= — NATIONAL ACADEMY OF SCIENCES. Vie a Varn. FIRST MEMOIR. ON THE BOMBYCINE MOTHS. SS. LIBRARY OF R. D. LACOE. For the Promotion of Research in PALEOBOTANY and PALEOZOOLOGY RETURN TO SMITHSONIAN INSTITUTION WASHINGTON, D. C. NATIONAL ACADEMY OF SCIENCES. Ne = Se ; Ss VW Ore Wa: FIRST MEMOIR. ON THE BOMBYCINE MOTHS. eS Oey. Zo, ANSON Sen Sp SEP 22 1958 | yf SLiBR ARYA Ql Sb! Blo P2X P| C2 oe MON OGTR ATE: ‘ OF THE BOMBYCINE MOTHS OF AMERICA NORTH OF MEXICO, INCLUDING THEIR TRANSFORMATIONS AND ORIGIN OF THE LARVAL MARKINGS AND ARMATURE, Asya Family 1—NOTODONTIDA-. BY ALPHEUS S. PACKARD. . oe - - 6 a a TA ; ‘ ’ aera | hs ee : » 7 inhi} 'r an! é 9 4 i oe »> ant 4 AB THE BOMBYCINE MOTHS OF AMERICA NORTH OF MEXICO. CLONER ENE IS: I. Introduction. | VI. Geographical Distribution of the American Noto- II. Hints on the mode of Evolution of the Bristles, | dontide. Spines, and Tubercles of Notodontian and other | VII. Phylogeny of the Lepidoptera. Caterpillars. VIII. Attempt at anew Classification of the Lepidoptera. TI. On Certain Points in the external Anatomy of Bom- IX. A rational Nomenclature of the Veins of the Wings bycine Larvie. of Insects, especially the Lepidoptera. TV. On the Incongruence between the Larval and Adult X. Systematic Revision of the Notodontid, with spec- Characters of Notodontians. ial Reference to their Transformations. V. Inheritance of Characters acquired during the Life- time of. Lepidopterous Larvie. I.—INTRODUCTION. For some years past the writer has been collec ing materials for a genera: account, systematic and developmental, of our North American Bombycine moths. The leading object or motif of the essay has been to collect materials for working out the origin of the larval forms of the higher Lepidoptera. The attempt has been made, so far as material and opportunity have allowed, to describe in as detailed a way as possible the transformations of our Bombycine moths, in the light of the recent very suggestive and stimulating work of Weismann, entitled Studies in the Theories of Descent (1882). Until within a few years the majority of descriptions of caterpillars have been prepared simply for the purpose of identification, or for taxonomical uses, and without reference to the philosophic or general zoological significance of these changes. The transformations of some of the European Sphingid have been very carefully worked out by Weismann, and also by Poulton, but it is believed that the life histories of the lower, more generalized families usually referred to the Bombyces, especially of the Notodontid, Ceratocampide, Saturniide, Hemileu- cide, Cochliopodide, and Lasiocampide, will bring out still more striking and valuable results, inasmuch as they, or forms near them now extinct, are believed to be closely similar to the stem forms from which many of the higher Lepidoptera have probably been evolved. The aim therefore in such studies should be— 1. To treat the larve as though they were adult, independent animals, and to work out their specific and generic as well as family characters. 2. To trace the origin of mimetic and protective characters, and to ascertain the time of larval life when they are assumed, involving— 3. The history of the development of the more specialized set (hairs), spines, tubercles, lines, spots, and other markings.! ! Besides the work of Weismann, compare also the suggestive papers of E. B. Poulton, in Transactions of the Entomological Society of London, 1884-1888, and my papers: Proceedings of the Boston Society of Natural History, xxiv-v, 1890-91. 7 8 MEMOIRS OF THE NATIONAL ACADEMY OF SCIENCES. 4, To obtain facts regarding the ontogeny of our native species and genera which, when added to what we know of the life histories of European, Asiatic, and South American Bombyces, may lead to at least a partial comprehension of the phylogeny of the higher Lepidoptera, viz, those above the so-called Microlepidoptera. The transformations of the Bombycine moths are especially noteworthy and useful for the purposes we have indicated, since the group is rich in stem forms, because of its probable geological antiquity, and because of the remarkable and significant differences presented by the larvie of many of the groups in the numerous successive stages of their larval life, these stages being characterized by distinctive and highly modified shapes, colors, markings, and armatures. These peculiarities, signalizing nearly each stage, were, we believe, evolved in direct response to the changes in their environment, in their mode of life, or to changes in their food plants, and the necessity of being protected through unconscious mimicry from the assaults of insects and reptilian and avian enemies. The transformations also afford the clearest possible evidence of the action of what Darwin calls ‘inheritance at corresponding periods of life,” and which Hieckel has tersely designated as “ homochronic heredity.” This fact, moreover, of inheritance at corresponding periods of life throws light on the problem so much under discussion at the present day of the transmission of characters acquired at different epochs during the life of the individual. We have devoted a section to a discussion of this question, or rather to a review of some of the facts which strongly suggest the truth of this principle. The characters, so unexpected and striking, as for those worked out in Heterocampa biundata, H, guttivitta, and obliqua, for example, as well as numerous other of the Notodontians and allied families, are plainly enough useless to the insect in the pupa or imago condition, and have evidently been inherited as the result of impressions or stimuli received from without at different periods in the life of the caterpillar alone. Such cases occur in many other Arthropods, especially in the barnacles, and in the Decapoda, as well as in the parasitic worms, but the causes can nearly as well be investigated in these insects, which are so accessible. Another series of problems is opened up by a study of the mouth-parts of the Bombyces and of their venation, which disclose facts intimately bearing on the genealogy of the Lepidoptera. In no other Lepidoptera has the agency of use and disuse, particularly the latter, been more marked. While the mandibles are present in certain of the Tineina and Pyralidina, they have totally disappeared from the so-called Macrolepidoptera, or higher and less generalized and primi- tive groups. In the Bombyces, particularly the Saturnians, the maxill, owing to disuse, have undergone great reduction, with complete loss of their original function. In another direction, i, e., in the veins of the wings, there has been a reduction in their number, and this is correlated with their loss of power of taking food, the great but weak wings of these colossal moths being of no use in seeking for food, which they do not need; as, unlike the swift visitors of flowers, the butterflies, Sphinges, and Noctuids, they are too feeble of flight to sip the nectar of flowers, or too short lived to need any nourishment. The geographical distribution of the Bombyces also tends to confirm the view that they are an ancient and generalized group, and to this subject we have given special attention. In the systematic portion of the work I have endeavored to arrange the families, genera, and even the species, in accordance with the probable phylogeny of the group. I have begun my account of the entire superfamily with what I regard as the most primitive family. The seven subfamilies of Notodontians easily fall into this arrangement; it is not difficult to perceive that the Gluphisiin and Datanin are the most generalized, and that the Cerurin are the most spee- ialized, whether we study the larve or imagines, though much the clearest light of course is thrown upon the subject by thelarve. It is less easy to indicate the true succession of the genera, though the way is made very plain in the subfamily of Heterocampinie. The proper sequence of the species in a large genus is always difficult to make out. It is obvious, however, that the old, unphilosophic method of designating such and such a species as the type of a genus, and then arranging all the others under it, is a thoughtless procedure. Usually MEMOIRS OF THE NATIONAL ACADEMY OF SCIENCES. 9 the type species is the most modified, that most unlike its congeners, unless, as is often the case, it happens to have been the first one of its genus to have been discovered and described. We have thought it better and more philosophical to begin with that species whose larva is the most simple and generalized, and then arrange in their natural order those whose larve are more and more specialized or modified, as regards the number and variety of their markings, or the complexity of their armature. In the genus Ichthyura, for example, the larva of I. apicalis (vau) is the most simple and generalized, not having the high tubercles and bright varied mark- ines of J. inclusa and albosigma. I have therefore supposed this to have been the first species to have evolved, and this decision is supported by the wide distribution of the species and the rather large number of varieties and subvarieties into which the form has been broken up. Tn the ease of the imago, that species which has plain wings without complicated bars and spots is more primitive than those with more complex markings. This course may at times lead to error and uncertainty, and involve more or less hypothesis or guesswork, but the simple attempt will lead to a more careful scrutiny of the larval character- istics, and to a profounder, more thorough, and better knowledge of the biology of the genus, and that of course is the aim in such work. Of course the systematic part of this or any other work of the sort is a necessary preliminary to all other higher endeavors to a complete history of the eroup from a morphological and biological point of view. On this account it is, we think, a great pity that some of the compilers of our check lists of Lepidoptera and other insects, and of our zoological text-books and other works of the sort, still persist to cater to the tastes, rather than true needs, of amateurs and collectors by beginning at the wrong end, i. e., with the “highest” forms rather than with the “lower” or more primitive. Such lists and works would have a far higher educational value and Jead to much better mental training if such compilers could have had some knowledge of the immense impetus given to the science and the new way of dealing with systematic zoology which has resulted from the labors of Darwin, Fritz Miiller, Weismann, and others. In describing caterpillars, particularly those of the Bombyces, I have been particular to dis- tinguish between the three thoracic and the ten abdominal segments, because the former usually differ from the abdominal segments in the number, arrangement, and relative size of the tubercles, warts, and other markings. The warts or tubercles also are grouped into dorsal, subdorsal, and supraspiracular rows (though this latter may in some cases be the subdorsal row), and an infraspiracular row or series. Tn order to obtain further material to finish and to perfect this monograph of the Bombyces, the author would like to obtain from collectors and students in all parts of the country, especially in the Sonthern, Western, and Pacific States, the egg, larve, or moths, in order to fill up gaps, as well as to afford material for illustration. Should anyone rear any of these Bombyces, with a view to publication, I should be greatly obliged for alcoholic specimens of the eggs and different larval stages, which might be sent after such deseriptions were published.' Such specimens would be carefully kept and returned. It will only be by such cooperation that we shall arrive at a fair knowledge of the transformations of this extensive group. This monograph could not have been prepared without generous aid from friends and cor- respondents, as well as from those in charge of the several museums mentioned below, whose hearty cooperation I now acknowledge. Iam specially indebted to Prof. C. V. Riley for the opportunity of freely examining from time to time his extensive collections, so rich in preserved laryze, both blown and alcoholic, the result of yearsof labor while residing in Mlinois, St. Louis, and in Washington, D.C. After presenting them to the United States National Museum, he has continued to allow me to examine the Bom- byces, and loaned me specimens of laryie as well as moths for study and illustration. He has also permitted the use of numerous colored sketches, made by himself or his assistants under his 1 It is earnestly hoped that anyone receiving this memoir will kindly reciprocate by sending the eggs and larve of any Bombycine moths not herein described, packed in tin boxes, to the author, at Providence, R. I., or during July and August, at Brunswick, Me. We still lack the eggs and young larve of Ellida, Lophodonta, Drymonia, and Notodonta. 