NG Nua bi LI Pa ay oY EGP, VAS RS Ly 55 bf 4 FD) Z =, 7 Aan) 4 NOVITATES ZOOLOGICAE. Vol. IX. SUPPLEMENT. a’ -& REVISION | | OF THE -LEPIDOPTEROUS FAMILY | SPHINGIDAE. BY THE Hon. WALTER ROTHSCHILD, Pu.D., AND KARL JORDAN, M.A.L., PH.D. (WITH 67 PLATES.) | ISSUED AT THE ZOOLOGICAL MUSEUM, TRING, APRIL 1903. | PRINTED BY HAZELL, WATSON & VINEY, Lp., LONDON AND AYLESBURY. 1903. ~“ ™~ A REVISION OF THE LEPIDOPTEROUS FAMILY SPHINGIDAE. Novirares Zoontocicr. Vor. IX. Suppeemen’. A REVISION OF THE LEPIDOPTEROUS FAMILY SPHINGIDAE. Hon. WALTER ROTHSCHILD, Pu.D.., AND KA £- z KARL JORDAN, M.A.L., Px.D. Y ne (WITH G7 PLATES.) IssuED Av tnk ZooLrocicAL Museum, Trine, Maren 1903, PRINTED BY HAZELL, WATSON & VINEY, Ly., LONDON AND AYLESBURY, 1903, it i PRINTED ny oegll HAZELL, WATSON AND aa LD, LONDON AND ‘ . xa" a ile Ihe Il. CO IyTRODUCTION , ’ GENERAL SUBJECT SYSTEMATIC SECTION CATALOGUE AND INDEX. NTENTS. PAGE vil XV Slo INTRODUGEION. oy researches in Sphingidac, which have been carried on conjointly for several years, and of which we now publish the results, were undertake with the purpose of giving a sound basis to the classification of these insects by an extensive study of their morphology. Classification is an interpretation of facts. The facts are to a great extent details of the anatomy and morphology of the beings classified. As it is, therefore, largely circumstantial evidence which guides the classifier, the first step towards a correct classification is to fiud out as many facts as possible. In interpreting these facts or characters presented by the individuals—the indi- vidual is the basis of all research—one starts with the assumption that what has been found to be true in the necessarily limited number of specimens investigated, holds good also in the vast multitude of individuals not compared. The possibility of an error in this respect can be lessened by the comparison of a large material of individuals. How large it should be, nobody can predict. To ascertain the extent of variation of the chief classificatory unit, the species, the material is never too extensive. On the knowledge of the extent of variation of the species of a family depends the stability of the superstructure of genera, tribes, and subfamilies. Au ideal classification could be drawn up, if all the species were known which are in existence and which have been. As this premiss cannot be fulfilled, we have to be content with the species that are known. And _ here, again, the foundation of the superstructure will be the safer the more species have been examined. We have endeavoured to comply with these three primary demands ou a classifier as far as it was possible for us: many facts, many individuals, many species. We have not restricted ourselves to a comparison of the pattern, general appearance of the body and wings, and of the ueuration, but have taken into the scope of our research every part of the skeleton of the imagines, and hence have given a broader basis to our conclusions than has ever before been done in this family. Since the structure of the Hawk Moths has never been peaneied to any extent—the Sphingidae being in this respect one of the most ( vii ) neglected families of large insects—we could scarcely ayoid discovering orgatis hitherto overlooked, aud throwing a new light on others which, though kuown to exist, had not been studied comparatively. The organ of friction on the clasper and eighth tergite, found in many males ; the structure of the inner surface of the palpi; the development of the pilifer and the antennal end-segment ; the special structure of the merum of the meso- and metacoxae; the diverse development of the abdominal spination ; the reduction of the pulvillus and paronychium of the claw-segments ; certain differences in the end-segment of the antennae ; the peculiar mid- and hindtarsal comb, etc, have never been taken into account in the classification of the Sphingidac. However, we were very much hampered in one respect. We should have liked to extend our researches in the same degree to the early stages. The Sphingidae Veing tor the greater part exotic species, the larvae and pupae are known only of a comparatively very small proportion. We were faced nearly everywhere by a lack of material. Though we have tried, during the years devoted to study of these insects, to obtain larvae and pupae from all countries where we have correspondents, we had very scant success, and scarcely any success in procuring the first larval stages. The larvae and pupae of many of the commonest Hawk Moths are still unknown quantities.” Nevertheless, our studies of the early stages have not been quite fruitless, inasmuch as they showed us that it is as dangerous to generalise from a few specimens or species in the case of larvae and pupae as it is in the case of imagines. The studies proved to us on the one hand that the deductions commonly drawn from the characters of European Sphingidae are faulty in many respects, and on the other hand