formed, and while it still retains its pupal skin, has been designated as the preimago. Where the pupal skin is not already dark in color, it grows con- siderably darker in the last few days before the insect emerges, and one is thus able to determine when the preimago stage is reached pl be “ . 7 mari) Cs 8 e \ogreme ot oneal odt tot ‘emRes ok tt exoted: Sar t opemtexg edd sodw ontmietegiiom elge eutt ee ity bas bewet. yligt ene setae nae =e Toe. od eka ‘caetenay tobnete 8 : re) -_ eyed? ,eidt esonentiat ceectdat | $3 pee ‘acied£d of stot beg t Cefoege hai n08, tise sd? equg ed? to erit: eas sotto ode ot .hemtos vite? exe tiobe ody 4 sot pestfetoege emos at sxe Beno ttass, ie Sf #n0 Jaereqenars, aaee zo ono sats vifatoedes ,eoziq, omit sigh bas epel od? ao Teeqge estkos adT -egisi add? no Seoant x ibeae ad caval écete aicT toes? alg te spaegreas | " Bs betangtans send sen te Siiedic tolor al Agab ybasr he toa st ade |. aves 1 oe eit oxried: eye wot taek eae of ssltek | III. EXTERNAL MORPHOLOGY The most important work on pupal morphology has been done by E. B. Poulton and Dr. T. A. Chapman. Poulton 'S1 in his paper on the "External Morphology of the Lepidopterous Pupa" dis- cusses a few pupal structures but does not attempt to name all of the parts or to locate any of them. So far as known he was the first to point out that the pupal structures were more than cases for the imaginal structures and objected to the terms pterothecae, ophthalmothecae, etc., as applied to pupae. Believing that Poulton's theory is correct, such terms have not been used in this discussion, nor the terms wing cases, antennal cases, leg cases, etc., but these are spoken of simply as wings, antennae, maxillae, etc. Chapman's papers, already referred to, discuss very fully some of the structures and describe their exact location; but included only few figures and one was left very much in doubt as to the identity of many of the structures and their location. W. Hatchett Jackson 'S1 published a very valuable paper on the "Morphology of the Lepidoptera" in which he discussed the external determination of sex in the pupa. A short discussion of the chrysalis was given by Dr. S. H. Scudder '89 and some of the parts were named. In a paper previously cited, Dr. A. S. Packard '95 gives many figures of pupae and names the parts, but his homologies were far from correct. It seems necessary, therefore, before proceeding further,to discuss the principal pupal structures and indicate their location by means of figures. The homologies given in this paper were determined by a series of dissections of pupae in various stages of development, 4 ve a drs, ee ay ay Rete , came toon. oat i a 3: ea motlwoe € rgolLoigqral fantetxt® ode tye evietoutse Tague: a watt Ie alk tagel oe “6 lea soe: badd tue fat ‘tue Rare eae ee Ps ad btiovwe ¢a .,- OF e808 961168 e! il 5° 20 nemode thadala ,ategeq itbesh bis’ peaue es ete owbcas wet - ce! qoedast tqohited: end: to ae eau. eit 1h xee To ie vo R18 20 Wt aevts eames ope ea vievoivaty teqgeq # at eb fod va tha Dts segue to wenag le a . Ly ee 17 ¢Ge1teo_ Bot vot hd Ao + ium wit eenoe te Of pret wy, 3 to shoes ye nGWaenksree wd raqeq ali? a nevis “estgg toned) oats 4 ay onete euvofisy ol eaqueg So, ete 12. the preimago being found most valuable for this purpose. Pupae of nearly every family mentioned in this discussion have been studied in this way, beginning with the Micropterygoidea and extending through the Hepialoidea, Cossoidea and other generalized families and including the Saturnioidea, which are believed to be the most specialized of lepidopterous pupae. The change from larva to pupa has been watched in many species and the subsequent: folding and soldering down of the appendages carefully noted. A large number of species have been bred and a study of the method of dehiscence, as shown by the pupal skin, has thrown considerable light on many instances where there was doubt as to the number of free abdominal segments, or where a suture was obscured by folds or other modifications of the integument. The three regions of the body, head, thorax, and abdomen, are easily recognized and each will be discussed in turn. There occur, on all of the regions of the body, in different families prominent projections and ridges of various types especially in the Papilionoidea. These projections have no morphological significance. The Head The usual sclerites found in the head of generalized insects may be located in lepidopterous pupae. The sutures are distinct in generalized pupac, but are obliterated in the more specialized groups. Vertex.-This is an area found on the dorsum of the head. It reaches its highest development in the Gracilarioidea, but is usually distinct in all generalized pupae. It is bounded cephalad bax tleqeaee aedso fan #€eblowacd ob tols lek. ed9 Ee a |S “ Inecpesdye edt has ae tor Im Yee at bed tee Reet ead aqua § A .Dbetom ylintemso segsbaegge’ ede. te awa. antretitae > ; sddaiehienoo mods send oldie [euig Bhd Ee anode 6s 908 1: I A ay + - me ch ae he Pou ae by. Awe seomzue. etae ‘wor sideutey ‘teom: r iz Me need ovad méleagoutb etdd wt beartsioa 1 ebtaet vaeve be godioartes yorzoIM edt dsttw at iondged wer eitt ed. os “Beveds exs doidw ,sebtoloxwiAe od pathy ¢ “2 . “maxt @eyiado sd? .oegeg evorstqobiqe® Ye bowstetooae § ftta od? to ybute s bax werd co gies ac Henne ‘to z to rodmuan edt of es siuob sew sues etedw eeonadadt s fot yd Detvosdo saw etvius & stéay Ie . Saemges coma » thomugetag ott to éaniteclyit i be tus .xetodt ,beed ,yood sd3 Bo aso tyes soxdt act) smut al beeevos ii od [i tw does ~oac ‘bes Ligeoet . tuersttis ot ,ybod edt to eagiget edt-to ee yiletosqee eeqyt ecvoliayv to eogost bse enoitoetorg ‘eolsolodaxom on evad ado Boer tq escent ‘.eebtonoke bast eT - A hoatlaxreney to Sved ally mi Hawor es?irofos Leaves 4m: eeiutue sat .ecqug exoretqohigel af Seiscol ed exoa sdt at betare¥iito one) He S2qed bes Hatensg aid | QUOTE OS of edt to muezod ed?’ ap tanet wexe qe et aes” st tud ,sebiotiellos1D edt of taemqoLlevesd senigit € tonal fodqec hebauod ef #1 eSquy hes stanecieg oman £3 ab ¥ Peri. ‘ ra ais ee ; ties aay & > S rt j Bea? Bi 13: by the Y-shaped epicranial suture, and may be seen in Figs. 3, 10, 29, 46, 49, 53 and 56; v. This area was referred to by Chapman and Packard as the dorsal head-piece. Front.