PSYCHIC.

THE WORK OF A DECADE UPON FOSSIL 1NSECTS, 188o-1889.

BY SAMUEL HUBBARD SCUDDER, CAMBRIDGE MASS.

[Annual address of the retiring president ofthe Cambridge Entomological Club, o Jan. 89o.

Some years ago I published an an- whatever cause, and thus endeavor tonotated and tolerably complete list of picture our progress as vividly as possi-papers on fossil insects. It contained ble. Indeed, the mere list of authorsnearly three times as many titles as would be wearisome, for one couldwere referred to by Hagen in his entomo- make a catalogue of the writings of thelogical bibliography nearly twenty years last ten years considerably longer thanpreviously, but, as the multiplication the entire list given by Hagen in 1863of periodical literature had brought in To be precise, I can cite 94 authorsa train of minor papers, largely abstracts and aboLt 225 papers published inand compilations, I remarked that the this decade, against 78 authors and aboutfar greater extent of my list was no 4o papers quoted by Hagen. Or toproof of an increased recent interest in picture it in another way, about onethis field of research, but thought it third of a complete catalogue of papersdoubtful whether in the intervening on fossil insects would belong to theperiod there had been as much activity decade just closed. Nor is the bulk ofas when the works of Heer were open- this literature its only value; it is quiteing the wealth of material at hand. as remarkable for its quality, for by farSo marked a change has now come the most important of the discoveries

about in this respect that I venture this yet made in fossil insects are embodiedevening to invite your attention to a in the researches of the last ten years,review of the advance that has been and there is no reason to suppose thatmade during the past ten years in this we have reached their conclusion.previously neglected field. In doing Note, first, the relatively great num-this [ do not by any means propose to bet of striking discoveries that havecite everypaperthathasbeen published, been made within this period. Thebut only to call your attention to the discovery and careful study of Silurianmore important or interesting, from scorpions in several different parts of

the world,--in Sweden by Thorell and niart inforlns me in a recent letter, a

Lindstr6m, in Scotland by Peach and considerable number of types alreadyHunter, and in New York by Whitfield, signalized in America, which indeedall brought out at very nearly the same we had a right to anticipate by the

time, are unprecedented in the annals comparisons that had been made betweenof this division of science. These were the forms already published fi’om otherfollowed ahnost immediately by Brongni- localities in the two countries, new

art’s surprising discovery of one or" the discoveries on one continent havinghexapods, Pa/aeoalatt,aa," in the Silur- repeatedly been ibllowed sooner or laterian of France, still the only known true by very similar finds on the other. The

insect in this ancient deposit. Coming abundance of cockroaches in both

dovn a stage later we have the remark- countries is fully sustained at Commen-able Devonian insect-fauna of New try, which hasyielded the wst number of

Brunswick, about the nature of which nearly six hundred specimens, or manythere has been so much dispute, first more than are known from all other

announced, it is true, before our period, carboniferous localities in the worldbut only filly published with figures of taken together. Still another strikingthe species in 880; a single addition discovery in the carboniferous rocks is

or two has recently been made to them the recent finding in Silesia ofcoleopteraby Matthew. With them must be the first time that these have been

classed the Devonian myriopods, the signal’ized at this early epoch, but their

earliest known members of that group, description is yet to come.

fully elaborated by Peach. In the car- These are the principal larger discov-

boniferous period we have the striking eries in the paleozoic series, but theywealth of forms from Mazon Creek and have been accompanied by the publica-other deposits in our country which I tion of many striking forms vhich

have described at variot,s times, includ- indicate the ancestral types of livinging so extraordinary a number of blat- insects, or by the better elucidation of

tarians that have ventured to call this types already known but whose signifi-period, so far as its insect-fauna is cance had not been understood. To

concerned, ’the age of cockroaches." specify some of these we may mention

These discoveries, largely due in this Palaeocampa and Aca,[erpeslescountry to the activity and zeal of Mr. among the myriapods the former with

