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' jij ' L ' »,..i Twii; E.^nLtEST WrxuEo Tv.sects of Amu. icai A Re-examixatiox of the Devonian Insects of New Brunswick, in tiik Light of Criticisms and of New Studies of other Paleozoic Types. a IjKrilEVlNG that soionce is littlo iidviuiciM] hy tin; acrimony c'ii<]fendered by controversial oMSiiys. the writer has always avoided replying to any criticisms of his scientific work, however destructive they might appear at first sight, until in the natural course of sub- sequent studies it became necessary to subject tiiem to public examination. Generally \nnr]\ time will then have elapsed, both [nirties may view the matter more dispassionately and, notwithstanding the delay, the truth is likely to be sooner reached. It is on this account that up to the ])resent time 1 have in no way noticed the objections which Dr. Hagen' made lour years since' to my interpretation of the wing-structure of the Devonian insects,'' although his criticisms were not always couched in the most tempcate language. Now. however, thai it has become necessary, for a work in hand, for me to review systematically the entire series of ])iileo/,oic insects, the Devonian wings have been stuilied anew, with the intention oi' profiting by the comments of so thorough and learned a student of Neuroptera. both recent and fossil, as Dr. Ilagen is everywhere well known to be; of profiting also by the greatly expended special knowledge I have myself gained in the last five years through the kindness of many correspondents (and especially of Mi". R. I). Lacoe). who have liber.illy I'urni.-lied me with a very considerable number of new paleozoic insect ty[»es, discovered in this coimtrx . Without wishing to discredit in the least the v^orth of Di'. ITageu's general opinion in <|uestioiis which affect the Neui'optera. 1 tliiid< it is only fair to point out dispassionately to the iiKpiirer of the future four things. 1 '. 'i'liat in the historic development of the broailer groups of insects, so far as we now an; acquainted with them, no important changes base transpired .s///'c paleo/.oic liuies; while oiu' perplexity regarding the proper relation of paleozoic; insects to uioiK'i'ii types is often very great, and our conclusions variabU'. 2'. That Dr. Hagen ha-' pulilisbed nothing upon paleozoic insecits (apart from the paper aliove refiM'red to ii|)on Devonian insect-) excepting scattered notes on a few which he referred to Termitina and wliieli. in all ])robal)ility. are no Termitina at all.^ 3". That lliei-c Is no evideiiee. hiii the contrary, thai [)r. Hagen in his investigations, uses the '-tlieory of de-cent" as a working liy|)othesis. without which no one who is stud3^ing any group of animals in the |)eriod ol its rise and most: ra[)id evolution can expect to do other- wise than stumble and wander astrav. To refuse to use it is to merit failure. 4°. That I null. M.H. Cciiii|.. Z.Mil., viir, 111). It. ^ Aiiiiiv. .Mem. Host. Soc. Nat. Hist., !>*«<». ' SiH> Pmc. Amor. Acinl. Arts Si-., xx, 167-173. - ..ifiiriiii mT"Tf°" BplftlMUT of tlu> Povoninii wiiijjjs. t)r. Iti^'(>n lias stti(1ie(t m nnfiirc oiity the fin jiiost vasex poorfr) reverses of the orij^niial spociiiiens. while soveral limes hofore and once siuct; tlic })ubliQti* tion of his criticisms 1 have careful] v studied lioth sets toi^ether* Gerephemera simplex. Tiiis insect was phiced I>y me in a distinct fiuuily ^'•roiip to which I y Brdu^iiiait had not lieen pul)lishcd. and I was not aware of the variety of ner.ratioii found in that ancient type. Now that this is known I am fur cliiii'd. notwith^^tandiiiL;- il< anomalous strneture, to hclieve that Gerepheinera more ni should iall in the same p'lieral u'l'oiip. and tlial ihr name oi Atocina sliould he droppe( <1 artieularh as a new stiiiK' makes me see that the neuiMtion wi ar an interpri'tation which le-:seiis the points nf di^tinetion lietweeii them, and icndei's the strncturt' of (lereph niera less