that one meets in the larvae and pupae with similarities in not nearly related species and conspicuous dissimilarities in close relatives, just as one finds such puzzles in the adults—puzzles which are stumbling- blocks for the classifier, and a source of pleasure for an inquisitive mind. The figures of the larvae and pupae are, with few exceptions, of little use, if drawn by amateurs who do not recognise and emphasise the essential points. The illustrations of pupae especially leave much to be desired. We draw attention to the following particulars, which we venture to hope will not be so often neglected in future by the artists who are trying to give an * As the ordinary, non-resident, collector in the Tropics shuns inflating larvae on account of the time it occupies and the trouble it gives him, we think it advisable to call attention here to the following very simple method of preserving smal] larvae. Put the larva, after it is suffocated by benzine or chloroform, into a glass tube heated over a flame, The specimen will contract, then expand and burst, and dry in this expanded state. Let the tube cool and take the larva out. In absence of a glass tube, one may use a piece of tin or anything that will stand heating. (ix ) adequate picture of a chrysalis: position of the glazed portions of the eye ; length of the tongue; anterior femur (externally visible or not); structure of stigmatical areas of the abdomen; shape and armature of the cremaster, The number of individuals and species of Sphingidae contained in the Tring Museum is considerable, there being in the collection nearly 16,000. speci- mens, belonging to 660-odd species. Though this material is vastly larger than that contained in any other collection, it was nevertheless uot sufficient to form the basis of a thorough revision of the family. Since many of the species of Sphingidae are very difficult to distinguish, and therefore the descriptions aud often also the figures not exact enough, it was necessary for us to examine the specimens on which the names proposed by previous authors were founded, in order to find out how many species are known and which are the proper names for them. It has been our good fortune to have our appeal for help addressed to private collectors and museums responded to with the greatest liberality.” Without this kind assistance it would have been altogether impossible to disentangle the synonymy and to bring the species which were insufliciently described into their proper place in our classification. We gratefully ackuow- ledze here the help received from the authorities of the British Museum, of the Museums at Oxford (Hope Department), Dublin, Paris, Bruxelles, Berlin, Dresden, Miiuchen, Stettin, Copenhagen, Stockholm, Vienna, Madrid, New York, and Brisbane; and we are no less grateful for the kind assistance rendered by H. Druce, L. W. Distant, W. Schaus, Colonel Swinhoe, M. C. Piepers, P. C. T. Snellen, G. Weymer, G. Semper, Dr. A. Pagenstecher, A. Bang-Haas (coll. Staudinger), P. Dognin, P. Mabille, Dr. W. J. Holland, and Charles Oberthiir ; who all either sent us photographs, types, and other specimens, or allowed us to ‘visit their collections and to study the material contained therein.— Maximas collegis gratias ! An important point for the satisfactory progress of our work was a comparison of the long series of types of Walker’s and Boisduyal’s descriptions, contained respectively in the magnificent collection of Mons. Charles Oberthiir and in the British Museum. A closer study of these specimens than had hitherto been attempted was absolutely necessary. For Boisduval, when visiting the British Museum in the forties of the last century, had named in manuscript and made notes upon the Sphingidae of that collection, which names were for the greater part adopted by Walker in 1856, but often applied to other species than those for which Boisduval had intended them to stand. This muddle became intensified by Boisduval, who, in his monograph published * Only two letters of inquiry have been left unanswered. ‘The names of the addressees may be passed over in silence, (=) in 1875, gave descriptions taken from his own specimens and applied the before- mentioned manuscript uames to species which he believed to be the insects he had so named in the British Museum, but which were often not the same. Moreover, Boisduval failed to recognise many of the Walkerian species, and deseribed them again under new uames. The confusion thus occasioned has, we hope, been successfully cleared up in the present Revision. There are 770 species contained in this Revision. Of these we have not seen the following :— Hyloicus francki, ). 135; known to us from the description.—Kausas. Lapara pineum, p. 1O1; known to us from the description and figures.— N. York. Lapara halicarniac, p. 153; known to us from the description aud figure.— Florida. Polyptychus goodi, p. 245; kuown to us from the description aud figure. — W. Africa. Smerinthulus (?) decoratus, p. 3023 known to us from the description and a sketch.