Lacoe, have been even more than paral- the curious and highly developed struc-

leled by the unexampled wealth rightly ture of the spinous hairs, the latter with

claimed for Commentry iu France by its possession of segmental organs or

Brongniart, who as yet Eas published branchial supports as well as stigmata

hardly more than an outline sketch to indicating a probable amphibious habit;

whet the appetite of the zealot. At A.nlracomar[us, Kreisc],eria, and

this place are found, as Mr. Drong- Geralinura the two former examples

January $9,. I’SYCttE. 289

of new extinct family types of arachnids,the last the first instance of the discoveryof the fledibalfli earlier than the terti-aries, and found at brief intervals on twocontinents other than this last of Kuta’sstriking discoveries in the Bohemiancoal field.might well be cited; the gi-gantic ephemerid, Paliug’enia, of Bohe-mia; l)asyleptus, an extraordinaryform of thysanura, a group not previous-ly known earlier than the tertiaries;Corydaloides, like the preceding, oneof Brongniart’s discoveries at Commen-try, remarkable for the extensive displayofbranchiae on the sides ofthe abdomenPetraMattina sublilis of Kliver (Stre-phocladus) with its strange neuration;lZrodia of England with its remarkablecoloration the gigantic Titanoasma,also from Commentry; the nymph ofEtoblatlina Woodward has publishedfrom England, showing the same modeof development among the ancient asthe modern cockroacles; and, finally,Phthanocoris, the only hemipteroidtype yet found in our own paleozoicrocks.

All these memoranda relate to theinsects of the older formations only, butthe statements regarding them in no

proper way indicate the immense strideswe have made in our knowledge of theearlier types. The decade has beenmarked not only by extensive and strik-ing additions to known types, far morethan doubling the number that had beenpreviously published; it has witnessedalso the advent ofmany original workerspreviously wholly unknown in this field,such as Beecher, Deichmfiller, Karsch,

Kliver, Kuta, Matthew, Peach, Sterzel,Thorell, and Whitfield but it has alsoseen the beginning of a new epoch inthe study of the earlier types, in that forthe first time the subjects have beentreated in much more than a scatteredway, by riffler discussions of the syste-matic status of the insects described, byattempts to systematize our knowledge,and by the treatment in single groups ofinsects from various or from all deposits,and not alone in the simple discussionof collections flora a given deposit.Let us hope that the constantly increas-

ing material and our larger knowledgemay permit in a new decade a furthercorrelation, by the comparative study ofinsects of different horizons, especiallyin the carboniferous age.

Previous to the last decade therehad been scarcely a single attempt at thesystematic study of all the older insects,or even of any of the minor groupstbund in the paleozoic rocks. Hagen,indeed, had treated briefly of the fewtermitina known over thirty years agoHeer had attempted a grouping of thecockroaches; and Goldenberg had sum-

marized our knowledge of all by an

attempted classification; but besidesthese I do not recall a single instancewhere any serious attempt had beenmade to collate in a broad way our

knowledge of paleozoic insects as a

whole or in any of the parts. Only be-cause it has so happened that the presentspeaker has been perhaps the most activeworker in this narrow field during thelast decade, is he obliged here to

mention mainly his own work, uince

290 .PSYCHE. January ,Sqo.

it has fallen to his lot, in however im-perfect a way, to attempt a more or lessmonographic treatment of the extincttype of archipolypoda, fbr instance,comprising most of the paleozoic myri-opoda; of the paleozoic arachnida as a

whole, in which he had been precededby this decade by Karsch, working on

much slenderer material and thereforeat much smaller advantage; also on thepaleozoic cockroaches, and on thespecies of Mylacris, a genus of cock-roaches known from several American

deposits; and on the genera allied to

..Dictyoneura, regarded as ancient typesof phasaida. Reference should herealso be made to Peach’s careful workon the carboniferous arachnida ofScotland. In my memoir on the cock-roaches, embracing the discussion of

fifty-eight species referred to elevengenera, it was claimed that their differ-ences from modern types were so fun-damental as to warrant their separationfrom all subsequent and from livingcockroaches as a distinct and equivalentgroup, called palaeoblattariae, andthat they could be further separated intotwo divisions, called respectively my-lacriclae and blattinariae, of wlich theformer was confined to the New World.Brauer has since questioned the valueof the palaeoblattariae as a group, andBrongniart has recently stated that inthe enormous crowd of cockroachesfound at Commentry, the m,ylacridaeare as numerous as the Matlinariae,which probably means that tlae fauna ofCommentry is older than that of theother carboniferous deposits of Europe