10 MEMOIRS OF THE NATIONAL ACADEMY OF SCIENCES direction, and has generously turned over to me all his notes on transformations, geographical distribution, etc., his contributions very much enhancing the value of this work. I am also indebted to the authorities of the American Museum of Natural History, New York, for the opportunity of examining the types of the late Mr. Henry Edwards, and a few types of Mr. Grote. Other material and types in the Museum of Comparative Zoology at Cambridge, Mass., of the Boston Society of Natural History, particularly the Harris collection, and the collection of the American Entomological Society Philadelphia, have been examined, and to the authorities in charge I am specially indebted. I should also acknowledge the frequent aid rendered by Mr. Henry Edwards before his death, and the labors of those who have in former years done much pioneer work in collecting and describing the Bombyces, especially of my friend, Mr. Aug. R. Grote, now of Bremen, Germany. Mrs. Annie Trumbull Slosson, of New York, has generously given me valuable material, and given me free access to her collection, and in this and other ways laid me under special obligations. Mr. H. G. Dyar and Mr. B. Neumogen have freely shown me their important collections, and generously loaned specimens for illustration and study. Mr, Dyar has in a number of ways rendered most efficient aid, and has my hearty thanks. We have together made a number of comparisons, and thus arrived at results which otherwise would have been less certain. Dr. J. A. Lintner, State Entomologist of New York, has opened his collection to me, and loaned me several cored drawings of larvee. From Rey. E. D. Hulst, of Brooklyn, I have received by exchange many specimens. Dr. R. Thaxter has permitted me to examine his very valuable collection of larve, now in the Cambridge Museum, and Professor French, of Carbondale, Il, has also kindly helped me. I am much indebted to Miss Emily L. Morton, of Newburg, N. Y., for eggs, Jarve, and the use of several colored drawings of Datana larvie, etc., and for notes on their habits. To Miss Caroline BE. Soule also I am under obligations for a fine colored sketch of Nerice bidentata. I am also indebted to the following entomologists who have aided ine with larvae, eggs, moths, local lists, ete.: Mr. O. S. Westcott, Chicago, Ill.; Mr. Tallant, Columbus, Ohio; Mr. Graef, Brooklyn, N. Y.; Mr. Trevor Kincaid, Olympia, Wash.; Mrs. Fernald, Amherst, Mass.; Mr. Charles Palm, of New York; Mr. William Beutenmueller, in charge of the collection of insects in the American Museum of Natural History, New York, and to others whose aid is acknowledged in the course of the work. To Mr. Joseph Bridgham, who has made the drawings of the larval stages, I am under special obligations. Besides the work of drawing, he has secured many of the larve, and shown the utmost pleasure in aiding me to the extent of his ability. It is to be hoped that the work of the lithographer will bring out the delicacy of color and fidelity in drawing of the artist. I have also had ten drawings of Walker’s types in the British Museum, made by Mr. H. Knight, of London, with the permission of Dr. A.Guenther, superintendent of the zoological department, to whom my hearty thinks are due; also for his courtesy in allowing me, with the kind aid of Mr, A. G. Butler, assistant in entomology, to examine some of Walker’s types. IT have also had copied in the plates a number of excellent colored drawings of caterpillars, made by the late Maj. John Eatton Le Conte, which were loaned me for such a purpose by his son, Dr. John Lawrence Le Conte, a few years before his death. They were made in Georgia, presuma- bly at Sans Souci, on the Ogeechee River, about 16 miles south of Savannah.! BRowN UNIVERSITY, PROVIDENCE, R. I. 1See Seudder's biographical sketch of J. L. Le Conte, Trans. Amer. Ent. Soc., Aug., 1884, p. 9. MEMOIRS OF THE NATIONAL ACADEMY OF SCIENCES. et II.—HINTS ON THE EVOLUTION OF THE BRISTLES, SPINES, AND TUBERCLES OF NOTODONTIAN AND OTHER CATERPILLARS.! It is not improbable that, as a rule, all caterpillars at first lived on grasses, herbaceous and low-growing plants generally, and that gradually they began to climb trees, as the latter became developed, and in time became adapted to an arboreal station. As is well known, no deciduous trees or flowering plants appeared in such numbers as to form genuine forests before the Cretaceous period, and about that time in geological history began to appear the kinds of insects which visit flowers and trees that blossom. The species of the great lepidopterous family Noctuidze, of which we have in the United States alone over a thousand species, are, as a rule, low feeders. Certain species of Mamestra and of Agrotis, ordinarily feeding on grasses and low herbs, will however, especially early in the spring, ascend trees and shrubs of different kinds and temporarily feed upon the buds; and in summer a species of Mamestra will ascend currant bushes in the night and cut off the young, fresh shoots. In the group of forms represented by Catocala, Homoptera, and Pheocyma we have true tree inhabiting caterpillars, and, like the Notodontians and dendricolous Geometrids, their bodies differ remarkably from those of the low feeders, being variously spotted and mottled with shades of brown and ash, to assimilate them to the color of the bark of the tree they rest upon, and are, besides, provided with dorsal and lateral humps and warts, to further assimilate them, in outline as well as in color, to the knots and leaf-scales on the smaller branches and on the twigs among which they feed. And then there is the small group of Noctuo-bombyces, represented by species of Apatela, Platycerura, Raphia, Charadra, and their allies, which closely “mimic” the hairy, penciled, or spiny arboreal Bombyces.’ It should, however, be observed that this is scarcely a case of mimicry, but rather of adaptation; the presence of hairs, pencils, spines, and bristles being apparently due to the caterpillars having changed their environment from herbs to trees, and being subjected to the same conditions as the Bombyces themselves.* In the exclusively low feeding caterpillars of certain groups of butterflies the body is usually smooth and adorned with lines and spots, while the general feeders and many arboreal forms are often variously spined and tuberculated, yet many spined caterpillars of butterflies feed on low herbs. The Sphingidie in part feed on low plants and in part on trees, and they do not, except as regards the caudal horn, exemplify our thesis. ‘This section is reprinted with some alterations from an article in the Proceedings of the Boston Society of Natural History, xxiv, 1890, pp. 482-515, 556-559. “Of 34 species of North America Noctuo-bombyces, whose transformations are known, all except 1 feed upon trees. (See Edwards’s catalogue. ) < 3It is hardly necessary for us to express our entire disagreement with the view of Mr. A. G. Butler, that these Noctuidie are really Notodontians, or in any way allied to them. It seems to us that the characters which he uses.to remove them from the Noctuide are superficial and adaptive. Nearly twenty-five years ago I satisfied myself, after an examination of the denuded head and wings, that the Noctuo-bombyces were true Noctuidw, and did not depart essentially from the typical genera. ‘While many, though not all, butterfly larve, as shown by Scudder and W. H. Edwards, have spine-like eland- ular hairs in the first stage, which may in some cases persist into one or two later stages, the body in many species, especially in those which are not general feeders, but select low-growing, herbaceous plants, becomes smooth and ornamented with stripes or spots. However, as a rule, butterfly larvie can not be divided, as the Bombyees, etc., into high and low feeders; yet from Scudder’s ‘‘ Classified list of food plants of American butterflies” (Psyche, 1889) the following facts and conelusions may be stated: Hesperidv.—Out of 45 species enumerated, all but 6 feed on herbs and especially on grasses, and those which feed on tall shrubs or trees, such as Dpargyreus tityrus and 5 species of Thanaos, stand at the head of the group, which, as everybody knows, is the lowest family of butterflies and nearest related to the moths. Papilionide.—Of the 6 species enumerated, 3 feed on trees as well as shrnbs and herbs; 1 of these, however (P. cresphontes), feeds on trees alone. None of this family are hairy or spined when mature, except P. philenor, with its peculiar flexible, spike-like growths. Pierinw.—Of 10 species, all feed on herbs, rarely on low shrubs, and none are armed with hairs, bristles, or spines. The other two groups (Lycenidw and Nymphalid@) are general feeders, occurring indifferently on herbs, vines, and trees, except the striking case of the 8 Satyrinwe, which feed exclusively on grasses and herbs (Z. portlandia, however, sometimes frequenting the Celtis), The very spiny Argynnis larvie feed on Viola. It should also be noted 1 MEMOIRS OF THE NATIONAL ACADEMY OF SCIENCES. Of the great group of Geometridee many kinds are arboreal (Dendrogeometrids), and in such cases are alihost invariably tuberculated in manifold ways. We know of no hairy or tufted caterpillars of this group or of any family below them, with the exception of the Pterophoride. The arboreal Pyralidie, Tortricide, and Tineidie live in such concealment, between leaves, or in buds, or as miners, that they differ little in their surroundings from the low-feeding forms, and ave thus scarcely ever tubercwlated or spiny; in fact, we can not recall one of these groups which are sv. The Pterophoridie are, to be sure, spiny, but they are low feeders, and their peculiar excretory sete (the Driisenhiirchen or glandular hairs of Zeller') are similar, as Dimmock has. observed, to the glandular or long hairs of plants; Miss Murtfeldt adding that “‘there is a very close imitation in the dermal clothing of the larvie [of Leioptilus sericidactylus| to that of the young leaves of Vernonia, on which the spring and early summer broods feed.” (Psyche iii, 390, 1882.) Returning to the Bombyces, all the Notodontians, without any exception, known to us have trees as their principal, if not exclusive, food plants. Thus, of the 37 species of this group whose larval forms are known, and which are enumerated in Mr. H. Edwards’s “ Bibliographical catalogue of the described transformations of North American Lepidoptera,” together with an additional species (Ichtiyura strigosa) omitted from the catalogue, all are known to feed on trees, unless we except Datana major, which feeds on Andromeda. It is noteworthy that the only species found thus far on a herbaceous plant is the caterpillar of Apatelodes torrefacta, which Harris found on the burdock, though usually it is an arboreal insect. This apparently omnivorous feeder resembles the species of Halesidota, all of which occur more commonly on trees than on herbs, and thus differs markedly from the majority of the Lithosians and Arctians, unless we except the Nolidw. Now the larva of Apatelodes is hairy, the long, white hairs having scattered among them black ones, with more or less black pencils, thus resembling the peculiar yellowish or white caterpillars of Halesidota, with their black tufts and pencils. Similar forms are some of the arboreal, hairy Noctuide, as Charadra deridens. It seems evident that the resemblance to each other in such different groups is the result simply of adaptation, brought about by two factors, the primary one being a change from a low-feeding to an arboreal station, and consequent isolation or segregation, and the secondary one being natural selection, the latter further tending to pre- serve the specific form. It will be seen by the following review that the North American Bombyces in general, with the exception of the Arctians and Lithosians, live on trees, and this will in general apply to the Old World species. In the group of Lasiocampidie, represented by Tolype, Artace, Heterocampa, Gastropacha, and Clisiocampa, the station is an arboreal one, none being known to feed on herbaceous plants. All the Ceratocampidie, all the Hemileuvidie and Attaci, the Platyptericide, all the Cochliopodidie (Limacodes), including both the naked and spiny genera, as well as the Psychid, live exclusively on trees. Of our North American Liparidie, all are arboreal in station, except the Californian Orgyia vetusta, which lives on the lupine. Finally we come to the Arctians. aud Lithosians, whose hairy, or rather setose, larvee in general feed on herbaceous plants and sometimes on trees, being in many cases omnivorous, while those of the Nolidw and Nycteolide whose history is known, are arboreal. Of the Zygienidie, including the Ctenuchidie, the species are low feeders, living on lichens, grasses, and other low plants, or upon vines. The Dioptid genus Phryganidia feeds on the oak. Of the Agaristide, some are low feeders, Euscirrhopterus gloveri feeding on Portulaca, while the majority prefer vines (Vitis, ete.). As to the boring habits.of the Hepialidse and Cossidie, which we now consider as independent groups, related to the Tineina, rather than belonging to the superfamily Bombyces, these seem to be the result of early adaptation. An examination of the food plants of the British species of Bombyces, taken from Stainton’s. Manual of British butterflies and moths (1857), gives the same results for the Old World, as will be seen by the following statements: that many moths, Notodontians among them, which in the Northern States feed on trees alone, in the Gulf States, according to Abbott, feed on shrubs, vines, and low plants, as well as trees. In reply to an inquiry, Mr. W. H. Edwards kindly writes me: ‘I do not think that the butterfly larva which live on trees are under more favorable conditions than low feeders as to healthiness or ease of rearing.” ' Revision der Pterophoriden. Linniea entom., 1852, vi, 356. Mentioned by Dimmock. MEMOIRS OF THE NATIONAL ACADEMY OF SCIENCES. 13 Nolidew.—Of the 3 British species, 2 feed on the oak and 1 on the hawthorn and sloe. Liparide.—Of the 12 species, all feed on trees and shrubs, except Lelia canosa, which lives on reeds and other water plants. It is tufted. Notodontidw.—Of 24 species, 1 ( Diloba ewruleocephala, which is smooth, with no protuberances) feeds on the hawthorn and other plants. Platyptericide.—Of the 6 species, 5 feed on trees and 1 on a shrub. Endromide.—The single species is arboreal. Psychide.—The 2 species, whose larval habits were known, feed on trees and shrubs. Cochliopodide.—tThe 2 species feed on trees. Saturniide.—The single British species feeds on the heather, a shrubby plant. Lasiocampide.—oOf 11 species, 5 feed on trees, the others on shrubs and herbs. Noctuo-bombyces.—All the British species are reported as ‘living on trees and shrubs quite exposed.” Bombycoidew.—All the species of Acronycta live on trees and shrubs. Influence of a change from low to high feeding plants, i.e., from living on an herbaceous to an arboreal station.