anomalous Dr. IIai:-en. on the other hand, refers it uuhe^ifatinirly to the modern ()(h)nata; hut a certain part of my de>;cri]>tion has heeii jilaiidy misunderstood hy him. The costal margin is represented hotli in the description and in the liLiiM'e as two to three mm. lonj^ ; in the fiifure it is not eij^'ht nun. innii-. as >tati'(l li\ him. 1 liave not said that "the mediastinal vein is never a depressed one in >uch insects."' Inn -the maiunua! would then he an elevated and the media-tinal a depressed vein, which | comliinat ion | is never tlie case," etc. lie ri\isf lliey wouM lie the only i'eature- incnn^istcnl with an ndmiate hypothesis, overlook injr the liict that the ne'vules they are -ii||iosed tu cnnnect ,ire stron,Ldy olili(pie, as never in t)donata and usually in other Nemopiera. '1 lie -nperinr oi'iLi'iu of the hrauches of the prin- cipal vein preserved, which he calls -proliahh' the -ector melius." is also entirely inconsist- ent with an Odiinale liypnt!ie-is. jml j.; the niD-t -alimt pDint in the win;: lu-xt to the luimerous ])arallel veins aliove it. Imt in this he does not at all refer. The superior origin of the Kranclies nf this vein, howevei'. i- not unknown in paleozoic wings as I hail supposed, havinir its eoimti'i'p irt in s,.veral of the l*roto|>hasmida. a> I'roiigniarl calN them, and I am now inclined to !ielie\c that this wing should Iind a iilac.- hei'e. s,)iii,.\Jirrc in 'he neiglihurliood of llaplophlehium. This would necessitate a dilli-rent :niil 1 thinly a -oniewliat more lalioual view id' the neuration. viz., that wiiat I hail looked upon a- the externomeijiau is th;' ".itei-uomeilian vi'iii. and that the externomediaii is the fnsi Inandied vein in the wing, counting from the costal horder. the '■mlerealary nei'vnie ' heing r-'ally a lii'indi of this, room I'oi' other hi-anclies heing found. it tlie main externomidiaii luaui h I'.lt the ciirvit ore nf the ape\ of the wing as soon as the other veins; this would hring the sH^rhi l..nd in the outline of the outer horder at the extremity ol the oiiteriiio-.t anal vein i -(, finpii'iii in insects), hut would give an ent)rmous expansion to the anal area, a- in the ra-e in -n\\ ther paleo/oic insects and as occurs in several of the l'rolopiia>mida. esp,.eially in the neiglil.<,ihood ui I lai)lo[)hh-hium. Dr. ftnj»en, exiimitted ttie revefse of the prlncipu! specimen, and say>' of It, it "strongly »'.onfirins my determination." ''It belongs," he says, ''to a part of the base of the wing which is not preserved" in the figure published. It shows, according to him, the '-sector trigonuli inferior," which belongs to the base of the wing and "is to be found only in Odonata, never in Ephemeridae." Unfortunately for all this, the reverse, now figured for the first time (in reverse) on fig. 9 of the acoompanying plate, is the exact counterpart of that portion of the better specimen (of whi(;h a figure was published in my former paper), which \\iin/arthe>it from the l)ase of the wing, and a piece of whi mm., as ' y , "' . Tip of will- >if IMiitc'iihomiTii, restored by IS done by the dotted hues ni the accompauynig sketch, n,-. na^nnvdata. woidd, on the most favorable showing, make a wing of ridiculously extravagant appear- ance ; the course of the known portion of the lower nuirgin will not allow us to suppose, at the outside, more than 5 mm., and probably not more than 2 mm. of the tip to be lost. n. Tho iiiirrowinj? of wlint Vh\ Jln^cn calls tlu> ''second ouhital Hpiiee'* is « comu»oj> fcafiiiv in Kplu'imM-itlat' (r.y. species of ("alliarcys. Clioroterpes. lllasturi.s Atalophlebia, Hlioenantlms. Cliloeon. etc.). tliough not often to cpiite sncli an extent, nor perhaps (|uit(« so rapidly, as here; and as this varies in dillerent sjx'cies ol'tiie same genus, it seems to h(j a very nniiuportant matter; tlie approach oftlie two veins, contrary to Dr, Ilagen's state* ment. is mentioned in my paper. 