—Sikhim. Sataspes ribbed, p. 474; known to us from the description and figure —Celebes. Euproserpinus euterpe, p. 615; known to us from the description. California, Arctonotus terlooi, p. 606; known to us from the description and fignre.— W. Mexico. Hippotion butleri, p. 760; known to us from the description and figure.— Madagascar. The names of which we have seen the types are marked with an asterisk (*) in the bibliography. Since Linné and Fabricius the Spdingidae of the globe have been five times classified, revised, or catalogued. Hitbner, in his Ver. beh. Schmettlinge (about 1822), was the first to propose a detailed classification of Lepidoptera. The work was, in spite of all its glaring mistakes, far in advance of the time, and was therefore almost entirely neglected by the contemporary entomologists, and sub- sequently forgotten or treated as not being worth consideration. We do not see any reason for rejecting the generic names published by Hiibner in the Verzeichniss. The definitions are insufficient and often incorrect, and the species considered, generically the same belong often to widely different groups, while close allies stand widely separated. That is quite true; but the badness of the classification and of the definitions is—perhaps unfortunately—no valid argument against the adoption of the names. If it were, we should likewise have to reject a multitude of names proposed by more recent authors, whose definitions Ga do not apply to the species generically defined, containing erroneous and quite misleading statements, or whose genera contain very heterogeneous elements, as do, for instance, many of Walker’s genera. Though, in the original definition of Protoparce (type: rustica), Burmeister stated that the pupa had no projecting tongue-case (which it has) ; though Staudinger erroneously said of his new genus Dolbina that it had only ove spur to the hindtibia, aud MHuwe made a similar mnistake in the definition of Smerinthulus ; and though the definition of Moore’s Hathia is so vague as to apply to a host of other SpAingidac as well (as many of Moore’s definitions do),—these names can aud will not be rejected on that account. There is no line to draw between good and bad definitions, sutticieut aud insuflicieut descriptions ; and every description is incomplete. The first after Hitbner to treat again upon all the SpAingédac was Walker, who, in the List of Lepidoptera Leterocera of the British Museum vol. viii. (1856), gave descriptions of all the known genera and species and numerous new ones. His bibliography is generally good, but his descriptions are often so bad that it is impossible to recognise the species without seeing his specimens. He has been much attacked on the Continent, and his names have been ignored to a certain extent by a few authors. Walker did not attempt w Classification of the Sphingidac. Ue simply described the genera in the order he thought proper, without bringing them into groups. His genera are very often as unnatural as many of Hiibner’s. Boisduval followed in February of 1875 with a monograph of the family, containing more exhaustive descriptions of the species and genera. The Sphingidae are divided in this work into a number of subfamilies, of which definitions are given, rather a rare occurrence in lepidopterological works of that time. In the nomenclature of the subfamilies he followed the old Freuch custom of employing scientific terms in a gallinised form—a bad custom, which was formerly in vogue also in the nomenclature of species and genera. The monograph, though far above that of Walker, had two great drawbacks: it did uot contain all the species described up to 1874, and names already employed by Walker were used again for other species (see above). We mention inci- dentally that Boisduval’s names have priority over those of Butler which were published in the Proc. Zool. Soc. Lond. of 1875. The Revision of Sphingidae by Butler—which came out early in 1877, not in 1876, as is quoted by some authors—is scarcely more than a synouymic list with occasional remarks. The genera are grouped into four subfamilies, but not defined, except the new ones. Though the definitions of the subfamilies are based almost entirely on the quite imperfectly known early stages, the grouping Is, nevertheless, au advance on Boisduval’s classification. ‘The work would have (x1) been much improved if Boisduval’s species and genera had been incorporated into the body of the Revision instead of being given as an appendix. The youngest work ou the Sphingidac of the globe is contained in Kirby’s Catalogue of Lepidoptera Heterocera (1892). As a list of names this catalogue has been of great help to us. The classification adopted in it has been much blamed by some authors as being arbitrary. But we think that one should not expect too much from a catalogue. Even the best is full of errors, as a cataloguer of insects cannot possibly have intrinsically worked out