-The front is the sclerite to which the antennae are attached. It is bounded by the epicranial suture on the dorsal surface and on the ventral surface by the fronto-clyreal suture, which normally extends for a short distance caudad from the base of each antenna and then transversely to the median line. In some pupae where there is a "shoving back" of the head parts as in the Pyraustidae and Sphingidae, the front is located on the dorsum of the head. The fronto-clypeal suture is usually not distinct except in very generalized forms. The superfamily Gelechioidea, however, shows it very distinctly. It is indicated mepaee. 1, 8, 36, 30 and 36; f. The front bears two setae on each side the meson in generalized pupae, which are often very conspicuous. Genae.-These sclerites are distinctly bounded in Erio- craniidae and Hepialidae (Figs. 1 and 8; g.) They are found laterad of the front and clypeus and mesad of the glazed eye. The mandibles are always adjacent to the genae at their lateral margins. Clypeus.-But very few pupae have the clypeus definitely bounded. The suture between the clypeus and labrum is seldom present, although it is often indicated by a furrow. It is then impossible to determine accurately as to its presence, but has been considered as if it were present. The boundaries of the clypeus are shown very distinctly in Fig. 1; cl. In the Hepia- ‘Bae wl a berTeter saw sews BGT” . _soeea teen tase oe aa otfrelog edd ah iaome ‘lent Snort fi oft yO bebmran se. ae A sede Loxdagy ede me bas coe aim ode 2 For Sineree Yibearon doo w oo tevenand weit ios sacetas dode 10 “load yatvede” @ ef etald etedm ot? eeblsatda® bae eahtids ~lo~ovaor?t ed?) obeed ott b cot Sestlarengs Yxevi at tqeox yeu, OL evade Stent webt . oe ya ,aeqed Box Heteney, pe aoess eft h r] \ . fs tonkvelb eit, sot stetoe uedtt. seae0 Mg (.5 78. bee £ ey htp wad tie hgee bas" heeex UMs BNOAVLS DBLé tnozt ott? Yo odd of ages speech sents Ore ontash 3d) oved Ssequqg net UteY eagle: —, fo eft seented. etotze edt 3 ‘perectbat metio ee ot Ayoostia 9 ot e# yYLoterupon pepe s ot i os lidae (Fig. 8) there is no clypeo-labral suture present although all the other head sutures are distinct. The clypeus can usually | be identified by the presence of the invaginations for the anterior arms of the tentorium which are associated with its lateral margins. This sclerite often bears prominent setae, and in the pupae of borers, often a distinct cutting plate or ridge. Anterior arms of the tentorium.-These invaginations are very distinct and are either small pores or slit-like openings. They are associated with the lateral margins of the clypeus and are distinct in most pupae (Figs. 1, 14, 19, 33; at). Labrum.-The labrum is usually distinct along its lateral and distal margins, but seldom separated from the clypeus by a distinct suture. Like the clypeus it usually bears setae which are especially conspicuous in the Eriocraniidae (Figs. 1, 3; lb). A peculiar development occurs in the Heliozelidae and some others where the labrum extends caudad over the appendages (Fig. 50). Pillifers.-This term is applied to the caudo-lateral projections of the labrum, which are so well developed in many Lepidoptera. They are very large in certain superfamilies, notably the Pyralidoidea and the Papilionioidea and their presence is easily detected by the lobes which are adjacent to the caudo-lateral angles of the labrum and often come to meet on the meson caudad of it, or are separated by a narrow piece of the labial palpi. (Figs. 70, 72, 74, 76, 77, 78, 79, pf). The mandibles figured by Scudder '89 (Vol. 3, Pl. 87, fig. 35) are the pillifers. There are often well developed pillifers present, however, when there are no external indications of their presence. a 7 . yi 4 os : | te tatcet-oomgte on batt nei te es ad? Io. seeteans att xe teh na? oy * 2) re 1 mad 26 = a4 ad jantotam Letetel ae? i dtiw begeisoesm Bie co bey “qutvotaet odd) om e “ pagina, 8Gl Ih oh .eeres tacaimoag gtasd qesto ett , i€ an ' wf * aye WV So ef 4 of é Pisa “don tteth & motte Pee eqottantgeval eseiT+.gustogmes gai ae emis sotantak ; al . .eeatnedo sAtfi-sile to eetog iihame. weer re sd DaB° tomnte i > o. : bs eS 2 zl eoiaitezs Leretsl sdv aain betetooe a oF Ts I > mee 22 ,Gf ,8£ (2 .eet?) eeqng fecm at 7Oneem ; xd 3 fo. sa ott betsis jee woblse ted .2orymeeG # t ¥ fw sates eucod ylipuet gt eberyis et) sat | aaa ESF .o0. 36 RET titmaseoitd ety. ee a sous dqamgo vi tee : 5 ; re ioe bac saebliexotieH -edy at exited tremgotovel ie ua i ait) ao yeygs off wev0 bebiag ehaetxe mvkowt 4 et otdT-.axedene i! a ’ ; ; re of ae ~ - % i ca . - : 4 , s al i . , - 4 i an “ ° 6, es meno of f yeh {ft 56 ere ne ids murda t eit 20 as Moe 7 p | ae a oe “ah i Paxvetefi-obuss sdt ot tasostbs ots aotde eedot ons ba) beroee Paces to Bebver aceem ed? ao teem Of: Gmpt ial base aur dek oa . ~*, = \s ns .telegq Laidel e4¢ lo eosty wotten s ed besexaaaa if | , « * ~ a{diiaen eft |. le Vey oo) a oF at . ecT .erettiitg eit ore (88 ight 08 eee eon) ai d ,tassetg exotiiita. beaoteveb, kts by ngeerg zed? to snot teotbat tt 15. Eye-pieces.-These are situated laterad of the genae and mesad of the antennae. There are always two regions to be noted, a smooth mesal portion, sometimes only a narrow band, but often a wider lunate piece, called the glazed eye-piece and the larger The latter is so called lateral portion, the sculptured eye-piece. because it is always sculptured like the adjacent parts of the | thorax. The sculpturing on the head is seldom like that found on the thorax and abdomen and, strange to say, the sculptured eye- Piece is always like the thorax, although it is probably an ex- tension of the vertex. On the dehiscence of most generalized pupae the eye-pieces are separated from the face-parts and remain attached to the conjunctiva which joins the vertex to the pro- thorax (Fig.43). In the specialized forms they remain attached to the face-parts. A peculiar modification is found in the Eucleoidea (Figs. 17, 19, 23; se, ge) in which the eye-pieces form movable flaps seemingly to protect and to cover the prothoracic spiracles which lie underneath. The glazed eye-piece probably represents the pupal eye. Antennae.