and synchronous or nearly so with mostof the cockroach-yielding deposits ofAmerica.Both Brongniart and myself have

also attempted new classifications of thepaleozoic hexapods as a whole, whichdiffer considerably in character, but

which cannot yet fairly be compared;first because mine discusses nearly allthe known types, but includes hardlyany of those found at Commentry, thenalmost wholly unknown, while Brong-niart, writing later, confines himselfalmost entirely to those of Commentrywith only an occasional allusion to pre-viously described types but principallybecause Brongniart’s work is, so fat, themerest sketch with hardly any structuraldetails, a forerunner of what he willsoon publish in extenso concerning thiswonderful fauna, while mine containsfull structural details as a basis for dis-cussion and generalization. In it Ihave endeavored to point out that theexisting orders of insects were not dif-ferentiated in paleozoic times except ina feeble way, prophetic as it were of thefuture, so that the Palaeodictyoptera, as,after Dohrn and Goldenberg. but withan extension of their usage, I had classedfor the first time all known paleozoicinsects, could only be separated into neu-

ropteroid, orthopteroid and hemipteroidgroups. These views, which urgedalso in a special paper showing the de-velopment of the insect-type in time,have been so strenuously opposed byBrauer and others, that their furtherdiscussion can hardly be profitable ex-

cept for those who have an unfortunate

January ,89o.] PS ’CI-E. 291

taste for polemics, at least until thefauna of Commentry, which will cer-

tainly double the field of observation,

gives us a fairer basis for judgment.Meanwhile it may be said that Brongni-art in his sketch hints by many of histerms that he has found the same diffi-culties as those which faced me, and hasbeen forced to admit a synthesis ofstructure in at least some of the oldertypes, which indeed the very laws ofevolution would render probable.At the beginning of this decade our

knowledge of mesozoic insects was verylimited it was almost entirely confinedto the researches of Germar Giebel,Hagen and Weyenbergh on the Jura ofEichstitt and Solenhofen to Heer’saccount of the Liassic insects ofAargauand to Brodie’s and Westwood’s publi-cations on the secondary insects of Eng-land. The horizon has been somewhatextended of late years by the thoroughdiscussion of the Bavarian insects byDeichmfiller and by Oppenheim; bythe careful exploitation of a new localityfor Liassic insects at Dobbertin, Ger-

man3" by F. E. Geinitz; by the con-siderable number of new generic andspecific types of cockroaches from thesecondary rocks of England describedby myself; by the repeated, though not

extensive, discoveries of Fritsch in Bo-hemia, adding interesting material forour very meagre knowledge of creta-ceous insects; and by the discovery at

Fairplay, Col., of a collection of triassiccockroaches of special interest and im-

portance.Among noteworthy contributions to

our knowledge of the insects of thisepoch may be mentioned Oppenheim’sstudy of the group he called ri2bido-rhabd[, which he regarded as a distinctorder and an ancestral type of lepi-doptera. The discussion of the structureof these insects- especially by Oppen-helm and Deichm/iller, has made clearmany points regarding the S,olenhofeninsects which have always been obscure,and brought about the agreement thatthe ripidor,adi must be regarded as

hymenoptera and in no sense prede-cessors of lepidoptera. Geinitz in hisstudy of the Liassic fauna of Dobbertinhas been able to extend considerably ourknowledge of the structure of that pre-vailing mesozoic type, Ortopleaia,known entirely by ":ts wings, and whichhe regards as phryganideous. In our

own country, the triassic cockroach-fauna of Fairplay, just referred to:slows an interesting transition from theolder to the newer forms, which goesfar to substantiate tle differences I havepointed out between paleozoic and latercockroaches; while the study of a largenumber ofspecimens of3/[ormolucoides,long but imperfectly, known from thered sandstone of Connecticut, has en-

abled me to render it in a high degreeprobable that this oldest known insect-larva was a sialid.