all the groups catalogued. Besides these five general works, there are numerous treatises dealing with the Sphingidae of certain restricted districts. Apart from a host of popular handbooks, there are two works on the Palaearctic Hawk Moths worthy of special notice. These are by Bartel, in Riihl, Grossschm. vol. ii., and by Tutt, Brit. Lep. vol. iii. Bartel gives lengthy and generally accurate descriptions, but relies too much on others, whose errors he repeats without having examined the insects himself and formed his own opinion. ‘Tutt’s work is of quite a different kind. It is the most intrinsic ever written on Palaearctic Lepidoptera. The third volume comprises ouly a portion of the Sphingidae ; the remainder of the family will appear in the fourth. The work will be of the greatest help to the scientist who knows the matter well enough to be able to distinguish between what is sciecutific and what appears merely in a scientific garb. The usefulness of the work could have been improved, we think, by a condensing of the contributions of the collaborators, and mistakes could have been avoided by the omission of references to foreign species with which the respective collaborators were not sufficiently acquainted. However, as it is, there is nothing written anywhere on European Lepidoptera coming up to it in thoroughness. The Indian Sphingidae are dealt with by Hampson in Blanford, Mauna Brit. India, Moths vol. i. (1892). The volume should be consulted with some caution, since many distinct species are treated in it as being identical. The species occurring in the Philippines are contained in Semper, Sc/mett. Philipp. vol. xii. (1896), where many figures of larvae and pupae are given. Miskin gave a catalogue of the Australian Hawk Moths in the Proc. Roy. Soe. Queensland vol. viii. (1891). A monograph of the North American Sphingidae by J. Smith is contained in the Trans. Amer. Ent. Soc. vol. xv. (1888). It is the best work on Nearctic Hawk Moths, though the classification is faulty in many respects, owing to Smith’s limited acquaintance with the forms not found in North America. The Cuban species are described and catalogued by Grote and Robinson in the Jour. Ent. Soc. Philadelphia vol. y. and vi. (1865, 1867), and again by ( xiii ) Gundlach in his Contr. Ent. Cubana (1881). The former paper was the best written on Sphingidae up to that time. The Central American Hawk Moths are enumerated and partly described and figured by H. Druce in the Biol. Centr. Americana, Lep. Het, A883—1896). The species occurring in the Argentine Republic are dealt with by Burmeister in his Deser. Rép. Argentine vol. v. (1878) and Atlas (1879). The deseriptions and figures of the earlier stages are of importance, Dnt several mistakes in identification ocenr. There is no list of the African species. Besides these more important works, which are purely systematic, there are treatises of another natnre, dealing with the markings and colour of larvae, like Weismann’s Studies in the Theory of Descent, anid Piepers’s paper on the larvae of Sphingidae published in the Tijdschr. Int. for 1897. All the systematie works referred to suffered from a lack of knowledge of the morphology of the Sphingidae. Nenvration, the organ generally relied upon in the classification of Lepidoptera, being of little assistance in the Hawk Moths, the authors seized upon any superficial characters, and thns were led astray. We hope to fill up that gap in onr science by the present Revision—at least to a certain extent. Nobody can be more aware of the incompleteness of our researches than ourselves. The subject is far too large to allow of being treated after a comparatively small number of years of study in any degree approaching completeness. Nevertheless, we may fairly claim to have given a new foundation to the study of the SpAingidae. The groundwork for future researches is there ; future authors will more clearly see their way, and be able to concentrate their efforts on the elucidation of the numerous points only cursorily touched upon in this Revision. We haye generally abstained from giving detailed description of any stage of the known species; but the keys to the genera and species, and the indication of some trenchant feature of shape, colour, pattern, or strneture under each species, will, we trust, be an efficient guide also to those who wish to nse the work as a means of determining the names of the material in their collections. Thongh we hope not to have missed any names, we have not attempted to give a complete bibliography of all the species. The work is divided into three parts :— I. General Subject. II. Systematic Section. Ill. Cataloene and Index. I. GENERAL SUBJECT. HE researches embodied in a work like the present are of two very different kinds. We had, firstly, to study the insects dealt with; and, secondly, to study the names bestowed upon them by previous authors. It may sound almost ridiculous, but it is nevertheless true, that in many cases the time one