-These are always attached to the front and extend laterad, curving to the ventral surface of the body mesad of the mesothoracic wings. They may always be identified without Bay trouble (Figs. 1, 8, 11, 15, 28; a). In pupae with broadly pectinate antennae as the Saturniidae the mesal portion is fre- quently elevated and has been referred to as the "stem of the flagellum" of the antennae. Labial Palpi.-These appendages lie adjacent on the meson caudad of the labrum except in the Eriocraniidae (Fig. 1; lp). They are visible in the majority of pupae (Figs. 1, 8, 15, 28, st wakiles aya te ete seed the dud .$fed wottem 8 Yaao somiteuos eepine Sane rpojiat ed? bas eosliq-eys Beeakg ony beliso, rede, or x - pes s6.e2 xettal of .298lg ONS betutohyas itt motte ait to étiag smeosths edt emits beaweafyse: evnnta ans Ngs bdet dedt etl wohies et Beod eae Re an tacrgiues ote exe Lbeacsgivos edt «yee, OF eyasiie prea modigibdie bathe ll iat ge vidsedora et, tt cigodeie nerodd eau ends axawie 64 neg Flom Ie Sones fieh. eh? a0 xot tev oH | _ a exe aco efiso-eoeeld sdf mort Bevesages ote escsig-e | q apoyo wat of wetvev edt entot aotiin evidcaipace oie he 1@ Jiewer ysdt entdt Deatladoesgs, ete mi . say ol cre el noited!Dihom: telavoeg 4A A hies 2 fy ; ‘ : ae om » mect Segelaq-evs edt coldw pas (ee. 66 eo ef a As e313) ae i fe ya: picerdioidg Sh&f LSvag, Ov bie JOLFO CI OF visa imens| bo a Pe « Nidadoig ecatq-ave hesehg ad? | Ak gene bes ait cote Pirie ae, Lagu eft as oe . -. Boe taott odd ot besicatia egepse ese sentl~ -2e3RS e. ee Dpe eft to soniiwe [ettaey, One gs) of Ivatero: bet SQeodiiv beititnebé é¢ eyes yea gemt segmihy oloswod? Pee ree-ed ote exowg ol: te Ge ieee eee at, “a | ‘i “Ie ie sort et neltss0oq febem oe? aebs Larcsoe exit ee pan st ‘ eff lo mete" sit em of Leprtetet need: ead bae bac eecnetas ef? +0," eit co tasoaths sii eegabasgge esedi~. sates Bebeadles : ?*.%.cm Wi -f .gtt) eabdiinersotid omg ak sapege euzdal, eds Be, i a Start i res : - pi | oe Pa 45, 61). They are frequently overlaid and concealed by the maxillae at their proximal end as in Figs. 61 and 67; lp. Often they are entirely concealed by the maxillae with the exception of a small V-shaped piece just caudad of the labrum (Fig. 73). This was thought by Scudder to be a special piece for covering the base of the tongue. Maxillae.- Where labial palpi are visible they occupy a mesal position, caudad of the labrum, with the maxillae laterad of them. When they are invisible and apparently absent, the maxillae lie adjacent on the meson, often overlying and concealing the proximal ends of the labial palpi as mentioned above. The maxillae (Figs. 1, 8, 17, 24, 25; mx) vary greatly in length but are never entirely lacking or concealed in the pupa. They often extend beyond the caudal margin of the wings, sometimes free and sometimes soldered to other appendages. The greatest development is found in certain of the Sphingidae where the maxillae do not extend beyond the caudal magin of the wings but the extra length is taken up in a loop at the proximal end which forms the so-called "jug handle" of Sphinx pupae. The maxillae are always measured on the meson from the caudal margin of the labrum to their distal end and are usually compared in length with the wings which are measured from the caudal margin of the labrum to their caudal margin on the meson. Maxillary Palpi.- Each palpus is represented on each side by a subrectangular or triangular area caudad of the eye- pieces and lying along the cephalic margins of the prothoracic and mesothoracic legs, frequently reaching as far mesad as the proximo-lateral angle of each maxilla. The normal position of edt ud betessnes hae bislaave inane ete: fe aet20 qf {ve bid 2 Jag lt Al ee Bae Semeeoey xeeae: ta : fwolvqeous eft. ct iw se SLL xeom $i ES fefsavace qloriine4 © efit “(89 op tt) ansel ede to Rebeea ‘touts adele heqeds- vi ; sf? gabzevoo to? ensiq Sa toeqera eee aabbiros Yd. Su vont yes oldietv ote fqleqniagges ‘shea oasis beactet ealiixem oit poke ,nyided eg? *. Sebinaa’. oft tac ec? ,tisads) piicetagge base wlitnivad ote qoute nen satieeoncs baw no iyizcevo getto ,csoeemrene a0 Morey: 4 istec: fantel say 4 abate Ltseay yeew (xh 368) Vas VE a ati hefio yedT .#qeq edt mi Defasodor so pabtoas hotels) ® bie eit eemivenoa ,suate-edd to- KESRES catate ene ? tracuc level teetses 3 ont , seyebaeqae code at bs — tox ob ebiilxen en?) exuitw oshtan tae ofd 6 aketede & ttanesl orfxe edt tod eyite eff to migam cebaeet! eed ebellet-oe sid easot dolicw bee femizorqg obs oe qoor ak : fexvesen eyarfe ein esifiizem ent cages zit Eas: ‘te mre f#teib sted? of mrroes edt te aigs se, tenteeo: gad mai S14 Solide aprtw sade ata diqaet a bedégmos eahase fettreco vied? et murdalbied? we pagzoe al sgt a “eve eit to hebgao (aeae Gatérane Pak 19 = aie oiosicidtotq eid to eaigies dttadgeo oil, gacle: ae ait 22 Deuem 142 6S gordeeet yey aompest, eget 9 it pee 5% 1 Oe coldéteoy Lamon: edt ol ftxam i549: 302% 8 oy ee ener - 7 - 7 ~~ - er a ae bts these appendages is discussed under the family Eriocraniidae and shown in part in Fig. 1; mp. They may also be seen in Figs. 28, 30, 32, 36, 38; mp. Structures which may be maxillary palpi are found in the genus Gracilaria (Fig. 47). The peculiar extensions of the maxillae in the Cossoidea and Eucleoidea are not considered as maxillary palpi (Fics. 15, 19, 323). Thorax The three segments of the thorax are always distinct. They are only visible on the dorsum, because the ventral and lateral surfaces are covered by the appendages. Prothorax.