4. What Dr. Ilairen calls the sector snl)nodalis does not rini unliroken to the tip, as \n all dragon ll«'s 1 have exainineil. Iml is lost in the reticulation shortly l)etoro the nnirgin. In looking ovr all tlie ancient types known. 1 lind none to which this insect may l>e at nil closely com])ared excei)ting tiie K|>hcmeridae ; uidess it Ite iJreyeria, ti) which it l)ears .lome distant resemhlanci". ;'.iid t) which it is not impossible that it is allied ; l)Mt it dillers strikingly in every detail I'rom that fornu and so much more clos(>ly reseml)les the F^pluMneridae of the present day th:it it would apptMr to lie soim-wiiat of a st rain to attempt to hring the-e two ;ire lookni upon as distinct externo- meilian and iuternomediau stems, wt- liiive au ;i(lilitiou;d rescudilauce in this insect to some of the Protophasmida. though not to jJreyeria : yet this repetition, '•which a])pears t(» havt' iio counterpart among living Kpheiueiidae." is iii realit\ a Icature constantly seen in paleo'/.oic wing*;, ami is indicative nuTcly ol simplicity and coumion (>rigin such as we should naturally look lor in early injects: and on this ground we may he justified in con- sid"ring this in«^ect as a representative of a distinct early type of KplnMueriileous insects, — which may he called the I'alephcmeriihuv Lithentomum Harttii. This insect I placed in a tlistinct family of Neuroptera proper, which IVom "•having its nearest atliuily to Sialina in juoderii tiuu'«." 1 proposed to cidl ('ronico^iaiinn. Dr. liagen also recogiiizf* its Sialidaii l'eatui'e< and compare-; the wiul:' lo that of ('hauliode<. adding •the paucitv of the otV-lioots of the sc;i|)ulai' lirauch i-^ liy no means exceptional . . . the living ( 'liaulioile< possesst'S oidy one. ' Here again i-^ an evident misa|ipi'eln'!ieem to be the uio-^t rational (Uie. and though Dr. ilageu seems to imply (though he. does not explicit ly state ) that the cliaracler above iueutione(l wa-; the only one laying claim to distinguish the Cronii.'osialina from the modern Sialina. 1 do not discuss this poiiii here, as I .-^hall soon do 'e group. T 8 SocaOtllMiM foSBili*. On nccoiinT, niuinly of a transverse roln near tlie biiso of tlie wing, •which I considered honioloi-oiis wi almost every otiier feature of the wing was (iistinetively n()U-()(h)nate and generally Sialidan, I looked on this as th(^ typo of ii distinct synthetic! group, which F called lloniotiietidae, u (iiniily ■•forming the eonnecting link hetweon the Ncuroptera projx'r and Pseutt'jnouroptera." Dr. llagLMi. wlio has not seen the single original, says, ''It is obvious that the wing helongs to the Sialina," and exphdns the so-called arculus iis the end of a horny basal ])art of the wing, such as is seen in Corydalis. "'rhe fragment." he says, "shows nothing foreign to the (.'orytIi.-tii> f.i-^ili-. Cnirri It'll >krt('h. HSMK wmmi* ( tlK> iicconipiinviiijr sketch covrortinjr tliat /onuorly jrjvon. in wliii-li tlie lines inarUod In loii^rer (lashes jrivt' tlie eoiijectmal eourse of the veins where they are not li(|ue iMiinclies <'onliued to the a|)ical fourth of the wing, as the externomedian vein: the lirauclies lielow this, as I'ar as hut not iniduding what I formerly considered the evternomedian vein, as hi-anches of tlie internonu'dian vein; and the re- mamiler of the nervules im|)inging on the lower margin, anil more closelv connected than the others hy cross-veins. a>i Immches of the anal vein. Thus interpreted, the wing falls into a group of paleo/oic insects which was perhaps the Mio>.t lunuerously repit'>;cnted of all the old neuroplerous types in carl)oniferons times, a group which is sejiarated I'rom :dl others liv the complete independence of the medi- astinal, and the lac k of any inferior hranches of the scapular vein. — a group to whiidi the name of llomotlu'tidae. with this completi' alteration of the featiu'es hy which it was at first char;icteri/.ed. may he appli<'(l. As the lack of inferior luanclies to the seupular vein is an attrilmte at the present day of nearly all l*seudoneuropter<^tauce in this case oi the "arculus" from the hase of tin- wing, hut the facts given almve -iliow that the explanation is uuneces.sary. Zenoneura antiquorum. A re-examination of both fragmc-nts ot' tlii-: wing in the light of Dr. Ilageii's statements shows my ligure to he correct in e\cry point wherein it ha^ heen criticized, except in the wholly insignincant nuUter ol' omitting to give with sutlicieiit distinetiu'ss what Dr. Ilagen has compared to the ••r«'curreut vein" of lleineri'hidae. ( hi the other hand. Dr. Ilageii !