—It appears, then, that the more typical Bombyces, such as the Ceratocampide, Hemileucide, Attaci, Notodontians, Cochliopodid, and Liparidie, are arboreal in their station, their bodies being variously protected by spines, spinulated tubercles, hairs, or tufts. The group is indeed particularly distinguished for the manifold modifications undergone by what are morpho- logically setze, and it is an interesting inquiry whether the great development of these spines and hairs may not have originally resulted from some change in environment, such as that from low-feeding to high-feeding or arboreal habits. It may be objected that the sete and spines were originally due to the stimulus arising from the attacks of parasitic insects, such as ichneumons and Tachine, or that, as hairy caterpillars are not usually devoured by birds, these hairs and spines have originated through natural selec- tion, and are danger signals, indicating to birds that the wearers of such hirsute and bristling armature are inedible. But while the final purpose or ultimate use of such an armature may serve the useful purpose of protection, and while natural selection may have been the leading secondary factor in the preservation of varietal and specifie forms of hairy and spiny caterpiilars, this does not satisfactorily account for. the initial causes of the growth of tubercles, spines, ete. If spines and hairs form hedge-like guards against the attacks of parasitic insects, why are they not developed as well in the great multitude of low feeders as in the less numerous high feeders? It may be said, however, that Huprepia caja is more subject to the attacks of ichneumons than almost any other larve. (A.G. Butler in Ann. and Mag. Nat. Hist., 1891.) Everyone knows how efficacious any hairs or bristles are in deterring ichneumons and Tachine from ovipositing on caterpillars, and it is well known that naked or slightly piliferous laryie are more subject to their attacks than those which are densely hairy or spinose. The eruciform type of larve.—In endeavoring to account for the origin of the tubereles and spines, as well as the hairs of caterpillars, let us glance at the probable causes of the origin of the caterpillar form, and of the more primary colors and markings of the skin. It was Fritz Miiller who, in his Fiir Darwin (1864), maintained that ‘the so-called complete metamorphosis of insects, in which these animals quit the eggs as grubs or caterpillars, and afterwards become quiescent pupie, incapable of feeding, was not inherited from the primitive ancestor of all insects, but acquired at a later period.” ! In 1869 Dr. F. Brauer? divided the larvie of insects into two groups, the campodea form and raupen form, and in 1871°-1873 we adopted these suggestive views, giving the name of eruciform to the larvee of weevils and other coleopterous larvie of cylindrical form, as well as to the larvie of Diptera, Lepidoptera, and Hymenoptera, all of which are the result of adaptation, being derivatives of the primary campodea type of larva. Brauer’s views on these two types of larvee were also adopted by Sir John Lubbock in his Origin and Metamorphoses of Insects, 1873. ‘Facts and Arguments for Darwin, with additions by the author, Translated from the German by W.S. Dallas, F.L.S., London, 1869. *Betrachtungen iiber die Verwandlung der Insekten im Sinne der Descendenztheorie. Verh. K. K. Zool. bot. Ges. Wien, 1869. ‘Embryology of Chrysopa. American Naturalist, Sept., 1871. 14 MEMOIRS OF THE NATIONAL ACADEMY OF SCIENCES. While the origin of the eruciform larvie of the Cerampycidie, Curculionidie, Scolytide, and other wood-boring and seed-inhabiting and burrowing Coleopterous larvie in general, is plainly attributable to adaptation to changed modes of life, as contrasted with the habits of roving, carnivorous, campodeiform larvie, it is not so easy to account for the origin of the higher metabolous orders of Diptera, Lepidoptera, and Hymenoptera, whose larve are all more or less eruciform. We are forced to adopt the supposition that they have independently originated from groups either belonging to the Neuroptera (in the modern sense) or to some allied but extinct group. Restricting ourselves to the Lepidoptera; as is well known the Lepidoptera are now by some believed to have descended from the Trichoptera or from forms allied to that group. We should, however, prefer the view that the Lepidoptera, Trichoptera, and Mecoptera had a common origin from some earlier, extinct group. The similarity of the imagines of certain of the lower Tineina and certain of the smaller Trichoptera is certainly very marked, the most significant feature being the fact that the mandibles in the two groups are either absent or minute and rudimentary. We have attempted, however,! to show that the larve of the Panorpidee, judging from Brauer’s figures and descriptions, are much nearer in shape and ornamentation to caterpillars than to case worms. Hence, it seems to us probable that the ancestral or stem form of the Lepidoptera. was probably a now extinct group, somewhat intermediate between the Mecoptera (Panorpide) and the Trichoptera. The primitive caterpillar.—We would suggest that the earliest type of Lepidopterous larva was allied to some Tineoid which lived not only on land but on low herbage, not being a miner or sack-bearer, as these are evidently secondary adaptive forms. It is evident, when we take into account the remarkable changes in form of certain mining Tineoid larve described and figured by Chambers? and by Dimmock,' that the flattened, footless, or nearly apodous mining larvie of the earlier stages are the result of adaptation to their burrowing habits. The generalized or primitive form of the first caterpillar was, then, like that of Tineoid larvie in general, and was an external feeder rather than a miner. The body of this forerunner or ancestor of our present caterpillars (which may have lived late in Carboniferous times, just before the appearance of flowering plants and deciduous trees) was most probably cylindrical, long, and slender. Like the Panorpid larvie, the thoracic and abdominal legs had already becom differentiated, and it differed from the larvie of Panorpids in the plant of the abdominal legs being provided with perhaps two pairs of crochets, thus adapting them for creeping with security over the surface of leaves and along twigs and branches. The prothoracic or cervical shield was present, as this is apparently a primitive: feature, often reappearing in the Noctuidie, and sometimes in the Bombycina, and always present in the boring larve of the Hepialidie and the Cossidee. As tactile hairs, defensive or locomotive sete, and spines of manifold shapes occur in worms, often arising from fleshy warts or tubercles, it is reasonable to assume that the piliferous warts of lepidopterous larvee are a direct heirloom of those of the vermian ancestors of the insects. In our primitive caterpillar, then, the piliferous warts were present, eventually becoming arranged as. they now are in ordinary Tineoid, Tortricid, Pyralid, Geometrid, and Noctuid larvee. Origin of the green color of caterpillars.—The cuticle may at first, as in that of caseworms and Panorpid larvie, bave been colorless or horn colored. But soon after habitually feeding in the direct sunlight on green leaves, the chlorophyll‘ thus introduced into the digestive system and into the blood and the hypodermal tissues would cause the cuticle to become green. Afterwards, by farther adaptation and by heredity this color would become the hue in general common to- caterpillars. Moreover, some of the immediate descendants of our primitive caterpillars were probably lighter in hue than others; this was probably due to the fact that the lighter-colored ones: fed on the pale-green underside of the leaves, this difference becoming transmitted by heredity. ''Third Report U.S. Entomological Commission. Genealogy of the Hexapoda, pp. 297-299, 1883, Also American Naturalist, Sept., 1883, 932-945. 2 American Entomologist, iii, 1880, 255-262; Psyche, ii, 81, 137-227; iii, 63, 135, 147; iv, 71. Refers to the larvae of the *‘Gracilaridw ” and ‘“ Lithocolletide” together with Phylocnistis. 8 Psyche, iii, Aug., 1880, 99-103. ‘See the important and quite conclusive footnote by Professor Meldola on p. 310 of Weismann’s Studies in the Theory of Descent, Vol. i (‘I have already given reasons for suspecting that the color of green caterpillars may be: due to the presence of chlorophyll in their tissues, Proc. Zool. Soc., 1878, 159.—R. M.”). MEMOIRS OF THE NATIONAL ACADEMY OF SCIENCES, LD Origin of the lines.—As Weismann has shown, the primitive markings of caterpillars were lines and longitudinal bands, the spots appearing from interruptions or what may be called the serial atrophy of the lines or bands. It is not difficult to account for the origin of the dorsal line, as this would naturally be due to the presence of the heart underneath. This dorsal line is, for example, wanting in the freshly hatched larvie of Spilosoma virginica and Hyphantria textor, but after the first molt of S. virginica there is a slight, diffuse dorsal line of no decided color, though after the second ecdysis it is decidedly whitish, or at least much paler than the surrounding dorsal region. In pale caterpillars the dorsal line may be darker. In the first stages of the two moths in question there are no lines or bands; only the piliferous warts. Whether the subdorsal or the spiracular lines were the first to originate is uncertain, bat probably, from what Weismann has concluded from his-studies of the Sphingidew, the subdorsal arose first. In the second stage of Spilosoma virginica the subdorsal lines are reddish lines extending between the two subdorsal rows of alternating subdorsal piliferous warts, the line becoming more decided, however, in the third stage of this species, there being as yet no signs of a spiracular or of any lateral line. In the freshly hatched larva of H. textor, however, what may be the first beginnings of the subdorsal line are elongated brownish linear spots inclosing the subdorsal row of larger piliferous dots, but not reaching the sutures between the segments. These patches, however, do not in the second stage unite to form continuous lines, but two rows of decided black elongated spots inclosing the black piliferous tubercles. In the freshly hatehed larva of Edema albifrons each of the two subdorsal lines is a row of elongated black spots connected on the three thoracic segments, but separated by the sutures along the abdominal segments. The spiracular line is seen in the same larva of the same stage to be a yellowish band ineclosing the spiracles, and there seems to be a tendency in some, if not many, larvie for the spiracles to be inclosed and connected by a parti-colored or bright line, and for this to have a darker (as in Edema) or lighter edging. Why the spiracles themselves are so apt, as in Bombyces and Sphinges, to be inclosed by a dark or conspicuous line remains to be explained. To return to the subdorsal lines in the pale-reddish larva of Datana, probably D. integerrima, these lines before the first molt are also inclosed by the two rows of subdorsal piliferous spots, and in both the first and second stages there are pale spiracular lines, which appear to be contem- poraneous with the subdorsal line. In the third stage a new dark-red line is interpolated between the subdorsal and spiracular. In the fourth stage the spiracular line has disappeared, and there is a supra and an infra-spiracular pale line on the now brown, dark skin of the caterpillar, Seen from above there are four pale lilac lines, but after molting two of them disappear, and in the last stage there are only two subdorsal lines to be seen, if my colored drawings, very carefully made by Mr. Brigham, are correct. We thus see that after the subdorsal and spiracular lines are formed, others are rapidly introdueed—and some may as rapidly vanish, as necessary features of certain stages—which, when they become useless are discarded. The admirable and most suggestive work of Weismann has placed on a sound basis the theory of the origin of the lines, bands, and spots of the Sphingidwe. The additional notes by Professor Meldola and the beautiful researches of Mr. Poulton have added to the strength of the arguments of Weismann. The lines, bars, stripes, spots, and other colorational markings of caterpillars, by which they mimie the colors and shadows of leaves, stems, etc., have evidently been in the first place induced by the nature of the food (chlorophyll), by the effects produced by light and shade, by adaptation to the form of the edge of the leaf, as in the serrated back of certain Notodontians, by adaptation to the colors of different leaves and to the stems, often reddish, shades of greens, yellows, reds, and browns being as common in the cuticle of caterpillars as on the surface or cuticle of the leaves and their stems or in the bark of the twigs and branches. We (and probably others) have observed that the peculiar brown spots and patches of certain Notodontians do not appear until late in larval life, and also late in the summer or early in the autumn contemporaneous with the appearance of dead and sere blotches in the leaves themselves. Now, to say that thesc wonderful adaptations and marked changes in the markings of cater- pillars are due to “natural selection,” and to let the matter rest there, is quite unsatisfactory. Natural selection may account for the elaboration of these larval forms with their markings after they have once appeared, but we want to discover, if possible, the original causes of such orna- 16 MEMOIRS OF THE NATIONAL ACADEMY OF SCIENCES. mentation, i. e., the primary factors concerned in their evolution. Weismann in his earlier work repeatedly asserts that these changes are due to the direct action of external conditions together with natural selection. Within afew years past many naturalists have returned to a more profeund study of the causes of variation along some of the lines vaguely pointed out by Lamarck.'