- This segment probably varies more in size and shape than any of the others. There are some forms as in the Gracilarioidea, Yponomeu toidea and others where the prothorax is very short on the meson (Figs. 53, 56, 58, 64; p) or even invisible (Fig. 54), but is very wide at each lateral margin. It is longer in the Galleridae and certain families of Noctuoidea than any other pupae examined. Prothoracic Legs.- These lie adjacent to the maxillae at their proximal end. The coxae are frequently exvosed, espe- Cially in generalized pupae where the appendages are free (Fig. 2; 11, 19; cxl) and dissection frequently showed a segment cephalad of the coxa, the trochantin, although there was no distinct suture indicated on the exterior, this being covered by the mouth-parts. The trochanter is a very small segment usually found at the caudal end of the femur when the leg is folded and would therefore be generally concealed by the tibia and tarsus. The femur extends from the trochanter cephalad to the caudal margin of the head. vi aC: ‘eat git al nese od oeils yaa yet. a ‘ee te. tae vad: “ame ce Bete estes ORETS: ytsnetze taliosedg eat oY att) atreftogite erase of be feiienoo $6n STA 4 ebiowlos® Bis nebingeed 4 suit Ae eokitga (88 (OL (OL. coma tated vial | xeog tA ey ey ton tte dt ayerle S18 KSSOL2 : 35 « fe igee eed) “a ies Bae Laserew edd eucanee awetob sit no! ‘okctety x a. sugboeqae eat ¥d pexsves os eon %) tis afte coo eelxav Yidedotq tTremeecje it es: a ri 4 arses? @e66 @2e @2enT G2845 0 oat 1 ve rs itouy es. ‘2 code exaite Kas eepiad 2 seen . neve 10 Cy chee Be OSES bt) TOE Ge. eat ba ? + ie ~ btdutood ‘to eol lime tas seo boa cabpnal cae eit wi } \, Ho SSA eKquee 70 es fitces ef} ot teecat ha ofl eeeet, ~/ened qtomrctits ; pene | i. a * \beeocee YLtiedpeta Sie. Seno eat fae ad ag a Ahi) sat exe aeyebooqge add oveutw esang i |. beletges éeenges & Bevows cisneipes® noktoseeth boe (2 Metatiaq tonttelh on aap at aguioct be wbtngsdoors oa Ph: ont cae-At00t edd eat 34 hed vist .Totvedtxe ead nt. P Sebuzo! wis te bavot ylieder 3 nemyee Lisae Yer Soot % ivi oe ed eroteieds hivow Sag Beptes et get oft nade swine? eli 4 estxe tumet ea? “- -ereter vet | atdks ot: ail bele ' : bast ede To ad fo fatuce ede at be csqee, It is frequently concealed by the tibia and tarsus which are the only portions of the prothoracic leg always visible, but they are often shoved slightly laterad so that a portion of the femur is exposed (Figs. 1, 8, 34, 32, 36; fl). The tibia and tarsus are seldom divided by a suture except in generalized pupae, where all the segments, even of the tarsi are readily distinguished (Figs. 1, worms, 338; 36, 45; 1>). Prothoracic Spiracle.- This is usually located on the dorsum between the prothorax and mesothorax sunk deep in the conjunctiva between the segments with an opening adjacent to the caudo-lateral angles of the prothorax. Its primitive position appears to have been much farther ventrad (Fig. 2 ps) and it is found in this position in the specialized Trichoptera. It re- tains this primitive position in the superfamily Eucleoidea and in the family Nepticulidae. The caudal margin possesses curious modifications in different families in the way of elevated ridges, tubercules, setae, etc., and in some of the Papilionoidea, par- ticularly in the families Hesperiidae and Lycaenidae there seems to be a definite external closing apparatus in many of the genera. Sometimes there is a tuft of setae, in others a plug or plate of somewhat honeycombed appearance. Mesothorax.- The mesothorax is usually considerably longer than the other segments in specialized forms, but in generalized pupae all the segments are more nearly equal. Mesothoracic Legs.- These are folded in exactly the same manner as the prothoracic legs and the femora are very seldom exposed,but may be seen in Fig. 1; f2. The coxae are frequently visible (Figs. 1, 48; cx2). The mesothoracic legs - iS t+ ect .(E% 766. 88 a8 See 4) “9039 a hi oxsitisrensa nl ¢qsone Ss ee a ce bob +e ‘nesovd bes sabbisedeet es ff bast sdi ai seo otom oie etoemges dd Lhe ebgiguee Tome? eft Boe epel otoprotozg ott (as) ‘em oeem scat .(&x6 S| be = Pe >=. i> i Qo G out? pares | ke ... ae 4 J thiegse sivieola [eat etre: ot tofite a ‘esealto adv Veetee Wo. siee co ei ered? 3 20 Lereeqas bedtoo 1es0 svau ef xer.ctt+omea edt =. 2eepdioger’ Tot ee ' ay a peril. toege al psémemyee TeAtc. ede Gees ‘7, * ae. beble? exe seeat, sessed piomnastosah @ s he ‘St :f .@2% of mace sd em, ted besa 4 me are usually longer than those of the prothorax. The tibia and tarsus of each leg are always exposed (Figs. 1, 8, 33, 30; 13). They lie on the venter between the prothoracic legs and the antennae. Mesothoracic Wings.- The wings of the mesothorax almost conceal those of the metathorax, except in the most generalized forms where the appendages are free. In most families they are the only wings visible on the ventral surface (Figs. 1, 8, 16, 17; wl). Tegulae.- The tegulae are the large lobes which cover in the adult, the proximal end of the wing. They do not form separate pupal pieces but are indicated in some pupae (Fig. 2;t). The tegulae are referred to by many authors es the patagia. The patagia are lobes of the pronotum which project over the mesonotum. Alar Furrows.- The furrows along each lateral margin of the mesonotum are designated as the alar furrows. They are best developed in the Aegerioidea (Fig. 37; af) although there are distinct depressions in many families. Axillary Tubercules.