has to spend over the nomenclature of a form, in order to clear np nomenclatorial muddle and to find out what form authors have meant to designate with a certain name, equals or surpasses the time one can, for certain reasons, devote to the study of the natural history of the animal. Surely this is wrong. The Natural History of the animal being the subject of our science, the accessory subject of nomen- clature should never have assumed such magnitude. It is waste of energy. However, we have patiently to bear the fruits of the sins of onr forefathers in science, and those who come after us will again mutter bad langnage. One may kick, bnt one has to suffer. We have no sympathy with those of onr contem- poraries who contribute unnecessarily to the burden, which is in itself superfluous, and detracts from the efforts devoted to our science. The system of naming the groups of individual specimens of animated nature has been invented as a help to the student of science, but it has been carried out from the beginning in such a way that it necessarily developed into a bother as well. Why? Beeause the PRINCIPLES OF NOMENCLATURE were not strict enough. Science is a republic where everybody may do as he likes. There are no laws which can be enforeed; and nobody can be prevented from publishing what he pleases. This freedom is a great boon to science. Unfortunately, the results of scientifie research and those of nomenclature are of quite a different standing. If the purported results of scientific investigations prove to he erroneous, they are repudiated and forgotten. If somebody propounds “laws ” of development which are found to be erroneous, science passes on without troubling any longer abont them. If somebody considers the battledore scales of Lycaenidae to be fungi, or the maxillary palpi of Pulicidae to be antennae, such statements are disproved, and are no further encumbrance to science. Statements of fact, and conclusions, once proved to be erroneous, no longer ocenpy the time of the scientific student; science is rid of them. Science can never get rid of a name of an animal or plant once published—unless quite a different system of designation be adopted than that employed since the time of Linné. We cannot simply ignore a name which is a record of an animal or plant. For we must keep a record at least of all the forms which have become known to science, since we cannot have a record of all the forms that exist and haye existed. Even names which are synonyms cannot be dropped ; they must ( xvi ) he carried on for two reasons. Firstly, if they were dropped and forgotten, they wonld in many cases be employed again for something else, and thus land us inevitably in a muddle. Secondly, closer research often proves that what was considered the same at one time is really different, A form may for a long time be lost sight of, but scientists will sooner or later become aware of the oversight, if the name is kept on record. Tor instance, in our case, the Hawk Moths described by Linné and Fabricius respectively as Sphinx thyelia and boerhaviae have heen treated as the same for about 120 years. When we looked up the original records, we found them to refer to two widely different insects belonging to different genera. However, if it is granted that it is necessary, for the sake of completeness of onr knowledge, to keep on record all the names siven to forms of animals and plants, it will also be conceded that if is an unjustifiable act—heeanse it adds unnecessarily to the burden—to suppress a name and replace it by another. Some of the older writers did not seem to think much of recording an already named species under a new name and treating the older name as a synonym. Tabricins—a great and influential man in his time—set a very bad example to his followers not only by his insufficient descriptions, but especially by his arbitrary changing of names. For no reasons whatever he superseded names given by Drury, Cramer, and others, by names of his own invention, and employed—worst of all—the rejected names for other species, thus entangling the nomenclature to such an extent that it is difficult to find one’s way throngh the impasse. With such an example before them, one cannot wonder that others followed suit. Boisduval especially seems to have found great pleasure in his names being printed. One cannot help smiling when one finds him coolly replace Papilio euchenor by a new name, “ «arion Boisd.,” and sees the mannseript-names which he had bestowed at one time or the other upon Sphingidae appear in his monograph of the family under species which had meanwhile been baptised by others. It may be comforting to an anthor who comes foo late to be nevertheless able to launch his names on the scientific world, but it shonld not be done. When Science was in its infancy, a little playing like this may have been pardonable, but nowadays there is no excuse whatever for playing at nomenclature. “The species described by Jones as conformis stands