■< incorrect in every statement of fact where his statement controverts mine. Jlehashad the mi-fortune to putili>h his sfatenieiils after an examination of cmly the more incomplete anil Ics- distinct of the two halves of the fossil, lie complains that ••the whole wing is shown iiy numerous parallel and very close longitudinal lines to have heen jjlaced heneath or ahove some jiart of a plant ; on account oi' these lines some parts of the venation are less distinguishahle. " In the hetter St. .h)hn s|)ccimen these lines do not in the slightest degree interfere with the neunition iiinintiiincd with till' St. .lolin spi'i'iuicn Ix'sidu liini. "Its hind nmr;>in," he siiys. '-is ii iittli' htdow the liind margin of thi; main wing." Only a mere Iragnumt of th(,' hind margin exists in the Hoston speeimen, and therefore the marking on the stone wliicli lie interprets as the hind margin of a second wing is reeogni/ahh' with little donht. and a glance at its rtdatioiis to the other liiu'S proves at onee thai it is simply one of the ••numerous parallel and very dose lougitmlinal" lines which he refers to a jilant. An e.\aminalion of the rcveise shows no such mark at all, and the glazed texture of the stone, peculiar to it whiu'e imy part of (he wing is found, does not extend, on either stone, heyond the limits of the wing as I have dcfinetl them. Dr. Ilagen would, however, prohahly hase his doulile wing theory more upon his next |)oinf: that the hranidies of what I have cilled the extcrnoniedian vein (foinidon the outer detached fragment) are elevated, while the '•corrcspouding sectors of the main wing" (hy which I suppose he refers particularly to the scapular and iuternoiuedian veinsof the piirts on the hasal pieci-) •■nvi^ de]»r<'sse(l.'' That is. that the detached fragment represents one wing, the Itasal piece another. lUit in (he St. .luhu s])eciiuen. aud indeed, though less clearly, in the IJoston speeituen. we have ahsolute proof of the inaccuracy of this view, since tlic externonu'dian vein, whether on lia'.s'c, elevated or depressed, according to whether oltvi-rse or reverse is examined; the same is (he casi- in (he reverse sense with llie intiTuomedian vein, which is elevated, on linth pieces, where the externomedian vein is dcpri'ssed. and rice nrsa. A little more familiarity with paleozoic wings would have taught Dr. Ilagen to ex|)ect this very feature. Dr. Ilagen is e(|ually unfortunate with the scratch of a tool on the costal margin »)f the I'oston specimen, which he would look for in vain on the reverse stoue. If it had heen shown on my former plate (lig. o) it woidd have lai i .'I mm. to the right of the outermost cross-vein figured ; his makinif the oflu'r existing cross-veins '•llurc/nrf very douhtful" is accordingly unwarranted. The upper lu'.inc'i of I he mediastinal fork is i-xactly as I have figured it. and not as Dr. Ilagen represents, as the St. .lolm specimen shows more i)lainly than its Hoston reverse. Dr. Ihigen's ••cilia" near this point are again the longitudinal liiu's of his plant. Tlu>re is no sort of re;>; on foi' claiming the •■fork, as it is called by the autlioi'." of (he s<'in)ular vein, as lu-loULrintr to an oveihiim' winir. since it does not exist. Dr. Ilaiicn says he is ••not ahle to classify the iii author." These are l)are assertions, uii.iccompanied l)y any ]U-oof whati-ver. and it would therefoie be waste time to consider them ; it will lie enough to say with regard to the lirst. that, if true, the wing cannot belong to the Nenroptera, as he himself claims it does. My own claim, suppiorted by direct evidence which can be examined into, was that it formed a distinct and extinct family of Nenroptera. My later study of a larger series of paleozoic as well as of existing types con- lirms my first cone liision. (f mt m '^^mmmm /T/-«»rjia*».