In the monographic treatment of me-

sozoic insects we have only to recordthe discussion of the rhipidorhabdi al-ready mentioned, and systematic re-

vision of the mesozoic cockroaches,based on a considerable collection ofEnglish forms new and old, lent me by

292 PS’CHE. January 89o.

that veteran in their study the Rev. P.B. Brodie, a work which included morethan seventy-five species, treated afterthe method employed in the revision ofthe paleozoic forms. The publicationof both these memoirs on the ancient

cockroaches, it may fairly be remarked,has since brought to light many more

new forms, so that during the past de-cade there have actually been added to

the number of pretertiary forms over a

hundred species of cockroaches, aboutequally divided between paleozoic (53)and mesozoic (57) times. A generalaccount of fossil cockroaches based on

these data was given in my "Cockroachof the Past," in Miall and Denny’s"Structure and life history of the cock-roach" (London, i886).

Passing now to tertiary times, we

naturally cannot expect to meet withdiscoveries of equal importance and in-terest to those which throw light uponthe origin of insect-forms, for it is a wellknown fact that the earliest tertiary in-sects are to all general intents and pur-poses identical with those of to-day.They differ no doubt specifically, andeven to a considerable degree generi-cally. Most of those so far recoveredfrom temperate regions indicate a thenwarmer climate, but, taken as a vhole,the grand features of insect-life appearto have been essentially the same sincethe beginning of tertiary times. By our

present researches upon them we no

doubt greatly widen our horizon, andas with modern types there always are

found problems of interest, so will therebe with fossil insects, however recent.

Activity in this field can hardly besaid to be relatively so great as in theothers, nor so great indeed as sometime ago when Heer and Heyden werepublishing extensively, but it neverthe-less has not been insignificant, and it isnoteworthy that more special workwith groups has been undertaken; thusBuckton has summarized our knowledgeof the fossil aphides, Schlechtendahlhas elaborated the psysopoda of Rott,Gourret the arachnida of Aix, Hagenand Kolbe the psoddae of amber, andI the termitlna of Florissant. I mightalso add the butterflies of Florissant, as

my paper, though not yet published,has been months in type, and the gen-eral results were given in a brief paperon "Fossil butterflies" in general, in my"Butterflies of New England." Akinto these can only be mentioned thepaper by Flach on the pleistocene cole-optera of Hfsbach, Schlechtendahl ’s

revision of Germar’s tertiary-fossils,Williston’s notice of tte FlorissantSyrphidae, and mine of the Florissantarachnida, my comparison of the Odo-nata of Florissant and Green River,the detailed study of Planocephalusfrom Florissant, regarded by me as a

new and practically headless type ofthysanura, and, finally, the discussion ofthe structure of this strange type and otthe supposed mite of the Rhenish browncoal, Limnochares, both of which Bert-kau regards as Galgulidae. To thisperiod also belongs my general survey ofthe ptl eontology ofFlorissant.

The additions to our knowledge ofthe amber insects of Prussia during the

anuary 89o.] ,PST’CHE. 293

past decade is surprisingly little. Be-sides the papers of Hagen and Kolbe onthe psoddae, already alluded to, wehave only a description of an Embia byHagen, of three species of 2Vothrus byKarsch, of two tothrideres by Stein,an account of tlephantomyia by OstenSacken, and a generic list of’ hymenop-tera by Brischke. Malfatti also de-scribes two small insects from theSicilian amber. A meagre showingindeed when the collections of un-

worked material are known to be soextensive. I have also noted but a

single paper on the insects found in therecent gum copal, a description of tvoor three species by Q.uedenfeldt.Here may fairly be mentioned a paper

or two on recent insects which throwlight on the structure of extinct types.One of these is the recent notice by deSelys Longchamps of the Japanesedragonfly Palaeophlebia, which hemakes the type of .a new legion, towhich he refers also tteterophlebia andother forms from the secondary rocksof England and Bavaria, and the ter-

tiary deposits of the Rhine. In theother, on the post-embryonic develop-ment of ulus, Heathcote points outthat the relations of the dorsal and ven-

tral regions of the body of the youngu/us correspond exactly with theirpermanent condition in Zuphoberia,a carboniferous myriopod; and he fur-ther holds that the traces of the divisionof the dorsal plates found in the archi-polypoda lend additional strength to thebelief that they are composed in moderndiplopods of two fused segments origi-