in my collection under the name of ase//vs mihi,” or something to that effect, is not only a foolish thing to publish, but is an intolerable crime, which should always be met by an energetic rebuff. Vanity has something to do with this kind of proceeding, though there is really nothing to be proud of in giving a name to a specimen and avoiding criticism by shunning publicity. int we do not quite understand what is the object of those who are busy publishing mannscript-names which are given by others and which they find on specimens in collections. As it is of no advantage whatever to science, whether it becomes known or not that a bird or butterfly which has a valid name stands in this or that mnsenm under this or that mannscript-name, there must he some other reason for wilfully increasing the list of synonyms. Is it to prove ( xvii ) that the respective authors of the manuscript-names were too lazy to write ont a description and make it public, or that they were not sure if the forms named were really new? Is it to show that the respective authors who gave names to individual specimens of one and the same species really did not know enough of the things they baptised? Is it to demonstrate the carelessness of the respective authors who bestowed, in the collection, a name on an animal for which a name had already been published? Surely if the authors of the names had intended to publish them, we may leniently assume that they would have found time to reconsider the matter before rushing into print. We should not pry into the private foibles of others, and thus detract from their fame. Only published matter is common property, which scientists are bound to critically examine. We have seen many collections with numerous manuscript-names, but, we are glad to state that the bad habit of naming specimens in collections without troubling about publishing a description is very much on the decrease— at least among scientific systematists. The habit has come down to us from a time when few people worked at the same group. There is another class of no less objectionable names which gives little credit to those who are responsible for their introduction. It is a matter of self-evidence that, if somebody claims credit for a discovery, he has to state what his discovery is. Let us assume that A publishes a note maintaining that he has found a new component of air, which he calls so-and-so, but abstains from explaining what it is he has discovered. Another, B, working in the same line, also finds a component of air, which he describes and designates with a name. Then A (or one of his followers) gets up and claims priority for his name.—Another case. The morphologist C announces that he has found in a certain group of animals a new secondary sexual organ, to which he gives a name. There the matter drops ; nobody can possibly tell what the new organ is. Some time after, several secondary sexual organs are discovered in that group, and described and named. Now the knowledge of the structures has become common property to scientists, somebody examines the preparations of (, and, finding that the naked name published by C applies to one of these organs, maintains that the name given by © should be employed for it instead of the later name,. which was accompanied by a proper description. There can be no doubt what the verdict of scientists would be in either case. Science is knowledge of nature. Anything new which does not increase our knowledge of nature is outside the pale of scientific work, and what we do not know is not yet part of science. Facts professed to be new, and new interpretations of facts, do not advance our knowledge if they are kept secret. We know @ priori that there are many facts to be discovered and new interpre- tations of facts to be offered. A naked name or technical term, however, does not tell us what is the nature of the conception for which the name is meant ; and as long as we are left without this knowledge, the name or technical term has no standing in science. Name and technical term are nothing but arbitrary means which science employs as a convenient abbreviation for expositions of b ( xviii ) facts and for the result of lengthy inductions. The name as such is not part of science ; we might employ a number, or a letter, or some other sign instead without interfering in the least with that part of knowledge which is thus designated. It is obvious that these deductions * which apply to science in general apply also to the nomenclature of classificatory work, if the work is meant to be scientific in all its branches. Facts and ideas in classification require explanation like any other facts and ideas in science. Families, subfamilies, and all the other classificatory units down to the individual varieties require exposition by definition. The definitions bring into order the chaotic mass of individuals which forms the subject of classificatory research. Howeyer, instead of operating with the defi- nitions, the