! It is noteworthy that Darwin changed his views somewhat in his Variation of Animals and Plants under Domestication, and laid more stress on the influence of the surroundings than in his Origin of Species. Neither Weismann nor other authors, however, so far as we know, have formally discussed the probable mode of origin of humps, horns, tubercles, spines, and such outgrowths in larvie. They are so marked and so manifold in their variations in form, and so manifestly related, and in fact have so evidently been directly developed by adaptation to changes in the habits of the Notodontian caterpillars and tree-feeding larvie in general that this group affords favorable material for a study of the general problem. Spines and prickles in animals, like those of plants, serve to protect the organism from externa] attack, and also to strengthen the shell or skin; they are adaptive structures, and have evidently arisen in response to external stimuli, either those of a general or of a cosmical nature, or those resulting from the attacks of animals. It is almost an axiomatic truth that a change of habit in the organism precedes or induces a change of structure. What has caused the enlargement and specialization of certain of the piliferous warts? As remarked by Sir James Paget, “* Constant extrapressure on a part always appears to produce atrophy and absorption; occasional pressure may, and usually does, produce hypertrophy aud thickening. All the thickenings of the cuticle are the consequences of occasional pressure, as the pressure of shoes in occasional walking, of tools oceasionally used with the hand, and the like, for it seems a necessary condition for hypertrophy, in most parts, that they should enjoy intervals in which their nutrition may go on actively.” (See Lectures on Surgical Pathology, I, p. 89, quoted by Henslow, who remarks in his suggestive work, ‘‘The origin of floral structures through insect and other agencies,” that ‘‘the reader will perceive the significance of this passage when recalling the fact that insects’ visits are intermittent.””) It is now assumed by some naturalists that the thorns, spines, and prickles of cacti and other plants growing in desert or dry and sterile places are due either to defective nutrition or to “ebbing vitality” (Geddes), or by others, as Mr. Wallace, to the stimulus resulting from the occasional attacks or visits of animals, especially mammals. It should be berne in mind that the great deserts of the globe are of quite recent formation, being the result of the desiccation of interior areas of the continents, late in the Quaternary epoch, succeeding the time of river terraces. Owing to this ‘Herbert Spencer says: ‘‘The direct action of the medium was the primordial factor of organic evolution” (see The Factor of Organic Evolution, 1886). Claude Bernard wrote: ‘‘ The conditions of life are neither in the organism, nor in its external surroundings, but in both at once” (quoted from J. A. Thompson’s Synthetic Summary of the Influence of the Environment upon the Organism, Proc. Roy. Phys. Soc., ix, 1888). Sachs remarks: ‘‘A far greater portion of the phenomena of life are [is] called forth by external influences than one formerly ventured to assume” (Phys. of Plants, 1887, 191, English translation). Semper claims “‘ that of all the properties of the animal organism, variability is that which may first and most easily be traced by exact investigation to its efficient causes” (Animal Life, ete., preface, vi). ‘‘External conditions can exert not only a very powerful selective influence, but a transform- ing one as well, although it must be the more limited of the two” (Ib., 37). ‘‘No power which is able to act only as a selective, and not as a transforming, influence can ever be exclusively put forward as the proper efficient cause— clusa eficiens—of any phenomenon (Ib., 404). 2Henslow also adds that ‘‘atrophy by pressure and absorption is seen in the growth of embryos, while the constant pressure of a ligature arrests all growth at the constricted place. On the other hand, it would seem to be the persistent contact which causes a climber to thicken.” It mayhere be noted that the results of the hypertrophy and overgrowth of the two consolidated tergites of the second antennal and mandibular segments of the Decapod Crustacea, by which the carapace has been produced, has resulted in a constant pressure on the dorsal arches of the succeeding five cephalic and five thoracic segments, until as a result we have an atrophy of the dorsal arches of as many as ten segments, these being covered by the carapace. Audouin early in this century enunciated the law that in articulated animals one part was built up at the expense of adjoining portions or organs, and this is beautifully exemplified by the changes in the development of the carapace of the embryo and larval Decapod Crustacea, and also in insects. For example, note the change in form and partial atrophy of the two hinder thoracic somites of some beetles, as compared with the large prothorax, due probably to the more or less continual pressure exerted by the folded elytra and wings. MEMOIRS OF THE NATIONAL ACADEMY OF SCIENCES. dd widespread change in the environment, involving a drying up of the soil, much of it alkaline, the direct influence on plant life must have been profound, as regards their protective defenses, and after spines began to develop one can well understand how their shapes should have been regulated for each species and preserved by the set of minor factors which pass current under the term “natural selection.” Animals may also, in some cases, have developed spines in response to a change of environ- ment. If we glance over the epochs of paleontological history we shall see that at certain periods trilobites, brachiopods, ammonites, and perhaps other groups showed a tendency to become tuber- culated, spiny, or otherwise excessively ornamented. These periods must have been characterized by great geological changes, both of the relative distribution of land and water and perhaps of climate and soil. Among the brachiopods, more spiny species occur in the Carboniferous period than in the earlier Paleozoic times.! Among the trilobites, although in Paradoxides and in other genera the gene and sides of the segments are often greatly elongated, we only find forms with long dorsal spines at the close of the Silurian and during the Devonian.’ There are no such spiny forms of ammonites as in the uncoiled Cretaceous Crioceras,* ete. These types, as is well known, had their period of rise, culmination, and decline, or extine- tion, and the more spiny, highly ornamented, abnormal, bizarre forms appeared at or about the time when the vitality of the type was apparently declining. Geddes claims that the spines of plants are a proof of ebbing vitality. Whether or not this was the case with the types of animal life referred to, whether the excess of ornamentation was due to excess or deficiency of food, it is not improbable that the appearance of such highly or grotesquely ornamented forms as ce:tain later brachiopods, trilobites, and ammonites was the result of a change in their environment during a period when.there were more widespread and profound changes in physical geography than had perhaps previously occurred. If the tendency to the production of spines in past geological times was directly or indirectly due to a change in the milieu, and if plants when subjected to new conditions, such as a transfer to deserts, show a tendency to the growth of thorns, or if those which are constantly submerged tend to throw out ascending aerial roots,' or if, like epiphytes, when growing in mid-air, they throw out descending aerial roots, I have thought it not improbable that tubercles, humps, or spines may have in the first place been developed in a few generations, as the result of some change in the environment during the critical time attending or following the close of the Paleozoic or the early part of the Mesozoic age, the time when deciduous trees and flowers probably began to appear. I have always regarded the Bombyces, or the superfamily of silkworm moths, as a very ancient one, which has lost many forms by geological extinction. We thus account for the many gaps between the genera. . Both the larvie and the moths differ structurally far more than the genera of Geometrids and of Noctuidie, and the number of species is less. The two latter families probably arose from the great specialization of type in Tertiary times; while evidently the great ‘Although there are spiny brachiopods in the Silurian, they become more common in the Devonian (e. g., Atrypa hystrix, Chonetes scitula, C. coronata, C. muricata, Productella hirsuta, P. hystricula, P. rarispina, ant Strophacosia productoides), and are apparently more numerous in the Carboniferous formation (e. g., Productus longispinus, P. nebrascensis, Chonetes ornata, C. mesoloba, C. variolata, C. salmaniana, C. setigerus (also Devonian), C, fischeri, ete., Productella newberryi, besides the Permian Productus horrida. 2 Besides Paradoxides, there are such forms as the Cambrian Hydrocephalus carens, the Silurian Dalmania punctata, Cheirurus pleurexanthemus, and Eurycare brevicauda, while the spiny species of Acidaspis seem to be more abundant in the Devonian than in the Silurian strata, but those which bear dorsal spines, such as Deiphon forbesii and drges amatus, are Devonian. ® Quite long spines occur in the Cretaceous species of Crioceras and Aneyloceras matheronianum of Europe, but none, so far as we are aware, in earlier times. 4See N.S. Shaler: Notes on Taxodium distichum, Mem. M. C. Z., xvi, 1, 2, and W. P. Wilson: The production of aerating organs on the roots of swamp and other plants, Proc. Acad. Nat. Sci. Phil., April 2, 1889, quoted in Garden and Forest, Jan. 1, 1890. Shaler conjectures that the function of the ‘‘ knees” is in some way connected with the aeration of the sap. Mr. Wilson shows that “ besides the cypress, other plants which habitually grow with roots covered with water (the water gum, Nyssa silvatica, var. aquatica, Avicennia nitida, and Pinus serotina) develop similar root processes; and what is still more suggestive, Mr. Wilson has induced plants of Indian corn to send roots above the surface of the soil by keeping it continually saturated with water.” It is to be observed that the aerial roots of the latter develop in a single generation. S. Mis. 50 2 18 MEMOIRS OF THE NATIONAL ACADEMY OF SCIENCES. group or superfamily Tineina and allied forms, in some of which the mandibles still persist,! and which in other features (besides haying, as in Nepticula and Phyllocnistis, nine pairs of abdominal legs*) show their-aflinity to the Trichoptera and Mecoptera, originated at an earlier date. As is well known, the Cretaceous land was covered with forests of oaks, liquidambars, maples, willows, sassafras, dogwood, hickory, beech, poplar, walnut sycamore, laurel, myrtle, fig, etc., at or soon after the close of the Laramie epoch, and this may have been the time, if not earlier in the Mesozoic, when in all probability the low feeding caterpillars of that time began, perhaps through overcrowding, to desert their primitive herbaceous food plants and to ascend trees in order to feed on their leaves. Darwin* has made the significant remark “that organic beings, when subjected during several generations to any change whatever in their conditions, tend to vary.” Further on he refers to the general arguments, which appear to him to have great weight, ‘in favor of the view that variations of all kinds and degrees are directly or indirectly caused by the conditions of life to which each being, and more especially its ancestors, have been exposed” (p. 241), and he finally concludes: ‘‘ Changes of any kind in the conditions of life, even extremely slight changes, often suffice to cause variability. Excess of nutriment is perhaps the most efficient single exciting cause” (p. 258). When, in Mesozoic or possibly still earlier times, caterpillars began to migrate from herbaceous plants to trees, they experienced not only some change, however slight, in the nature of their food, but also a slight climatic change, so to speak, involving a change in the temperature. Insects ‘Dr. A. Walter has discovered the presence of minute rudimentary mandibles in the European Micropteryx callella, Tinea pellionella, Tineola biseliella, Argyresthia nitidella, Crambus tristellus, and two genera of Pterophorida (Sitzungsb. Jena, Ges. fiir Med. u. Naturwiss., 1835). I have also detected them in Coleophora coruscipennella and in another Tineid of a genus as yet undetermined. 2The larve of Phyllocnistis have no thoracic legs, but have eight pairs of membranous retractile abdominal legs and an anal pair. (American Entomologist, iii, 256.) Mr. H. T. Stainton kindly informs me that the larvae of Nepticula have no thoracic legs ‘‘ but possess nine pairs of abdominal legs,” which, however, bear no hooks; ‘ they look like so many fleshy prominences.” ’The Variation of Animals and Plants under Domestication, second edition, revised, London, 1888. In the same work Darwin says: ‘‘Nathusius states positively (pp. 99,103), as the result of common experience and of his experiments, that rich and abundant food, given during youth, tends by some direct action to make the head [of the pig] broader and shorter, and that poor food works a contrary result.” Darwin also states that ‘the nature of the food supplied during many generations has apparently affected the Tetigth of the intestines, for, according to Cuvier, their length to that of the body in the wild boar is as 9 to 1, in the common domestic boar as 13.5 to 1, and in the Siam breed as 16 to 1” (Ib.,77). See also the cases mentioned by Semper in his Animal Life, ete., pp. 60-62, and Neumayr’s Stiimme der Thierreichs, 1889,123. Virchow claims that the characters of the skull depend on the shape of the jaw, this being due to differences in food; and here might be quoted the witty remark of Brillat-Savarin, ‘‘ Dis-moi ce que tu manges, je te dirai ce que tu es.” The most remarkable case, and one directly applicable to our subject of the probable cause of the growth of spines, is that cited by Prof. J. A. Ryder: ‘‘Even certain species of fishes, when well fed and kept in confinement, not only spawn several times during a season, instead of only once, as I am informed by Dr. W. H. Wahl, but also when kept from hibernating, as he suggests, tend to vary in the most astounding manner. The wonderful results of Dr. Wahl, attained in the comparatively short period of six years, show what may be done in intensifying the monstrous variations of Japanese goldfishes, through selection, confinement in tanks and aquaria, with comparatively limited room for swimming, plenty of food, etc., all of which conditions tend to favor growth and metabolism, and the expenditure of energy under such wholly new and restricted conditions as to render it almost certain, as he thinks, that these factors have something to do with the development of the enormous and abnormally lengthened pectoral, ventral, dorsal, double anal, and caudal fins of his stock. Some of the races of these fishes have obviously been affected in appearance by abundant feeding, as is attested by their short, almost globular bodies, protuberant abdomens, and greedy habits, as I have observed in watching examples of this short-bodied race living in Dr. Wahl’s aquaria. In these last instances we are brought face to face with modifications occurring in fishes under domestication which are infinitely in excess, morphologically speaking, of anything known among any other domesticated animals. That the abundant feeding and exposure to a uniform temperature during the whole year and confinement in comparatively restricted quarters have had something to do with the genesis of these variations, through an influence thus extended upon the metabolism affecting the growth of certain parts of the body, which have tended to become hereditary, there can searcely be any doubt” (American Naturalist, Jan., 1890). Darwin states that in India several species of fresh-water fishes “are only so far treated artificially that they are reared in great tanks; but this small change is sufficient to induce much variability ” (Variation of Animals and Plants under Domestication, ii, 246). MEMOIRS OF THE NATIONAL ACADEMY OF SCIENCES. 