nally distinct; which the doubling ofthe internal organs and of the meso-blastic segmentation also indicates.Among the new tertiary fields which

have been opened, and which havegiven rise to some of these researches,and to others upon which I must nottouch, are Felek in Hungary by Stauband others, Kutschlin, Bohemia, byDeichmtlller, and various localities inupper Alsatia by Foerster, in the last ofwhich about a hundred species havealready been found, though none haveyet been worked up. Peat beds havealso begun to be sounded, and notes oftheir contents have been made by Frfih,Geinitz, and Hollingworth, while sinai-larly recent deposits have yielded a littleto Brongniart, Kendall, and Sordelli.Wilkinson and Woodard have alsoshown us that insects may be expectedfrom the tertiaries of Australia. To some

places in our country I will refer later.The more general diffusion of knowl-

edge regarding fossil insects has beenmarked during the past decade. Im-portant new discoveries have found their

way into journals and into papers beforescientific bodies, to such a degree thatit is hard for the bibliographer to keeptrack of them. But besides these wehave had very full analyses of the largerpapers, among which those given by deBorre to the. Belgian entomologicalsociety easily hold the first place. Bib-liographies, like those of Malfatti andmy own, annual reviews of the litera-ture, like those given by Bertkau,Trouessart, Dalton, White, Marcou andothers; general compil,tions of col-

294 PSYCItE. [January 189o.

lated material covering the whole field,like the excellent series published byGoss; others more or less partial or

local, like the lists of Lacoe and Brong-niart, or the dictionaries of Lesley andMiller, or some papers by Brodie aladGoss; still others which pass the wholesubject under one general review, likeone of my own and those of Mauriceand Vidal y Careta,- all these haveserved to advance in one way andanother an interest in this departmentof science and to bring more or lessorder out of previous confusion or mis-

understanding. The most pretentiousof these undertakings is the generalsystematic survey entrusted to me byZittel for his "handbook of paleontol-ogy," in which for the first time sincePictet and Giebel, or for more than

thirty years, a systematic technicaltreatment of the entire series of fossilinsects, myriopods, and arachnids was

attempted, including tolerably full deft-nitions throughout the paleozoic seriesand to some extent in the later, with a

fullness and variety of illustration never

before given. To gather together, as Ibelieve is there doae, even the smallestreferences and weld all into a connectedwhole would have been almost impossi-ble, had not begun at least twentyyears ago a systematic card reference-catalogue in which every such allusion

great or snall is entered and which hasbeen constantly perfecting aad kept upto date. For English readers, the text

of my contribution to Zittel’s Handbuchwas also published ,by our Geologicalsurvey, with a somewhat fuller treat-

ment of the tertiary series, but withoutillustrations.And now, in bringing this too long

address to a close, you may perhaps askwhat the outlook is for the future. Iventure to predict that it will be quiteas brilliant as the past. In the firstplace, publications bringing the wholeknown series of discoveries in systematicorder up to date, like that just pub-lished, alvcays have a tendency to bringout new facts and discoveries. Again,new localities are being found, and infact, the public has as yet only tasted ofthe good things of Commentry andFlorissant, the richest known fields inthe world, respectively, for carbonifer-ous and tertiary insects. When Brong-niart tells us that he has six hundredcockroaches alone at Commentry, wemay well hold our breath, and it is notto be believed that he will delay, longerthan he is compelled by the very richnessof his field, the publication of the resultsof his study on the other insects whoseclassification has already been outlinedby him. As to tlorissant and our othertertiary fields, the work of illustratingthe insects, for which thousands ofdrawings are already made, has, owingto unavoidable engagements, marchedfar ahead of text; but a volurne, withdescriptions of over five l’mndred insects,including mainly the lower orders, andwith over eight hundred figures, isnearly ready for the printer. It willshow that Florissant alone is as pro-ductive as all the tertiary fields of Eu-rope taken together, if we exclude theinsects found in amber. Yet during the