systematist employs, for the sake of brevity, names for them, thus simplifying reference. Every name is a term for a definition, It follows from this that a name which 7s vot a term for a definition—7.e. for which no definition has been given—has no standiug. Naked names, with which classification has been favoured in abundance, are no valid terms; they become so only from the time when the fact or idea is published for which they are meant to be employed as a convenient means of reference, and therefore cannot take precedence over a name which has been defined before that time. An anthor who publishes a name for a genus, variety, family, etc., either has some kind of defiuition in his head—and then he should not keep this definition a secret,—or he has not—and then he should not propose a name for something he does not know, and of which therefore he cannot be certain that it exists at all. The action of an author who publishes naked names is as indefensible as would be that of a describer who published names for the respective subspecies, for instance, of those Oriental Papilios which are as yet not known from certain islands, but which doubtless occur there, and which are certainly different from the subspecies of all other places. We appeal to secretaries of scientific societies and to the editors of scientific journals to suppress all new names which are not accompanied by some kind of definition. Systematic work should no longer adhere to the bad habits of the middle of the last century, when the Linnean method of classification, though so young in years, had already become weak as if from old age and had lost its vigour, and classifying was to a great extent more a pastime than a science. A catalogue of names like Dejean’s, containing thousands of nomina nuda, published there for the first time, is, we hope, an impossibility in our time; but single nomina nuda still at the present day appear even in works professing to be scientific. It follows further, that, if we do not wish to jeopardise altogether the efficiency of nomenclature as a convenient means of reference and communication, and thus efface the motive which induces scientists to burden themselves with @ nomenclature, it is absolutely necessary that a definition should be replaced * We understand under deduction the process of reasoning by which we conclude from a general law the correctness of single cases; under induction the process of reasoning by which from single cases a general law is formulated. ; (Cx) only by one and the same name, and that a certain name should apply only to one and the same animal everywhere. Whoever adheres to this principle of stability of nomenclature must concede that this end can only be attained by adhering to the first defined name for every animal or plant. No compromise is possible. Personal preferences for euphony, so-called purity of language, etc., must be sacrificed by all those who sincerely advocate stability of names ; there is no help for it. A publication is meant to distribute knowledge acquired by the author. The publication of a new scientific fact or idea is meant to enable the reader to understand what is discovered. As even nowadays names without any attempt at exposition are considered valid by a good many systematists, it cannot be wondered at that the definitions published are not always so precise as to advance the knowledge of the reader beyond the fact that something hitherto unknown to the author is defined. We have the description, and do not know what to make of it; we have a name, and know perhaps what it signifies philologically, but not what its meaning is in natural excuse for unnecessarily vague definitions science. There is no of varieties, species, genera, etc. ; but there is much in the method of so-called systematic work which explains the frequency of inadequate descriptions. Incompleteness is an inherent character of classificatory research ; the best definition is not complete, and may, therefore, any day become insufficient for recognising the species and variety defined, or must be modified in the case of genera and the higher classificatory cate- gories. A character apparently not worth mentioning to-day may become very important when more allied forms are known. But then, what is the use of having descriptions at all? They are nothing but a record sufficient for the time (or meant to be so). They do not profess to be final, though the author may aim at finality. Here, as everywhere, the advance towards completeness is gradual. As our knowledge increases, the definitions of species, genera, etc., become widened or restricted. The definitions change in scope, while the name which was valid for the original definition remains the same. This contrast between a stable nomenclature and a labile knowledge is a nomenclatorial evil and a souree of much disagreement among systematists. If we tried to make the names as labile as is our knowledge of nature, the remedy would be worse than the evil. Albeit