19 living in trees or shrubs several or many feet above the ground are certainly exposed to a more even temperature, as it is colder at night even in midsummer within a few inches of the ground, say about a foot, the usual height to which grasses and herbs grow. The changes, therefore, by day and night are greater at the surface of the ground than among the leaves and branches of a tree. Moreover, forests, not too dense for insect life, with glades and paths to admit the sunlight and heat, must necessarily have a more even temperature and be less exposed to cool winds, and less subject to periods of drought than grassy fields. There is also a less free circulation of air among grasses and herbs, which may be more or less matted and lodged after heavy rains, than among the separate and coarser leaves of trees, such as the different species of oak, which in North America, at least north of Mexico, harbors a far greater number of species of insects (over 500) than any other plant known. On the whole, forest trees support a far larger number of kinds of phytopha- gous insects than grasses or herbs, and may this not be due to better air and a freer circulation, to a more equable temperature, perhaps of a higher average, and thus lead insects to eat more? May not the plump bodies of the larger silkworms, as the larval Attaci, the Ceratocampids, and especially the Cochliopodid (Limacodes), be in some way due to their strictly arboreal environ- ment?! ; When the ancestors of the present groups became fairly established under these changed conditions, becoming high feeders, and rarely wandering to low herbaceous plants, we should have a condition of things akin to geographical isolation.’ The species would gradually tend to become segregated. The females would more and more tend to deposit their eggs on the bark or leaves of trees, gradually deserting annual herbs. For example, the females of the Attaci and their allies, as well as the Cochliopodidie, may have at first had larger wings and smaller bodies, or been more active during flight than their descendants. Their present heavy, thick bodies and sluggish habits are evicently secondary and adaptive, and these features were induced perhaps by the habit of the females ovipositing directly upon leaving their cocoon, and cocoon-spinning moths are perhaps as a rule more sluggish and heavy-bodied than those which enter the earth to transform, as witness the Ceratocampide compared with the cocoon-spinning silkworm (B. mori) and the Attaci. Spinning their cocoons among the leaves at a period in the earth’s history when there was no alternation of winter and summer and probably only times of drought, as in the dry season of the Tropics at the present day, the females may have gradually formed the habit of depositing their eggs immediately after exclusion and on the leaves of the trees forming their larval abode. The females thus scarcely used their wings, while (as in Callosamia promethea) the males, with their larger wings, lighter bodies, broadly pectinated antenne, and consequently far keener sense of smell, could fly to a greater or less distance in search of their mates.2. The principal of segregation* so well worked out by Mr. Gulick, to which Mr. Romanes’ theory of physiological selection is a closely allied factor, if not covering the same ground, would soon be in operation, and the tendency to breed oniy among themselves, rather than with the low feeders, would more and more assert itself, until, as at present, arboreal moths, as a rule almost, if not wholly, oviposit exclusively on the leaves or bark of trees. 1 The fat, overgrown slugworms (Limacodes) may be compared to the overfed, high-bred pig, which eats. voraciously, has little need of rooting, and takes but little exercise. Where, as among cave animals, there is a deficiency of food, we have a constant tendency to slimness, to an attenuation of the body. This is seen in the blind cave arthropods, such as the blind crayfish, blind beetles, blind Cxecidotiea, ete., compared with their allies which live under normal conditions. (See the author’s memoir on the Caye Fauna of North America, etc., Mem. Nat. Acad. Sciences, iv, 24, 1889.) 2The secondary sexual characters so marked in Bombyces are perhaps the result of their peculiar arboreal habits; so also the apterous tendency of Orgyia and a few other forms, especially the arboreal Psyechidie (Cceticus and Thyridopteryx), as well as Anisopteryx and Hibernia. The larvie of the Nyssia feed on trees or low plants. It may be questioned whether any wingless female Lepidoptera live on herbaceous plants. Contrast with them the grass-feeding species of Noctuidie, as those of Agrotis, Leucania, ete. 3In fact nearly the whole group of insects is an example on a yast scale of the principles of segregation, geographical isolation, and physiological selection. As soon as the ancesters of insects acquired wings their milieu was changed, The air rather than the earth became their habitat; the acquisition of wings introduced them to a new world of existence, and free from the attacks of creeping enemies and other adverse conditions to which the terrestrial Myriopods and Arachnids were subjected; the winged insects living a part of their lives, and the most important part, above the surface of the soil, multiplied prodigiously, the number of species being estimated by millions when we take into account the fossil as well as the living forms. 20 MEMOIRS OF THE NATIONAL ACADEMY OF SCIENCES. Coming now to the origin of humps, fixed or movable, and of spines, the change from herbva- ceous to arboreal feeding grounds doubtless affected not only the shape of the body, causing it in many cases to be thiek and fleshy, but also led to a hypertrophy of the piliferous warts common to all lepidopterous larve. The change was probably not necessarily due to the stimulus of the visits and attacks of parasitic insects, because the low feeders are, if anything, at the present day at least, more subject to injury from them than arboreal caterpillars. The cause was probably more pervasive and a result of a change of the environment, such as is seen in the growth of thorns on desert plants, or the knees of the cypress and other water plants, or the aerial roots of ochids and other epiphytes; and that they may have originated with comparative suddenness seems probable when we bear in mind the aerial roots of corn artificially produced in the lifetime of a single individual; though it should be taken into account that plants are far more plastic than animals. If the reader will look at the recapitulations we have given at the end of the detailed life histories of certain Notodontians, it will be seen that not only are there different adaptive charae- ters in the larval, pupal, and imaginal stages, but that the larva itself in its different stages is wonderfully adapted to different surroundings. 1, At first some, indeed most, species live socially on the underside of the leaves near where they were born, and thus concealed from observation. Many have glandular hairs, while the tubercles are more or less uniform. : 2. Toward the end of Stage II and in Stage III they feed in exposed situations on the upper side of the leaves, and at the same time appears the showy style of ornamentation both as regards colors, hairs, and tubercles, approximating to that of the mature caterpillar, whose life apparently is conditioned by its bright colors and bizarre trappings. The smooth-bodied, green larve of Gluphisia, Nadata, Lophodonta, ete., are the primary forms.! Their shape, coloration, and retired habits ally them biologically to the larve of the European Panolis piniperda and other smooth-bodied, green caterpillars with reddish or yellowish stripes, which feed on trees. These smooth larvie are, however, rare and exceptional, especially in North America. But now, owing to a change in the environment, there arose a tendency to the hypertrophy of the normal piliferous warts, and in the actual life history of the caterpillar the tendency manifests itself in the third stage of larval life. We are inclined to believe (1) that the hypertrophy of certain of the tubercles was effected in a comparatively sudden period in consequence of a comparatively sudden change from herbs to trees, and (2) in response to a sudden exigency; (3) that the spines and stiff, dense spinnlated hairs were immediately useful in preventing the attacks of parasitic insects, while (4) the poison glands at the base of the tubercles (in the Attaci, etc.) served to render them distasteful to birds, (5) the bright colors serving as danger signals. The Lamarckian factors (1) of change (both direet and indirect) in the milieu, (2) need, and (3) change of habit, and the now generally adopted principle that a change of function induces change in organs? and in some or many cases actually induces the hypertrophy and specialization of what otherwise would be indifferent parts or organs; these factors are all-important in the evo- lution of the colors, ornaments, and outgrowths from the cuticle of caterpillars.* ‘I am however inclined, since writing the above, to regard Datana and Pygiera as the most primitive forms of Notodontians, the smooth-bodied larvie of Gluphisia being secondary and adaptive forms. 2R. Marey: Le transformisme et la physiologie expérimentale, Cours du Collége de France. Revue scientifique, 2° Série, iy, 818. (Function makes the organ, especially in the osseous and muscular systems. ) See also A. Dohrn: Der Ursprung der Wirbelthiere und das Princip des Punctionswechsels, Leipzig, 1875. ‘It is possible that the close resemblance of the warts, projections, and spines of certain arboreal caterpillars which so closely mimie the spines, leaf scars, and projections of the branches or twigs or plants, has been brought about in a way analogous to the production of spots and lines on the body of caterpillars. Darwinians attribute this to the action of ‘“‘protective mimiery,” but this expression rather expresses the result of a series of causes to which we have endeavored to call attention. The effect of dark and light shades and the light and shade in producing the stripes and bars of caterpillars are comparatively direct and manifest; but how can thorns and other projections on trees and shrubs affect caterpillars directly? Given the origination by hypertrophy of warts and spines, and it is then easy to see that by natural selection caterpillars may have finally become adapted so as to mimic similar vegetable vrowths. Our object is to endeavor to explain the causes of the primary growth and development of such projections, i. e., to lay the foundation for the action of natural selection. MEMOIRS OF THE NATIONAL ACADEMY OF SCIENCES. 21 a The following table is an attempt at a classification of some of the structures arising from the various modifications of the primitive piliferous warts or tubercles common to nearly all, if not all, smooth-bodied lepidopterous larvee. As is well known, the term “hair” does not properly apply to the bristles or hair-like structures of worms and Arthropoda, as morphologically they are not the homologues of the hairs of mammals, but arise, as Newport first showed, through a modification and hypertrophy of the nuclei of certain cells of the cuticle. Hence the word seta, as suggested by Lankester, is most applicable. A.—TUBERCLES. a. Simple and minute, due to a slight thickening of the hypodermis and a decided thickening of the over- lying euticle; the hypodermis contains a large unicellular gland, either for the secretion of the seta or for the production of poison, . Minute piliferous warts. (Most Tineid, Tortricid, and Noctuid larvee. ) . Enlarged smooth tubercles, bearing a single seta. (Many Geometrid and Bombycine larvee.) . Enlarged spherical tubercles, bearing a number of setie, either radiated or subverticillate. (Aretians, Lithosians, Zygenide, including some Glancopinee. ) . High, movable, smooth tubercles, having a terrifying function. (Schizura, Xylinodes, Notodonta, Nevice.) 5. Low and broad, rudimentary, replacing the “caudal horn.” (Charocampa, the European Pheosia dictwa, and dictwoides. ) b. More or less spinulose or spiny (disappearing in some Sphinges after Stage I). 1. Long and slender, usually situated on top of the eighth abdominal segment, with microscopic spinules in Stage I. (Most Sphingidie and Sesia.) 