- In the genera Tropea and Telea of the Saturniidae, there is found a large tubercule at the base of each wing, with sometimes an additional smaller one. The edges of these tubercules are strongly chitinized and somewhat roughened and serve to cut the cocoon for the emergence of the moth. They are probably assisted in this by the peculiar development of the wing sclerites of the preimago, which protrude into these tuber- cules and are sometimes found to have cut the pupal skin at the apex of the tubercule. ms eit bee enol ciperadéotg add iceowied, rethev eat... siottiosen edt Yo egatw. od? =. meet gipemonsoRs ex tlievenss doom edge ege0o axe xexodfates edt to sec vatt soli lest teed of .O0Tl. Of vegubasgys addi sou ton ob wont ate et? 20 Soe tamizorg, ante ca? | eaeoban nde ae) WOE chad s0ngshp % a (8) Ye baeoqxe eyenis e348 ‘get tone Hl 0 )%) soetute Lexdmev ent oy efdhely is ‘5 ww esdol ental sftoeva BAlvged ea eee tae | mi: aaoa at beotenthal er ded ageei¢g r6teqg edt a6 eaten; os Yigg yd do? Desteten ors GAs og ct isyo.deep ougesio hia) margnete sis to, esdel - [evetsad dese grote enonss > ont < eeort 2 Yor? .ewo rust weld eft de betengiesd eta acti ata eveut?d Susednse (te 730.928) sebio bregek sas a 4 ,eeilins®? yosm ai eno sexetg bfeT be jor? aueatg edt BT ows togsedey oul reed of? 3s sluotadet’ egital © bawot ef Sten? BabhL be exT .eno “tetlomm Senolttbe! ne peadtemoe ad fa ox tad vemos Bae Berkaieigo yiguotte are se locresae fiom eft 1 _piregasme ede tok wooogs eds 710 ot ro ¢rexgofeveh tellwoeg eft yd aint at Betetage, f * i+ suedt otal ebsriddeg Cold lomemteng eae, te, r6 t ta nite lequy ett tue evad ot Smug? wap 20. Metathorax.- This segment is longest in generalized forms where its length is nearly equal to that of the mesothorax. Metathoracic Legs.- The tibiae and tarsi of the meta- thoracic legs are never normally exposed for their entire length but are concealed by the other appendages excepting at their distal end. Only a small portion is visible in specialized pupae, and the appendages are often wholly concealed (Figs. 1, 8, 36, 45; 13). Metathoracic Wings.- These are usually covered by the mesothoracic wings excepting for a narrow strip along their dorsal margin. In a few families a narrow strip of the metathoracic wings is visible on the ventral surface caudad of the mesothoracic wings (Figs. 3, 9, 13; wo). The Abdomen The abdomen consists of ten segments of which three, segments 8-10 are always "fixed", that is, they possess no power of independent motion. In the generalized forms there is motion possible between all of the others. A segment is said to be movable when there is movement between its caudal margin and the segment caudad of it. In many pupae the movable segments are | capable of being "telescoped" so that only their caudal margins are visible. Proleg Scars.- The scars of the larval prolegs are found on the ventral surface near the meson (Fig. 11, pac) and are often conspicuous. Tubercule Scars.- Those families in which the larvae have prominent tubercules show very definite scars in the pupae. These are especially noticeable in the Saturniidae. uc By eaee o an, ae”, if, 4 a ed ot bisa ef tnemgea 4 ,evedto ett to, Lie gentee iF . sre atnsepes aldevos pall cegug yeas) é st ez br Hew isn ene | at tasgant at tasagee ere? Aya gel wv itne pistt 10? Besome: Caceres Teveu baad, wn Be (OAL. welt) belesonoo yefodw gerze eae se mitoeem eit io bebNaD See Tt wWe Lexiceay gilt, ie wid fran Labvieo siedd yleo Seat) oe "heqonaelee® seied 9 ts (osq .If .g2%) power att taun énetave Letigey evial oft dole of eetiies? sgoat ~, 65008 Se ames i TOR Om ey Ke reid. oe fesce yltaem er atesoh ex soit t o ieus? bas seated edt = Bool sibaroisatel oY wit te aaitosoke emgnbeuags tethe See em be Lao saa beg tlizioeqe af eldtehy es no kt nee {iene cas ° vi betroveo vilsues ote stent) > ames pinervor fed? yoola gira wortke ® tOT Bo eegeees ini on fare wotrer, 2 s6ht) wet wen #8 : Caw ae aa ron cu@edog yede (ely tsad Fhe at eyewla al p-* a Pt + } on = +4 ft 3? beridersngn ea7 al .. aide icon fabuso ots weywied Sasaron pe BiTesy oie enolforg Iavtel eof i BiKOE OT +, Sage a cry att at exaoe stintted .viev weit & so ceatedet aie eablinwgtad eat mt oidseotvom na esa | é a 7 - 7 ns af a gg shen treme SR ETM 7 F 49 ‘a, Setae.- There are usually setae present on the abdomen and these are arranged much as those of the larvae. . They are often very inconspicuous, otherwise they might furnish good taxonomic characters. There is often a dense covering of secon- dary setae over the entire surface as in some gelechiids and lasiocampids. The Pterphoridae retain a spiny armature similar to that found in the larvae. Spines.- These are found covering the dorsum of the abdomen in generalized pupae (Fig. 49) and larger ones are also found at the caudal end of the body (Figs. 37, 31). They are arranged in rows on the segments in Tineoidea and Tortricoidea (Figs. 27, 31, 39, 42). | Flanged Plates.- In the pupae of borers the flanged plates are best developed, but are found in other pupae as well. Figure 9 shows them well developed on the dorsum and also shows @ well developed ventral plate on the seventh segment. They are usually developed along the cephalic margin of the segment and prevent the telescoping of the segments. Genital Openings.- In the male the genital opening is situated on the ventro-meson of the ninth abdominal segment. It is usually either a slit-like opening as in Fig. 5 without any adjacent elevations, or with a distinctly elevated tubercule on each side as in Fig. 8go and occasionally is situated in a slight depression. In the females there are two openings which may or may not become confluent. These may be mere rounded pores or | slit-like openings and are associated apparently with the eighth and ninth segments. The boundary lines between segments 8 and 9, a ss - \ ? ale ~ $ r , oe we, uae ts wet oft { Ai se an 3 * ’ ; : iam & hy rs wv ‘Si - hoe iW aw * we 4 res < a 5 - 8 - a Fk de © Me: ; Pr ” 4 M ©. ie saa Lenimohde witape + hn GG .arh al #6 Ba2 }, Betevets yitonttesh «@ “olan, ea yee ent cel t hetendté et vwilenoteac¢go Eas One ite sankneqo wt 61s ene? aelenpt: oat ‘gi od, Cea geet T “ vide ashes betetovess ene ine bee erat jes ted euntl Yxebaved AT and 9 and 10 are rarely distinct on the meson and where they are distinct it seems as if the caudal opening were associated with the tenth segment. In the more specialized pupae the caudal margins of the eighth and ninth segments are more strongly curved cephalad near the meson than in the male (Figs. 34, 44g0) and the segments are dove-tailed together. The presence of the two openings apparently represents the more generalized condition (Figs. 7, 17, 28). They are confluent in Podosesia syringae (Fig. 36) and Archips argyrospila (Fig. 44). Anal Opening.