-/iiW:.*;5v:r:v: Tliis new rt'vii'w of the Dcvouinn iiH(>ots iilt(»r«* Momcwliiit tho ronfluj^lonM wFiicIi Vfc ]^ro* vioii.ily u'liclii'il, (i('ri'|tlu'iiH'iii i^ now coiisidt'i't'd a niciiilx'r of tin' f^roup I'l'otopliasinidii, formcrlv lookcil ii))oii tin its lu'iiri'st ally. Itut from wliiidi it was rcj^ardrd as distinct. Tho Htructurt' of lloniotlictus is shown to Im« did'crt'iit from what was formerly snpposod, in tiikinz iVom it its prcxinned Odonatc allinilit's, Init its position is otherwise retained, and the relation of the major part of the Devonian insects to later, carhonileroiiH types, i^4 shown to lie more inliinati' than was supposed. This latter conclusion has heen reached maiidy l>y a >liidy of forms discovered since the former paper was |)riMted and wlii(di arw yet unpiiltlished ; ami it is tlie oidy point in whitdi the thirteen several general C(»nclusion.s Iniiiiidated in my previoii'^ paper reipiire any es-iential modification. It is even still true that iiotwithstandin;; the discovery of ;rreati'r unity lielween the I)e\onian and carhonif- erons insects, the little liiuna of St. .Inlm has features whicdi instantly stamp it as distinct from thi' ciirlioniferous; since, while most of its uK'iidiers heionj; to restricted groups which occur in carltoniferous deposii-i. they are in most cases very diiVerent from the later meni- l)ers of these ^'roups. • Instead, therelore. of the five sjiecies' liein;,' divided; — none to Kpliemeriilae, two to Odonata. ami three to Neuro|)lera proper i and "proltalily" to Sialina). as claimed by Dr. llap-n. we find none whatever li(don};imr to ( >doii.il,i. Imt of the two so claimed, onu referalile to an ancient type of Kphcnu'ridne. ami one to an ancient ty|)e of I'hasmidu ; while the rem;iiiiinjr three lieioui; to as many distinct families of ancient Neuroptera, doiilitlcss rel.ited to. Iiut still distinct from, .^ialina. two of wlii(di were well represented in carhoniferous times. The third, however, hail, so far as yet discovered, no repres«'ntative even ainonji' paleozoic insects, and has special inteii'si from its distinct resendilance to the carlioniferous I'rotophasmida, — a group aflcrwanls dilll'rentiated as a special type of another lader. FXI'LANAIloN n|' TIIK I'l.ATK. I vcntiiri' tc. .■1.1.1 Id tliis |.:i|.i T :i |i|-.t>' fii^rax cl luiiiiy vi:irs ;i.^t'ili.> crr.it- iiii.i wlii.li my rriiie li:is lifin ilniwn. All tlic liLjiircs cxcept- iiii.' til.'. •"' MIC iif tlip n.ilnral si/c. I'iu. 1. J/'DiK't/ii tii.i j't'isi/is. Vl-^s. '2, ;',. LitliCiit'Uiiiii,, ll'irtlii. 'I'liesi' litfiins ililTvr frmii tliosc ..^ivcii in my (iirmcrly |ml)lislicil plate ton idiisiilcralilf ilc^rci. Fij^. 4. Jh/nrrifil.i r, fiiafiis. Fi^'N. ;'i, <», 7. X'H'iii, iii-'i finfl'/n'inim. V\\x. 'i is maili' ii|i fn'iii si'veral camera skctclics, atul is cnlarijl'il aliuiit "Jll iliamctiTs. Fi'.'s. s. l'.t. '\'\u-<- n|iiisiiii Imlli ol.vifsc ami reverse, as lliey (irii,'innlly iipIX'iircii. Fig. !t is till' . nil' that has never lieen ti«iirc.l lirl'.ir.-. All ur nearly all uf tlicsc willies .ipiiear in rcverscil |Misiti.in on the plate. FIl'. slioiilij have licen tnine.i :i little. Fi._'s. 1(1, II. l'lathf the outir maru'iii not far fr..m tie- tip; this I suspect is simply a scries ofcross veins ami wi>s niuant to rcprcsiiil only that; it is, however. <^ivcii jir.'.'i^cly in this way in my ori^rinal ilr.awiiiLj. Oiiiillin^i DvM'iitus, lull liii[iirlrii)ii. e pro* suiitla, Ht'd, in 'il, aut- i is fiiliiritfil lly„l.i>ciinci. • in rt'Vi'VKfil 11 bit of the lo n-pri'suiit ^ 10 \ \ N^^ M >.uH.l(M -Ir'i . -i •issmann SruJp'