2. Smooth subspherical warts. (Zygenidie, e. g., Chaleosia, East Indies); or elongated, but stillsmooth. (Altacus atlas, and a species from Southwestern Territories, U. 8. A.) . Subspherical or clavate spiny tubercles of many Attaci; the spinules usually short. . Spinulated spines or elongated tubercles of Ceratocamipd and Hemilucidw. (H. io and H. maia, etc.) . Spike-like hairs or spines, (Samia cynthia, Anisota, East Indian Hypsa, Anagnia.) . Antler-like spines. Early stages of Heterocampa biundata, guttivitta and obliqua. ) who St hm Oo o B.—Sere (‘‘Hairs,” BRISTLES, ETC.). 1. Simple, fine, short or long, microscopic or macroscopic sete, tapering hairs, scattered or dense, often forming pencils. (Many Bombyces, Zygienide, Noctuo-bombyces, Apatelie ) . Glandular hairs, truncate, spindle-shaped or forked at the end, and secreting a more or less viscid fluid. (Many in Stages I and IT of Notodontians, many butterfly larve, and in the last stages of Pterophoridie.) 3. Long, spindle shaped hairs of Apatelodes, Apatela americana, figured in Harris Corr., Pl. II, fig. 2; also Packard’s Guide, fig. 236, and the European Tinolius eburneigutta Walk. 4, Flattened, triangular hairs in the tufts or on the sides of the body of Gastropacha americana, or flattened, spindle- shaped seales in the European G. quercifolia.' 5. Spinulated or barbed hairs. (Most Glaucopides, ete., Arctians, Lithosians, and Liparidie, and.many other Bombyces. ) bo C.—PSEUDO-TUBERCLES. The filamental anal legs (stemapoda) of Cerura and Heterocampa marthesia. 2. The long suranal spine of Platyptericidie. THE USUAL POSITION OF THE MORE SPECIALIZED WARTS, HUMPS, OR HORNS. Everybody has noticed that the horn characteristic of larval Sesiv and Sphinges is uniformly situated on the back of the eighth abdominal segment and no other, and that when it is absent, as in Choerocampa, ete., it is replaced by a small, low, and flattened tubercle, the segment itself being somewhat swollen. The larval Agaristidie (Alypia, Eudryas, Copidryas, Psychomorpha, ete.) have a prominent, gibbous hump on this segment, or at least this segment is more or less prominent and humped, not only in this family, but also in certain smooth-bodied Noctuidie, as Amphipyra, and Olygia versicolor, ete. 7 In many Notodontide the first abdominal segment bears a conspicuous hump, sometimes forked, often ending in a seta. In the larval Ceratocampidie, either the prothoracic segment or the second and third thoracic segments bear high conspicuous horns and spines. They may be roughly classified as follows: ‘See my article in Annals and Magazine of Natural History. Ser. 6. ix. pp. 372-875. 1892. 299 MEMOIRS OF THE NATIONAL ACADEMY OF SCIENCES. Prothoracie segment.—With a large subspherical tubercle on each side bearing numerous radiating hairs (Lasiocampide of first stage) or pencils of hairs (Parorgyia); two antlers (H. guttivitta, biundata and H. obliqua). Second thoracic segment.—Two high slender spines. First stage of Anisota senatoria, A. stigma, and Dryocampa rubicunda. Third thoracic segment.—Two spinulose pappose flaps, Empretia stimulea. First, second, and third thoracic segments—Each with a pair of high spines, Citheronia regalis and Hacles imperialis. Second and third thoracie segments. —Bach with a pair of long horns, Sphingicampa bicolor. First and third thoracic segments.—In Stage I of the European Aglia tau (Poulton). First abdominal segment.—Movable tubercle in Schizura and Xylinodes. Bighth abdominal segment.—The caudal horn of Sesia and most Sphingidwe, Pheosia, and Endromis, Bombyx mori, and other species—Sphingicampa, Eacles, Citheronia, and Aglia tau (Stage I). So far as I am aware no one has suggested why these horns and high tubercles, and often pencils of hairs, are restricted to these particular segments. As a partial explanation of the reason it may be stated that the presence of these high tubercles, etc., is correlated with the absence of abdominal legs on the segments bearing the former. It will also be noticed that in walking the apodous segments of the caterpillar are more elevated and prominent than those to which the legs are appended. They tend to bend or hump up, particularly the first and the eighth abdominal, the ninth segment being reduced to a minimum, and the tenth simply represented by the suranal and paranal plates, together with the last pair of legs. As is well known. the loopers or geometrid worms, while walking, elevate or bend up the part of the body situated between the last thoracie and first pair of abdominal legs, which are appended to She seventh uromere. Now, in the larva of Nematocampa filamentaria, which bears two pairs of remarkable filamental tubercles rolled up at the end, it is certainly very suggestive that these are situated on top of the loop made by the caterpillar’s body during progression, the first pair arising from the second and the hinder pair from the fourth abdominal segment. It seems, therefore, that the humps or horns arise from the most prominent portions of the body, at the point where the body is most exposed to external stimuli; and the force of this is especially seen in the conspicuous position of those tubercles which are voluntarily made to nod or so move as to frighten away other creatures. Perhaps the tendency of these segments to loop or hump up has had a relation of cause and effect in inducing the hypertrophy of the dermal tissues entering into the composition of the tubercles or horns. Analogous positions are in the vertebrates utilized, as in spiny, osseous fishes, or the sharks, the horned Amphibia, or horned reptiles and horned inammals. The prominence of the foundation parts, from which the tubercles arise, may lead to a determination of the blood toward such places, and thus in well-fed or overfed (possibly underfed) individuals induce a tendency to hypertrophy, which once set up in early generations led to the production of incipient humps which became more developed as they proved useful and became preserved in this or that form by natural selection. On the other hand, the hypertrophy of certain piliferous warts would tend to cause an arrest of development or a tendency to atrophy in the piliferous warts of adjoining segments, And in like manner may the simple set have become hypertrophied on account of their great utility as deterrent organs, and become wonderfully modified in this and that direction in such and such forms, until they became in recent geological times the common and normal inheritance not only of scattered species but of certain genera in scattered families, and even of entire families. It is to be observed, as one will see by referring to the special larval histories and the recapit- ulations which we have appended, that in the species of Schizura the evolution or hypertrophy of the movable or nutant tubercles begins in the third stage at about the time when the young caterpillars leave their common birthplace on the underside of the leaf and seek more conspicuous feeding grounds on the outer edge or on the upper side of the leaf, where they are exposed to the visits of ichneumons, or Tachine, or carnivorous Hemiptera, or to the onset of open-mouthed insectivorous birds. At the same time arise the bright colors, spots, and stripes, the very peculiar v-shaped silver or yellowish-white mark characteristic of the species of Schizura—these are per- MEMOIRS OF THE NATIONAL ACADEMY OF SCIENCES. 23 haps danger signals—though later in life the brown shades and green tints, so like the green leaf with its serrated, blotched, sere-patched edges, would often deceive the most obscrvant of birds. In regard to the nutant.or movable tubercles, it may be observed that a slight motion of these appendages may suffice to scare off an approaching ichneumon or Tachina. If most insects have, as supposed by Exner and by Plateau, more imperfect vision than has formerly been attributed to them, so that they are extremely nearsighted and only clearly perceive bodies when in motion, then even slight movements of these tubercles, while the caterpillar itself was immobile, would probably be sufficient to frighten a parasitic insect and deter it from laying its eggs on the caterpillar. GROUPING OF NOTODONTIAN LARV4 ACCORDING TO THEIR AFFINITIES AND ALSO THEIR ADAPTATION TO ARBOREAL LIFE. As is well known, the larvee of this family vary greatly in form and ornamentation for a group of such moderate numbers; and the following synopsis has been prepared in order to show this great variety in as graphic a manner as possible: 1. Body smooth, not hairy, with red and yellow spots. Gluphisia. 2. Body smooth, moderately hairy. Datana. 3. Very hairy, the body almost totally concealed. Apatelodes. 4. Body smooth, hairless; with no humps or tubercles, of a noctuid shape; anal legs never elevated; color green, with yellow lines, the latter sometimes edged with reddish; feeding less conspicuously than any others of the family. Nadata, Lophodonta, ete. 5. Body with two dorsal tubercles; also hairy. Ichthyura. 6. Body smooth, polished; a single hump, surmounted by a horn on the eighth abdominal segment. Pheosia. 7. Back 2-8-humped, serrate, body smooth, not brightly striped. Notodonta, Nerice. 8. Body smooth, gayly striped, eighth abdominal segment gibbous. Hdema, Dasylophia. 9. Body smooth, with nutant tubercles on first and eighth abdominal segments; end of body uplifted. Colors green with brown patches simulating dead blotches on leaves. Hyparpaa, Schizura, and Nylinodes. 10. Body with stout spines and with spiny tubercles on first and eighth abdominal segments. Schizura unicornis. ; 11. Body smooth, tapering; anal legs normal, often with two prothoracic tubercles, enormous in early stages. Heterocampa guttivitta, biundata, and obliqua, 2. Body smooth, striped; anal legs normal. Heterocampa manteo. 13. Body with two dorsal prothoracic tubercles; anal legs filamental; each ending in an eversible flagellum. Macrurocampa marthesia. 14. Body with two lateral prothoracic tubercles; anal legs filamental, each ending in an eversible flagellum. Cerura. 15. Body doubly humped on the abdominal segments; filamental anal legs. The Old World genus Stauropus. So far as I have gone in the examination of the structure of the moths, this succession of genera roughly corresponds with the classification of the family. Judging by the moths alone. Datana stands at one end of the series and Cerura at the other. Perhaps Cerura has generally been placed at the end of the group because of its fancied resemblance to the larva of Drepana, but this is deceptive, because the long caudal filament of the latter genus is simply a hypertrophy of the suranal plate, and the anal legs themselves are atrophied, while in Cerura they are enormously hypertrophied, probably owing to their active use as deterrent appendages. SUMMARY. One would suppose that the two genera Nadata and Lophodonta, with the Old World genera Pterostoma, Ptilophora, Drymonia, Microdonta, and Lophopteryx! (of the two species L. eucullina, which is humped on the eighth abdominal segment, connects with the plain-bodied L. carmelita 1 The first larval stages of the following genera are still unknown, and the author would be much indebted for eggs or alcoholic specimens of the larve of the first and later stages: Ellida, Lophodonta, Drymonia, Notodonta, 24 MEMOIRS OF THE NATIONAL ACADEMY OF SCIENCES. and the above-mentioned group, Pheosia, Leiocampa) should properly, by their smooth, noctui- form shape, stand at the bottom of the family, as being nearest related to the primitive forin of the group. But until we know more of the earliest stages it is best to suspend our judgment. 1. The more prominent tubercles and spines or bristles arising from them are hypertrophied piliferous warts, the warts with the seta or hair which they bear being common to all caterpillars. 2. The hypertrophy or enlargement was probably primarily due to a change of station from herbs to trees, involving better air, a more equable temperature, perhaps a different and better food. 3. The enlarged and specialized tubercles developed more rapidly on certain segments than others, especially the more prominent segments, because the nutritive fluids would tend to more freely supply parts most exposed to external stimuli. 4. The stimuli were in great part due to the visits of insects and birds, resulting in a mimicry of the spines and projections on the trees; the colors (lines and spots) were due to light or shade, with the general result of protective mimicry or adaptation to tree life. 5. As the result of some unknown factor several of the hypodermic cells at the base of the spines became in certain forms specialized so as to secrete a poisonous fluid. 6, After such primitive forms, members of different families, had become established on trees, a process of arboreal segregation or isolation avould set in, and intercrossing with low feeders would cease. 7. Heredity, or the unknown factors of which heredity is the result, would go on uninter- ruptedly, the result being a succession of generations perfectly adapted to arboreal life. 8. Finally the conservative agency of natural selection would operate, constantly tending toward the elaboration and preservation of the new varieties, species, and genera, and would not cease to act in a given direction so long as the environment remained the same. 9. Thus, in order to account for the origin of a species, genus, family, order, or even a class, the first steps, causing the origination of variations, were in the beginning due to the primary (direct and indirect) factors of evolution (Neolamarckism), and the final stages were due to the secondary factors, segregation and natural selection (Darwinism), F IIl.—ON CERTAIN POINTS IN THE EXTERNAL ANATOMY OF BOMBYCINE LARVA. Homology of the “flagellum” of Cerura, ete., with the planta of the other abdominal legs.—We have in a former! article, in describing the larvee of Macrurocampa marthesia and of certain species of Cerura, called attention to the nature of the stemapoda?