- This is always situated on the meson near the caudal margin of the tenth segment. It sometimes shows as a circular opening (Fig. 7ao) but is usually slit-like (Figs. 8, of, 17). It is usually surrounded on each side with several prominent wrinkles or folds. Anal Rise.- The anal opening is frequently situated on the summit of a mound-like elevation known as the anal rise. The setae on this rise are very conspicuous in certain families of Tortricoidea (Fig. 38, ar). Abdominal Spiracles.- Spiracles are always present on abdominal segments 1-8. The spiracles of the first segment are covered, so far as observed, by the wings, except in the super- family Eucleoidea and the family Nepticulidae. The spiracles of the eighth segment are never functional and show no distinct opening. Spiracular Furrows.- On the cephalic margin of the movable segments cephalad of the spiracles are found furrows which frequently extend almost to the meson on both dorsal and ventral aspects. They occur in several families, as the Liparidae and ge ort ered bat ne Hp itt. no soalte ks: yom ‘aa or a diiw Bete fooses omer yatneqe tetuise edt tt em, amove +2 | fabuso sad saorg howtlstosqa stom eat at. -taongen tl bevive ylgootte estos! ome etiemges eate bas Htdgte edt 9 10: edt bar (ooh ,O¢ - gta) miam Set ae net? node od't * om : gwt eft Lo sooebeng ent tedtegod beLtet-ovob mm rokt thao bes iLeredten etom edt etaseetgert yitas eemninyy slesenhe? ai dmeriingn ozs yout». (88. (bb aft) sligeotygxa egtdonk boat rowem eng no Dedage ‘te geyewle ef efaT -amimeqo Lamhe nwode esaeliemos tI . treme? dtsed om? to nkgteu feby wait), efit-site eUteves ad tye) Goat ight) gutneqo % tevevee Aokw ebte sows Se yiewey at + a bled a0 seliak 5 no betautia yidaeupert ef. aise Lars eT eetA f ait -euit lane od3 ea mota sodtevele elti-fagom £ to 4 to wetiinet afaireo al esque Laeatyo Vaev on8 ents oa ie (88 grt) | ‘ do tnesetq eyswia ete eeloeriqd - -pesoezige J ansooyial ere taemyee Joxrtt edt to eofoatige edT (eat veconadil -teque oft at dqooxe .egatw off YS bevneedo ae tat to ss ele tae alee obbttiootsqel: yiime? eatd: bets seb toe is gonivelh of wots. bits fenotionel even: reo sanagen eft Yo otytam otLedgeo edt a0 - mosey | doltir ewortyt Savot ot aoloetige edt Pied beLadage. eitney Dae Lae toh dtod ato noeom vist ot reomie. ; Py Sta hast ' bop exblanqid edd 66 oott tna {a teves at suoeo ved | Wid ” Tawer } J one yal A 4 ie M ' he Ay bie >! eh ey ke et Fe Geometridae, but are best developed in the Sphingidae where they are lacking in but a few genera. They are usually separated by sharply carinated ridges and are of various types but their func- tion is unknown. Cremaster.- The cremaster is a prolongation of the tenth segment and is not found in the more generalized pupae. It was homologized by C. V. Riley with the suranal plate of the larva. It is of various lengths and shapes and often bears setae at the distal end. Two types of cremaster are shown in Figs. 41 and 44, cr. Its length is measured on the ventral surface from its junction with the curve of the ventral surface of the body as in Fig. 44 where ab represents the cremastral length. IV. CLASSIFICATION As no classification of the Lepidoptera based on pupal characters has been attempted hitherto, consequently nothing had been done to determine what characters were of value in defining superfamilies, families and genera. It has been necessary, therefore, to base specific,generic and other characters on those found in such material as could be secured. The present inves- tigation has been limited by the difficulty in obtaining repre- sentatives of many groups. It is not expected, however, that the tables and descriptions given here will do more than furnish a basis for later work upon the subject. It is hoped that these will call the attention of entomologists to the vast possibilities opened up by the use of the taxonomic characters available in pupae. More than that, further studies on the different groups will make it possible to identify the insect at another stage in ite® edt at peqotevet teed ‘eta. ae it roe vllavet Ore yet tenes wet » tuo" af ud esqyd sooliey to té Bas eoghis bedsn tea wom f a Y ‘ ) : a me A) fda 3 goits,colorg & el Teiegmeze Gt: -. 82 aeee ie +I of ry hesileteney erom emt ni Lavo tom at rel adit to stele Lameswe edt ¢ ot be yoriA He 438 wf jecie brs ssqace Bas. siggas at eno Bt bas f 1qf% ot owes Ome Ter aRgeaE tg eat owt 1 met (6itue Less oe +; go Petieebx el tigas vhbed ett to eostwe [eidaev sit Fo. Svaue Ste same ftonel Laatesmeto, od? BiReueuget UOT TAO OTS ES lipt aie SD 23s leqguq mo. bes retaobiged. edd, to gotdeclTiaeage batt a: iquétie aged call « . ir yi } ~ D ; ri & ' griaizean o4 tuacly on hints eb >t a ees : i t8ee08 eelitmg : nan ae me r — wetitn 'F er i Pe ¢ 1 } Sede ais ‘ 43 oad de mis ad a a St Viegee ad A T2 og Se fy ca: 4) 5 Y ‘ 9 : iy - ¢ Om ee * Tee Be Oy A ae * = LG fe Leb teiean S'S fintetdo mi ytin: setatl og . ny 7 aes oP reas. if. _ he soaqns UOTE: n P ui . ij y . * ‘ Hii dete ++ axon of Li bw- exedgetig y taoddeteones ae , pests ts颒 Beqod ef #1 . toeteee, ob “soek F108 teter* gaititivtseod fakv odd oF ate tno Lomere: oo atodtnetie : » i ‘ * : i 4 ~ ; i Lea ‘ P’ ‘ > 4 v We m ; i i ih ah a4 tee _ - . ‘ we t : = e J ~~ ' d } Lat 1 , = 7 , - « , ~? f 1 - ¥¥ be | i of . * 3 5 - ) a! Z : 4 4 of ff * \ o ii te rE ~ < ~~? tealied a " 4 * 4s “ar ( v i a } od ¢ Vo me ae Cy wu i if ‘es U © yviie We 4 _=* : ne > es es a) Ton Fy hy sp A ; ‘ it . 