January 89o.] I?’. TC[-IE 295

past summer, in explorations for theGeological survey, I found that thestrata of a considerable tract of country,certainly many, probably hundreds of,

square miles in extent, lying in westernColorado and eastern Utah, xvere packedwith fossil insects as closely as at Flor-issant, where they occupy a lake basinof relatively small proportions: whetherthese new localities will excel or even

equal that place in the variety of theirfossil treasures, is yet to be determined;but there can hardly be any doubt thatwe shall soon be able in our westernterritories to rehabilitate successivefaunas as successfilly as has been done

with many of our vertebrate types, andas has not yet been done for insects in anycountry in the world. Nor are we

confined to our later beds; insect de-posits have now been found in a scoreof places in our extensive carboniferousseries, and it is in no way improbablethat we may find our own Commentryto double the value of the French dis-

covery. What we really need is a score

of trained workers to "go in and pos-sess the land." No one would welcomethem more heartily than one who isalmost a solitary worker in the Ameri-can field.

THE AMERICAN PLUM BORER ’EUZOPHERA SEMI-FUNERALIS’WALK.

BY STEPHEN AI,FRED FORBES CttAMPAIGN IIL.

Although various boring insects have

occasionally attacked the plum, thesehave been species whose principal inju-ries are done to other trees, and no dis-tinctive plum borer has hitherto beenknown in this country. Among these

incidentals enemies are the peach borerSania exltiosa) the flat-headed

apple-tree borer CArysobotrisfemo-rata) the so-called pear-blight beetle

(Xyleborus pyri), and one of the twigborers (ElalShidiou villosum). Sotne-what recently a newly imported Europe-an bark beetle, Scolytus ruulosus, hasattacked a variety of fruit trees, the plumamong them, but by none of these insectshas any constant and serious injury beendone to the latter fruit, so far as I am

now aware. In a species first described

(in this country) in 887, and whoseimmature stages have remained un-

known until the present time, we haveour first example of a borer devoted, so

far as now known, to the plum alone.This species was first reported to me

as injurious z August 887, in a letterfl’om Farmingdale, Sangamon county,illinois, accompanied by a few borersfound in young Chinese plum trees

(Prunus simoni), one of which was

nearly killed by them.The attack was described as most

general near the forks of the trees,especially at the bases of the lower limbs,but the larvae were sometimes found an

inch, or less, within the earth. The

Submit your manuscripts athttp://www.hindawi.com

Hindawi Publishing Corporationhttp://www.hindawi.com Volume 2014

Anatomy Research International

PeptidesInternational Journal of

Hindawi Publishing Corporationhttp://www.hindawi.com Volume 2014

Hindawi Publishing Corporation http://www.hindawi.com

International Journal of

Volume 2014

Zoology

Hindawi Publishing Corporationhttp://www.hindawi.com Volume 2014

Molecular Biology International

GenomicsInternational Journal of

Hindawi Publishing Corporationhttp://www.hindawi.com Volume 2014

The Scientific World JournalHindawi Publishing Corporation http://www.hindawi.com Volume 2014

Hindawi Publishing Corporationhttp://www.hindawi.com Volume 2014

BioinformaticsAdvances in

Marine BiologyJournal of

Hindawi Publishing Corporationhttp://www.hindawi.com Volume 2014

Hindawi Publishing Corporationhttp://www.hindawi.com Volume 2014

Signal TransductionJournal of

Hindawi Publishing Corporationhttp://www.hindawi.com Volume 2014

BioMed Research International

Evolutionary BiologyInternational Journal of

Hindawi Publishing Corporationhttp://www.hindawi.com Volume 2014

Hindawi Publishing Corporationhttp://www.hindawi.com Volume 2014

Biochemistry Research International

ArchaeaHindawi Publishing Corporationhttp://www.hindawi.com Volume 2014

Hindawi Publishing Corporationhttp://www.hindawi.com Volume 2014

Genetics Research International

Hindawi Publishing Corporationhttp://www.hindawi.com Volume 2014

Advances in

Virolog y

Hindawi Publishing Corporationhttp://www.hindawi.com

Nucleic AcidsJournal of

Volume 2014

Stem CellsInternational

Hindawi Publishing Corporationhttp://www.hindawi.com Volume 2014

Hindawi Publishing Corporationhttp://www.hindawi.com Volume 2014

Enzyme Research

Hindawi Publishing Corporationhttp://www.hindawi.com Volume 2014

International Journal of

Microbiology