® or filamental legs of those caterpillars, and their generally undisputed homology with the anal legs of other Notodontians. P]. XXX VII, fig. 9, represents the anal legs of Dasylophia anguina in its first larval stage. It is intermediate in form between the normal leg and the stemapod. It has no ecrochets, but the planta, of which the “flagellum” of Cerura and H. marthesia seems to be the homologue, is retracted and the retractor muscles, one of which is divided, are much as in the filamental legs of Cerura, ete. Note on the modifications in the tenant or glandular hairs of the thoracic feet.—As is well known, the thoracic feet of caterpillars are five jointed and end in a single claw, with apparently a rudimentary one at the base. Usually, besides the unguis or claw, there is a tenant hair, which is generally spine like, but besides these appendages there are sometimes more or less flattened, lamellate setee, which are curious and worthy of notice. In Parorgyia parallela, besides the unguis ' Proceedings Boston Soc, Nat. Hist., xxiv, 1890. *The term ‘‘tails” or eandal filaments is too vague for these highly modified anal legs; hence we propose the term stemapoda or stemapods for those of Cerura and Macrurocampa. The derivation is Gr. cryyua, filament, zoic, rode, leg or foot. Mr. J. Hellins, referring to these organs in Buckler’s Larvie of the British Butterflies and Moths (Roy. Soe., ii, 138), remarks: “ But now through Dr. T. A. Chapman’s good teaching, I regard them as dorsal appendages, somewhat after the fashion of the anal spines of the larvie of the Satyridie.” This, I am satisfied, is an error. After repeated comparisons of the filamental anal legs of Cerura with those of Macrurocampa marthesia, and comparing these with the greatly elongated anal legs of young H. unicolor as figured by Popenoe, and taking into account the structures and homologies of the supraanal and paranal flaps, one can scarcely doubt that those of Cerura are modified anal legs. MEMOIRS OF THE NATIONAL ACADEMY OF SCIENCES. 25 and the spine-like tenant hair, there is a lamellate, flattened hair. Pl. XX XVII, fig. 10, represents the end of a thoracic leg of Heterocampa manteo. Besides the unguis and tenant hair at the end, there are two singular, thin, flattened, oval leaf-like sete arising near the middle of the joint. The use of the claw and tenant hair as grappling organs is quite apparent, but the function of the singular lamellate hairs is a matter of conjecture. Hints on the origin of the prothoracic or cervical shield.—Not only in the wood-boring Lepi- doptera, such as the larvie of the Hepialidie, and the Cossidie, as well as the Sesiide, is there a well-marked cervical shield, but also in the grubs of Cerambycidie, and some other Coleopterous families whose larvie bore in hard substances, and in such groups this hard, chitinous plate serves to protect the base of the head and adjacent parts of the body most exposed to injury. Developed in the borers of widely different orders, and obviously of direct use to the animal, it has probably arisen in response to an external stimulus, an extra quantity of chitin having been developed by the hypodermal cells of the tergal arch of the prothoracic segment, which by friction has become thickened, just as the skin of the sole of the foot in savages becomes thick and horny in those accustomed to go barefoot in dry, rough places. In the lower lepidopterous families, as the Tineina, Tortricidie, Pyralide, as well as in the low-feeding Noctuids, which hide under stones, sucii as the cutworms, a well developed cervical shield is génerally present. In the Bombyces, which feed exposed both on trees and on herbaceous plants, the cervical shield is rarely even well developed, but there are sporadic cases of its development, and especially of its appearance in the early stages and of its suppression in later larval life, which are of interest and merit notice. In the Notodontian genus Cerura, the prothoracic segment is unusually broad and flat above, although it is not smooth, chitinous, or polished; whether its use is to support the large lateral tubercles or to resist pressure and friction is a question. In the first stage of Dasylophia anguina there is a small cervical shield (PI. XX XVII, fig. Le), which bears four glandular sete on each side of the median red dorsal line. In Datana integerrima, a small, transversely oblong, conspicuous black cervical shield is present in the freshly hatched larva and in the subsequent stages. There is, however, no shield or rudiments of one in Edema albifrons or in Heterocampa and Macrurocampa. In the other Bombyces there is no genuine shield, but in the first stage of some forms the two dorsal piliferous warts on the prothoracie segment are more or less enlarged and sometimes coalesced so as to indicate that the shield may have been formed by the enlargement and coalescence of these warts. The supraanal or suranal plate—This plate, the poder of Kirby and Spence, in Bombycine and Geometrid larvie, both as to its shape and ornamentation, affords excellent characters for distinguishing species, and we have found it of great use, especially in describing Geometrid caterpillars. It varies much in shape and ornamentation in Notodontid, also in Attacide .and Ceratocampide. In Noctuide it is not, so far as we know, very characteristic. It seems to be especially developed in those larve which constantly use the anal legs for grasping, while the frout part of the body is more or less raised. It is thus correlated with enlarged anal legs. Morphologically this plate appears to represent the dorsal arch of the tenth or last abdominal segment of the body,! and is the “anal operculum” or lamina supraanalis of different authors.” This suranal plate is in the Platyptericide remarkably elongated, forming an approach to a flagellum-like terrifying appendage, and in the larva of Aglia taw forms a long, prominent sharp spine. Its shape also in Cerura caterpillars is rather unusual, being long and narrow. In the Ceratocampide, especially in Anisota, Dryocampa, Eacles, and Citheronia, this plate is very large, the surface and edges being rough and tuberculated, while it seems to attain its Inaximum in Sphingicampa, being triangular, oe in a bifid point. ‘See my note, “‘The number of abdominal segments in Hepdupte rous larve 1885, pp. 307, 308. 2Compare E. Haase, ‘On the constitution of the body in the Blattide.” Ann. and Mag. Nat. Hist., March, 1890,. 297-234. Translated from Sitzungsb. Ges. Naturf. Freunde zu Berlin, Jahrg., 1889, 128-136. fees ican Naturalist, March, 26 MEMOIRS OF THE NATIONAL ACADEMY OF SCIENCES. The ninth abdominal segment is unusually well developed in the Attacidie and the Cerato- -campidie, sometimes, as has been previously stated, bearing a true “caudal horn,” which takes the place of that usually growing on the eighth segment. In the Rhopalocera, the suranal plate is in general, especially in Hesperidw and Papilionidie, small and rounded, much as in the Noctuidie, but in the Nymphalide it is more or less specialized, and remarkably so in the larva of Neonympha phocion and other satyrines, where it is greatly elongated and forked. (See figures in Seudder’s “ Butterflies of New England;” also W. Miiller’s figures of larva of Prepona.) The paranal lobes.—These are the homologues of the two anal valves (valvule of Burmeister, “the podical plates” of Huxley) observed in the cockroach, and occurring in nearly all, if not all, insects. In Geometrid larve they are full, fleshy, lobe-like, or papilliform, bounding the areas on each side, and appear as if projecting backward from the base of the anal legs. In the Ceratocampide these paranal lobes are not well developed. In the larva of Cerura they are much as in Geometrid caterpillars, where they end each in a seta. The paranal forks.—We have already called attention to these two bristles in our description of the larve of Cerura. (Proceedings Boston Soc. N. H. xxiv, p. 553.) They are well developed, arising from the end of a papilla projecting directly backward. Their use has been indicated by Mr. John Hellins,! who refers to a pair of sharp points underneath the anal flap, ‘which are used to throw the pellets of frass to a distance.” Occurring in Notodontian and other arboreal caterpillars, notably the tree-inhabiting Geometrids, they are wanting in Noctuide (including Acronyeta and Catocala), Sphingidie, and Rhopalocera, as well as the lower Geometrids and the Microlepidoptera, and are not developed in the Sphingidie. In Ichthyura (Clostera) they are slightly developed. In the European Urapteryx sambucata (received from M. P. Chrétien) these lobes are very large, papilliform, and setiferous, and in our Cheerodes, ete., they are similarly developed and the use of the two set or the fork is undoubtedly the same as in Cerura. : The infraanal lobe.—My attention was first called to this lobe or flap while examining some Geometrid larvee. It is a thiek, conical, fleshy lobe or flap, ending often in a hard chitinous point, and situated directly beneath the vent. In appearance it is somewhat like the egg-guide of the Acrydii, though the latter is thin and flat. Jts use is evidently to aid in tossing the pellets of excrement away so as not to allow them to come in contact with the body. In a large not iden- tified Geometrid worm, which lives on the ash, this flap is large and conical, ending in a blunt chitinous point. In a large geometer belonging to another genus, the tip is sharper and harder, and in what is probably a larva of Endropia, while the paranal forks are well developed, the infraanal lobe ends in a stiff bristle. Whether this infraanal lobe is the homologue of the ninth urosternite or ventral plate [ will not at present undertake to say. Glandular sete.—Among the Notodontide the freshly hatched larvie of several genera are provided with glandular hairs of various shapes. In Datana integerrima they are clavate; in Dasylophia anguina they are clavate, somewhat flattened, and are dark, but clear at the tip,’ while in all the other caterpillars we have observed that the glandular hairs are confined to the body, those on the head tapering to a point, and apparently not fitted for secreting a fluid; those on the head of Dasylophia are glandular, all ending in a slight transparent bulb. Other genera of this group will probably on further investigation be found to possess glandular sete in their first larval stages. They occur in the freshly hatched larva of what is probably a species of Heterocampa, also in Nadata gibbosa, Ichthyura inclusa, and Pheosia rimosa. It is to be observed that the freshly hatched eaterpillars of Ceratosia tricolor Smith are provided with glandular hairs. They are flattened at the tip, which is slightly tridentate, with 1'The use of these I find explained by Mr. Hellins in his description of the larva of C. bifida in Bueckler’s Larvie of British Butterflies and Moths, ii, p. 142, as follows: ‘‘At the tip of the anal flap are two sharp points, and another pair underneath, which are used to throw the pellets of frass to a distance.” Similar dungforks are very generally present in Geometrid larvie, the paranal papilliform tubercles being well developed, though we have not seen them in use. I have noticed a caterpillar of C. borealis in the process of defecating, and with the forceps pulled off a pellet which was held by the two spines of the paranal tubercles. Mr. Dyar tells me he has both seen and heard the caterpillars casting their pellets with the aid of their spine against the side of a tumbler. 2PILXXXVII, fig 11. Glandular hairs of Dasylophia; a, of body; b, of the head; ¢, of prothoracie shield. MEMOIRS OF THE NATIONAL ACADEMY OF SCIENCES. 27 grooves passing down the shaft from the notches between the teeth. They occur not only on the back and sides of the body segments, but also on the sides of the abdominal legs. The occurrence of such hairs in this genus is interesting from the fact that they have not yet been observed in Aretians, to which this moth has been referred, nor in the Noctuide, among which it should be placed, since no Arctians have when hatched smooth glandular hairs.! I1V.—ON THE INCONGRUENCE BETWEEN THE LARVAL AND ADULT CHARACTERS OF NOTODONTIANS. As is well known to zoologists, from the writings of Fritz Miiller and later students, in groups of animals which generally undergo a metamorphosis, two or more species of the same genus may differ remarkably in respect to their early life, one species passing through a complicated meta- morphosis while a closely allied form has a direct development, hatching in the form of the adult. The embryo, however, in the latter case rapidly passes through a series of changes, constituting a premature, abbreviated, or condensed metamorphosis, epitomizing the ordinary early stage of its metamorphic allies. Thus the lobster differs from the other marine macruran crustacea in having a condensed metamorphosis before hatching from the egg, rapidly passing through a nauplius and a zoéa phase. Itis so with some crabs. All the fresh-water Decapoda, notably the crayfish, have no postembryonic metamorphosis. The fact that the embryo exhibits a condensed metamorphosis shows their origin from metamorphic forms. These are perhaps the most remarkable cases of incongruence between what may be closely allied genera and even species. Also two allied species of Gammarus may differ in toto as regards the mode of segmentation of the yolk, total cleavage occurring in one marine species (G@. locusta) and partial or peripheral cleavage in two fresh-water forms (G@. pulex and fluviatilis). Examples of such great divergences in larval or early life, or in the condition in which the animal is hatched, in species closely similar in adult life, are not uncommon in worms, Echinoderms, Molluses Soweecen besides insects, and the phenomenon is with little doubt due to the changed Poaditions of the environments to which forms with such exceptional modes of development have been exposed. The principle, then, of divergence or incongruence of larval charactersin forms whose adults are closely allied has been established in the lower classes of Metazoa. The most remarkable and puzzling case, perhaps, is that of Balanoglossus, whose Tornaria larva is so much like that of Echinoderms, while the adult is a protochordate animal. As a matter of fact this does not