4 \ es ad gO ist a - nie afta 2 Pe Pon es : ES ‘io ae © ,hoe6em oie ido i : - ‘ 10 A * $ ,5 1% 7 . Dorsum of abdomen with a covering of small spines, usually over the entire length of the segment and not arranged in distinct rows; if spines are arranged in rows then the maxillary palpi are absent; vertex always longer than the prothorax on the meson. GRACILARIOIDEA . Dorsum of the abdomen with a distinct row of spines along the cephalic margin of the segment, with or without a caudal row; spines seldom found elsewhere on the segment but, if present, then the maxillary palpi present and well developed. E. Caudal row of spines never present on the dorsum of the abdominal segments; maxillary palpi always present. TINEOIDEA . Caudal row of spines always present on the dorsum of the abdominal segments; maxillary palpi usually present. F. Distinct cremaster never present; setae never present on the anal rise; wings narrow and pointed; large spines always present on the venter of the tenth abdominal segment. AEGERIOIDEA . Distinct cremaster usually present, if not then setae present on the anal rise; wings broad and never pointed; large spines never present on the venter of the tenth abdominal segment. TORTRICOIDEA BB. Movable abdominal segments never present cephalad of the fourth; appendages never free from each other and usually soldered to the body wall. C. Lobes indicating the presence of pillifers always present, except in Gallerinae (Fig. 69) and Oeneinae (Fig. 80). D. Maxillary palpi usually present, if absent, then abdominal segment seven is movable in the male, the body covered with a spiny armature and both prothoracic and mesothoracic legs extending cephalad between the eyépieces and the antennae, the former reaching nearly to the cephalic margin of the glazed eye, or a deep furrow lined with setae present on the dorsum between the ninth and tenth abdominal segments; antennae never clubbed at the distal end; femora of the prothoracic legs usually visible; labial palpi very seldom visible, except as a small triangular or polygonal area caudad of the labrum and between the pillifers. PYRALIDOIDEA to mutete aa [lawes fi \ P ot em ee pe Wie dy de i> a t TKomMtyee oF ‘ Pyare ‘ so ort ie ~4 : - ee 4 eager Latha hie weisvOS | va 3 one bere Ar ha) . Maxillary palpi never present; antennae always clubbed at the distal end; femora of the prothoracic legs mever visible; a deep furrow lined with setae never present on the dorsum between the ninth and tenth abdominal segments; labial palpi never visible except as small triangular or polygonal areas caudad of the labrum between the pilliters and often entirely concealed. PAPILIONOIDEA CC. Lobes indicating the presence of pillifers never present. D. Mesothoracic wings on the ventral surface at meson usually extending considerably beyond the caudal margin of the fourth abdominal segment, if not, then the body depressed, mostly in the thoracic region, the incisions between the movable segments very deep on the dorsum and venter and less deep at the lateral margins and the caudal part of the antennae always adjacent on the meson for a considerable distance; abdominal segments 1-4 usually longer than the other segments; epicranial suture always present. E. Maxillary and labial palpi present and well developed, and a large portion of the prothoracic femora always exposed; if maxillary palpi are not present then the fronto-clypeal suture never visible; prothorax dis- tinctly shorter on the meson than at each side, so that each half is triangular in outline; appendages soldered to each other but not to the body wall; fronto-clypeal suture never visible; antennae with the caudal portion very rarely touching and not monil- iform in appearance. YPONOMEUTOIDEA . Maxillary palpi usually present, but labial palpi and prothoracic femora seldom visible, if visible, then the fronto-clypeal suture distinct; prothorax usually the same length on the meson and at each side so that each half is subquadrangular in outline; appendages usually soldered firmly to each other and to the body wall; body usually ovate in outline, broadest in the thoracic region and usually strongly depressed; fronto- Clypeal suture usually visible; antennae usually monil- iform in appearance, the caudal portion always adja- cent on the meson, usually for some distance; if only touching, then the fronto-clypeal suture is distinct. GELECHIOIDEA DD. Mesothoracic wings on the ventral surface of the body at meson rarely extending beyond the caudal margin of the fourth abdominal segment; if so then maxillary palpi never present; abdominal segments 1-4 or 1-65 rarely longer than the other segments; epicranial suture seldom visible. . i 4 4 -” 5 n< > ‘ 2 ° - é Pr & a. ¢ q ; ; - i iS veo 20 © . P oe “ute 8 : , . \ sou. @ ve ' ' ; ’ ; hy se eed ;. ne iSO 45 OwISy YOOoC 2 4 « “pl « f oh < earn , . 2 \ \ on J on RR eee So iy | * + . e 7 | j : vy free ; - sham 3 . : ‘ f= > it i : ' ¥ : t i ares Se b 4 i ; ; joe On Ps ) Lite sf ’ q hd ~ « - Oe Med ~~ : 5 Part eft ov a al an ~* . ~~ \ . (o a , iy 5 | i erin) | r= i ; : : : ° ae f‘wteo fl Ame sre gi duet | a 4 ; 4 : » » why BL Fae b C= s wees. | q a rs r, ; rl ‘ : ‘ ? . : » - ia os : AS. ; ‘ ‘ ‘ 4 4 , - - - oo my Ps ; , r a ‘ 7 om - ‘ . ; eyeas ' J & _ : oa ’ j Lid : ne + 5 : ' 2S . aun ee ’ ¥ * ; 4 q Pee f< J a = : we : 0 -_ se oS - és . eal? aan 9? : s cc Oo PASS it ~gqldeuot 7 AT? ye (+ “- > eee SOR 4) wi bMioocs. obes. et } tir Py . Labial palpi usually present and well developed and from one-fourth to one-fifth the length of the wings, if not visible then the body usually shaped as in Fig. 104, the abdomen with setae arranged around the larval verrucae, and usually flanged plates on the abdomen and a cremaster present, or with a more or less dense covering of setae never arranged around larval verrucae, the body never of the shape in Fig.104 and flanged abdomen nor a distinct cremaster. . Labial palpi usually present and well developed and the prothoracic femora usually exposed, or if not then both prothoracic and mesothoracic legs reaching cephalad to the eye-pieces; if both labial palpi and prothoracic femora are wanting then the body of the type in Fig.104, the abdomen with setae arranged around larval verrucae, flanged plates usually pre- sent on the abdominal segments and a distinct cre- master often present; body never with a more or less dense covering of setae, except arranged as mentionel above; maxillary palpi occasionally present. NOCTUOIDEA . Labial palpi sometimes present, body never with a cremaster and always with a more or less dense cover- ing of setae which are never arranged around larval verrucae; prothoracic femora never exposed; maxillary palpi never present. BOMBYCOIDEA . Labial palpi never visible, unless represented by small triangular or polygonal areas caudad of the labrum; body very seldom with visible setae. F. Suture extending between the proximal ends of the antennae and separating the clypeus and front always present and very distinct; antennae never broadly pectinate so that the width is one-fifth of the length; spiracular furrows often present. G. Antennae usually considerably broader near the proximal end, their greatest width greater than that of the prothoracic legs; antennae usually more than three-fourths the length of the wings, if not, then the epicranial suture is present, or the cremaster is wanting, or if present bifurcate at the distal end or bears hooked setae; dorsum of the abdomen usually with a deep furrow between the ninth and tenth segments; scar of a caudal horn never present on the dorsum of the eighth abdominal segment; labial palpi. ‘as small triangular or poly- sonal areas caudad of the labrum. NOTODONTOIDEA Mn See ee VI Sale @. hey iy F ye f ? ne, Tren 1 “ ‘, eae ur Aa 5 re vat ; We ie ‘21 wre dcutaeeed ue yy ed ista 1aah C ee arty iat ot Bi vinT=eno mezs") 7? I ba thOd at? ned aldiety tos ths i = ch iv LeRonds eit; ROL: pit ae Wit yLfepad bas yeeousiev Levies ef coce ey seteBnesg, 2 bis pemohcs 1 vay oetee fo Seksoyeo wunal west Oo. 4. vad. SOG aay (SBOUTTEV favrtal o ') £ som: nomarde’ Begdatlt bre i s.. TOS Re es SeuB telay fertied | ‘vitesse stone? olentanrorgd (2 jousm Fae bdiacr oa itod a nett [ r (tt :ee0@la-s¥e) eam aa Baten ro od sa hives cs atones oheexoszord yoLdée Sid POR eS at egyt a1 bad Ami tov Leviel Bauows . » jaotmatde: edt ie taee 3 pix Pe reat iA | ctl LTE6FGAM . f one ,¢ Qe to. gn iterge eansh ‘oprog lated eabl fine. Perec. don Igiag Late evenle ine, toteemere j >, i tavern =! tide aetee To t DSe.QU S120 SfoBtocsot, _Seorsiey PASI sovon 2¢iaq 218i Tua eidgeasy soven.f Lag tuztals isgocylog to selogaetee Diane : avty toblés cis. yiod Gammdan Eby : it ceewted go isaetxe (Sgr be ‘tS 9 > siriacedes ' Dm Sanat j Bit hz tonitesh wey. aes ee +) - pe ir bet étie deh oo Sten se ao" b 73es ieFid evoungt aivoarine items gi sword yidssebleden el Cespear seamed | id idbiw ‘esteetg sled? bas) tap eeorg Latse ga isnot oteazedtoxy Sift “S Bee) t afsaal egt emp ruot-eatdy | nas @xou- = : etutve La tosto te ode nods hoe V2 td @ sy Uh zo ,erbéiioe oh, cesenmext paid. % Sylood etasd so’ bee Late rd odd, ta i > gseb # téin Yiteban. amo ods. ‘edt Pahveo o tBOe eae. itaet has. ttota,.. a dtdgte adt to Mee i eee faenetg Tevane ; vi Af sity: Sime ge hy tam Leite ‘ dreminse BA ay .aetdal edt te, Behves Beets oe Ti > < wan, : me , Fy) a wai? : Meme alin idkatiaiee, aunties ~ = te GG. Antennae rarely very much broader near the proximal end, usually filiform, their greatest width seldom greater than that of the prothoracic legs, if so, then the cremaster is never wanting, nor bifurcate, nor with hooked setae; antennae never more than three-fourths the length of the wings; epicranial suture never present; dorsum of the abdomen never with a deep furrow between, ninth and tenth segments; scar of a caudal horn usually present on the dorsum of the eighth abdominal seg- ment; labial palpi never visible. SPHINGOIDEA . Suture between the proximal ends of the antennae and separating the front and clypeus obsolete for the greater part of its length; antennae always broadly pectinate and the width at least one-fifth of the length and often wider; spiracular furrows seldom present. SATURNIOIDEA A. Pupae with Functional Mandibdles. Among the Trichoptera, from which the Lepidoptera are Supposed to have descended, and which are known to be very closely related, there are many pupae which have functional mandibles. They function, though, merely to assist the pupa to escape from the cocoon. Among the generalized Lepidoptera the pupae of one . Superfamily, the Micropterygoidea, have large mandibles which serve the same purpose as in the Trichoptera. Superfamily Micropterygoidea The most generalized lepidopterous pupae known belong -to the superfamily Micropterygoidea, which includes two families, the Micropterygidae and the Ericraniidae, characterized by the Presence of functional mandibles. Except for fragments of the head, no pupae of the Micropterygidae have been described, but they are undoubtedly the most generalized, because they possess } i+ ta0n 28haote 200m (rev ELORA * ; | syn ¢fLeng OLE ht wi Dees «oars [om RON rc? whee ees i pale fi] roty S43 hw eet he Gi aohlse ate ‘tr hcte vou ot totemiero ett@edy , 06022. eBee ‘ -gatea betood ole. ied) eeeomys bg ton od: 1 itgael ede edtavelsanith: taneous teres iy %% mworoh (iieeend. TENE Stones Lathes ics pagrity din ewsad wortae?: qeeb # cf te 2evar memohds ott ron Sapieo 82 20 t£08 -atasknse \atier shite iAyke sad to mesos nite eo treeets stole tv. raven fotee Tetdas Sha — =) to - «<& ~~ £% ¢ es oft te és Lemdzory em raawsed etutuc a to? +a foed (sty lo Tete FIOCLe edt th. t AL Sot bas STOLL Te uz@etans Bh ee | th! cape bad Soe TEsEAIS aie GftLT-sio F: ' ta dibiw way eee PAT ERS vehegss . ewout: Telsoas tee Thy > TG nike itsneg egdRo dotde eevaae wode soredsos O67. 88) Bed) Se Saborao; tied saad? srom ot hipgtze, foe ad sats i ut ’ a AM ip ! Ree Y. et. tapth edt Yo, Senume tewh. ai mueadal See egegylo say noawd edie fatae atl oF heseerage efenats: one hafoetts ste seldtines of* Moker oF 4 latoo atl an wecbread) mete ters 5 ort sositte Laine See vn Febeiso afi booyed 4g saps oD : iedtel eat , aent ote J { eat, 50m 1 ae taiveuve Sia Bae sneered oa nites * bsdoie ata seleiiasm: edT, @- 2 + sary kg ect vk ‘Be etuse fatce out. : Civeod tote Gent .vbod eff: eo ie tisitan {Lereteal-oleiges ae teivotio tsdwauee