affect the classification of these animals. Zoologists have not thrown forms with a direct development into distinct groups where the adults have not shown any differences; at the same time no one would unite the two species recognized as such which presented no easily observed differences if one had a direct and the other a metamorphic development. In the present state of our knowledge it may be well to at least provisionally mark the differences between the two forms, so divergent in their early life, by giving them distinct names, and thus emphasizing the fact that of the two closely allied forms one has diverged from the other through having been subjected to a different set of external influences, whatever such conditions may have been. Systematic zoology has undergone within the last thirty years an entire change. Our present systems of classification are now attempts to arrange animals in the order of their probable appearance, 1. e., phylogenetically, and as the subject is yet in its infancy, and our attempts provisional and tentative, we are obliged to give great weight to any differences in the larval conditions of animals with a metamorphosis, because such differences were undoubte:dly due to differences in the environments of their parents. Indeed if it had not been owing to changes in the physical and biological environment, animals would never have risen beyond the dead level of the lowest Protozoa. Such reflections as these and a knowledge of the mode of development of the lower classes of invertebrates are all-important to the students of insects, especially of the metamorphic orders, UPS XXXVII, fig. 12. Gieadaiee hairs of Ceratosia tricolor, a, from the Poel thoracic and fact erdgenna seg- ment; b, those on the first and second abdominal legs. 28 MEMOIRS OF THE NATIONAL ACADEMY OF SCIENCES. Neuroptera, Coleoptera, Mecoptera, Trichoptera, Lepidoptera, Diptera, and Hymenoptera, where there are so many and perplexing cases of incongruence or divergence in larval forms whose parents are very closely allied. It is worthy of notice that in respect to Diptera the veteran dipterologist, Baron R. von Osten Sacken, remarks of the nemocerous flies: “An arrangement of the imagoes based upon such prin- ciples will of necessity be justified by a more or less tangible correspondence in the characters of their larvie. This structural correspondence, this parallelism of larvee and imagoes among the Nemocera, suffers, as far as I know, but one exception, Mycetobia pallipes and Rhyphus. In both almost identical larve produce flies belonging to different families.” (Berliner entomolog. Zeit- schrift, Bd. xxxvii, 1892, Heft iv, p. 418.) In the copy kindly sent me by the author a second case of Anopheles and Dixa is mentioned in the printed copy, but struck out by the author in the emended copy. Eyeryone is familiar with the fact that there is a nearly similar incongruity between the larve of the Muscide and the flies. Many new facts bearing on this subject appeared in Portchinsky’s article on the habits of the necrophagous and coprophagous larvie of Muscidee, of which an English abstract by Baron R. von Osten Sacken appeared in the Berliner ent. Zeitschrift for 1887. After speaking of the wonderful power of adaptation of these larve to their environment, he states: Distinctly related species belonging to different genera issue from larvie almost indistinguishable from each other. And again closely related and almost indistinguishable imagoes, species of the same genus, differ in their oviposition (size and number of eggs), and their larve follow a different law of development (as to the degree of maturity the larva reaches within the body of the mother and the number of stages of development it passes through). In one case even (Musca corvinw) larvie of the same species were found to have a different mode of development in northern and southern regions of Russia. Here also it is evident that the cause of the incongruity is due to the fact that the larvae, for the time being different animals from the adult, are modified by their environment, the similar surroundings and habits of the larvie of quite different genera causing the larve externally at least to closely resemble each other. Whether they are so similar in their internal organs remains to be seen. Dr. C. W. Stiles, who has studied so carefully by microscopic sections tapeworms of externally similar form, and which can not be separated by external characters, tells me that the internal organs seem to afford excellent specifie and generic characters. Lepidopterists in general do not hesitate to base their systems of classification on the larval as well as adult features. They in general regard their systematic arrangements of the imagines as more or less provisional, and all acknowledge that it is immensely satisfactory, even after they are pretty well satisfied with th eir arrangement of the adults of a group, whether a genus or family, to work out the larval stages and to check their classifications based on adult features by the larval characters. In many cases they may be led to change the position of a species or genus, or to split up a genus or species. But, after all this, the fact that so many larvee, even in the same group, are hatched with such different shapes and characters; the fact that some are so much more simple and primitive than others, opeas up most perplexing yet interesting questions and problems. We may, however, be able to solve these, and in the present group of Bombyces it seems to us that the different larval forms, some primitive and geueralized and others more or less modified or specialized, give clues. to the phylogeny of the groups which we confess we had not expected. And in this memoir we have endeavored, though often it is mere guesswork, to drop the old- time method of putting the type species first and then ranging the others after it in an ill-assorted group, and have attempted to begin with what has seemed to us to be the ancestral form of the group, following with the later forms. This can be best accomplished by taking into consideration the caterpillar, beginning with the generalized forms and ending with the later more modified or specialized forms. In such a large genus as Heterocampa this is not difficult to do. For example, as we shall see hereafter, the larva of H. manteo is as simple and generalized as any, while that of H. wnicolor is the most modified, with its semi-stemapoda, from which Macrurocampa, with its fully formed stemapoda, may have descended. And then, while Cerura, with its stemapoda alike in all the species, is often or generally placed first in the group, it is evident that it was descended from some Heterocampa-like form through Macrurocampa. Aided by our knowledge of MEMOIRS OF THE NATIONAL ACADEMY OF SCIENCES. 29 the larval forms, especially of the earliest stages, it is not difficult to construct a genealogical tree of the subfamilies Heterocampine and Cerurine. When taking into account the larval stages of the entire family, even with our present imperfect knowledge, it is easy to see that Datana stands at the base, is the more generalized primitive form, and was perhaps the first to diverge from the stem-form of the family. The first author to cali attention and at the same time to treat in a philosophic way of what he has ealled “the incongruence of form relationship, between larvie on the one hand and imagines on the other” is Weismann, in his well-known work entitled Studies in the Theory of Descent. In Chapter II of the second volume, entitled “Does the form relationship of the larva coincide with that of the imago?” he points out certain incongruences between the larval and adult characters. Ee claims that “neither the group of Microlepidoptera nor those of the Noctuina, Bombycina, Sphingina, and Rhopalocera can be based systematically on larval characters,” adding the quali- fication, ‘Several of these groups are indeed but indistinctly defined, and even the imagines present no common characteristics by which the group can be sharply distinguished.” Within the families, however, he states: “There there can be no doubt that in an overwhelmingly large majority of cases the phyletic development has proceeded with very close parallelism in both stages; larval and imaginal families agree almost completely. On the other hand, ‘in the butter- flies a perfect congruence of form relationship does not exist, inasmuch as the imagines constitute one large group of the higher order, whilst the larvie can only be formed into families.” But in this case Weismann does not seem to be aware that the imaginal Rhopalocera as such is quite an artificial group, and that the imaginal families recognized by Bates, Scudder, and others have perhaps more equivalent, congruent, or nondivergent larval forms than his remarks would seem to imply. But without attempting to enter into an exposition or criticism of Weismann’s general statements, his whole discussion being most suggestive and stimulating, we will turn to what he says of the Notodontide : An especially striking case of incongruence is offered by the family Notodontide, under which Boisduval, depending only on imaginal characters, united genera of which the larve differed to a very great extent. In fact, in the whole order Lepidoptera there can scarcely be found associated together such diverse larvie as are here placed in one imago family. He then refers to the short cylindrical caterpillars of Cnethocampa, which, however is not a Notodontian, but a Lasiocampid. He then briefly refers to the larvee of Harpyia (Cerura) and the saterpillars of Stauropus, Hybocampa, and Notodonta. Without giving further attention to the family, he returns to the butterflies. This family, then, presenting ‘‘an especially striking case of incongruence,” we will briefly discuss, referring the reader for fuller details to the figures on the plates. In the first place, as a matter of fact, the more one becomes familiar with the Lepidoptera and their larval forms the easier it is to distinguish the larvie by their ‘‘ family” characteristics, premising, however, that the term family is of very uncertain meaning, and that different authors differ as to what to call a family as much as they do what to designate a species. But no one, we think, need to err in correctly picking out or identifying any Bombycine larva except, perhaps, a few Notodont laryie, which are liable to be confounded with certain Thyatiride, and the hairy Noctuidie, but even then a careful examination will show family differences even when adaptation and modification have nearly bridged over the fundamental differential characters. In this work I have divided the family into seven groups, which may be for convenience regarded as so many subfamilies. I was first led to do so by the larval characters alone, but found that this classification would also apply in general to the moths, so that there proved not to be so much incongruity as was expected. There appear to be, then, seven larval subfamilies and seven imaginal subfamilies. Others may not agree with this view, but it is the most rational classification I have been able to make. Beginning with the most simple forms of larva, those of the Gluphisine, which, both as regards those of the Old and New World, are tolerably constant, the adults certainly differ notably from those of other subfamilies, as also do the larvee and pupie. 30 MEMOIRS OF THE NATIONAL ACADEMY OF SCIENCES. The remarkably woolly and penciled larve of Apatelodes are congruous with the very distinct imagines of the subfamily Apatelodine, which are so well defined by their structural characters. The hairy and brightly banded larve of the Pygerine, so unlike those of other Notodontians, are paralleled by the general appearance and structure of the moths, so much so thatthe group was regarded as a distinet family (Pygieride) by Duponchel. The larve of the European Pygierinie are hairy and gaily striped, and related in much the same way to our larval Datana as the imago is to our imaginal Datana. The larvee of the subfamily Ichthyurine, represented by only a single genus, need not be confounded with those of any other division of the family, though there is a great deal of plasticity within the limits of the group. The most generalized species is the larva of I. apicalis (vaw) and its allies brucei (multnoma), since it has no large specialized tubercles like those of inclusa and albosigma, and the latter species differs, both as regards larva and imago, from J. inclusa. The incongruence in this group is not greatly emphasized. But in the two next subfamilies there is a striking lack of congruity between the larva and moth, both in the genera and species. Among the Notodontinz we have Hyparpax, whose imago is so different, in the shape of the wings and in the color of the body and wings, compared with any other genus of the group or even of the family; yet the larva is very nearly allied to those of Yylinodes and of Schizura. A remarkable case of incongruence is the larva of Schizura concinna. This well-known caterpillar, with its formidable armature of long hobnail-like spines and its gay head and swollen coral-red dorsal hump, would seem to be the type of a distinct genus, and yet from a study of its adult character it is not separable from the other species of Schizura, and we have dropped the genus (@demasia we originally proposed for it from the lack of stable differential characters. The freshly hatched larvie, however, is undistinguishable from that of other Schizurie yet known, and perhaps we have done violence to the principles of classification in not allowing it to remain in the genus we originally proposed for it. At all events, it with other Schizurie evidently had a common pareutage, and it has diverged since it first molt farther away from the stem than others of its cospecies and may be regarded as an incipient genus. It is also plain that the causes which have acted upon this organism have from the first been of a quite different nature from those which have been efficient in causing fixed variations in other directions, resulting in the fixation of the other species of the genus.