we cannot do away with the evil altogether, we are at least able to mitigate its severity by the application of a dose of common-sense. We all know that the number of specimens on which the definitions of new species and varieties are based—it is of no consequence for our argument which classificatory category the reader designates with the word “species ”—is extremely small compared with the number of individuals existing. Innumer- able species have been described from single specimens. Though this specimen or these few individuals may have been abnormal, though the definition is after- yards found not to cover the entire species at all, as an original definition seldom does—unless it is so vague as to cover other species as well—the name given to the first-named individual or individuals is accepted for the vast ( xx ) number of specimens which are later found to belong to the same species, be they practically identical or be they very different in appearance. Preference is given to the first name, though the species may later have been much better described under another name. Nobody with a sense of responsibility will now- adays re-name a species, variety, or genus of which he knows that it has a name, on the ground that the original definition does not apply to all the specimens of the species, or all the species of the genus, for which the original name is now employed. Everybody who agrees that for the sake of a stabile nomenclature the first name should strictly be preserved, gives to the first individual or individuals which became known to science an importance in respect to nomenclature which none of the later-discovered specimens can acquire. Now, if a definition is not sufficiently precise to recognise by it the species or variety, there is one way of solving the riddle, accepted by all systematists, we think, If there should be somebody who objects to this means of finding out the meaning of published but insufficiently defined names, and advocates that such names should be dropped, he will doubtless retract the objection, if he comes to think of the consequences. To drop such names, though theoretically justi- fied, is impossible, as such a procedure would give the careless worker and the ambitious amateur of the worst sort an excuse for inventing new names wholesale. The means referred to of ascertaining the meaning of an original definition is the comparison of the original specimens. If they are not pre- served (or if the author has based the name on an inaccurate figure or on an insufficiently precise description of an earlier writer), the name cannot take priority over another name; it may be put down as a query synonym under some species with which the definition agrees best, or may be enumerated as species indeterminata at the end of the catalogue of the group. There are very few defined names of Sphingidae which we cannot refer with certainty to any species known to us: Sphinw ixion and Sphinx belis of Linné; Sphinx leuco- phaeata and Chaerocampa thalassina of Clemens; Smerinthus decolor, Sphinz trojanus, Chaerocampa brasiliensis, Macroglossa tristis, and Oenosanda chinensis of Schaufuss, are examples. If the originals are there and are sufficiently well preserved, we may be spared all difficulties, or we may get more deeply entangled in the meshes of nomenclatorial controversy according as we find one or more originals. Let us consider the two cases separately. (1) If the species (or variety) was based on one individual, or, at all events, if only one individual (authentic, of course) is preserved, and there is nothing in the deseription which distinctly points to the definition being based on seyeral different specimens, we are quite certain of what the name applies to. And that is all we require. (2) If the species was based on several specimens, we may find that they belong to one species (or variety), or to more than one :— (a) If they are actually of one species (or variety), there is again no uncertainty about the application of the name. But we must remember that to pronounce two or more individuals to be specifically the same is nothing else ( xxi ) but a conclusion, and that a conclusion may be erroneous. Those who have some experience in systematic work will know that every now and again it happens that the specimens which one author considered to be the same species (or variety) are proved by another to represent several. The reader will find a number of instances illustrating this experience, if he looks over the synonymy in the present work. Therefore, what appeared to be certain may become uncertain again, if there are more than one original specimen. Some authors will, indeed, accept the identification even if it is based on some such mistake, because they consider a name far too unimportant to justify a strict adherence to principles, if a change of names is involved. However, the majority of classifiers will oppose a name which is incorrectly applied. This spirit of opposition against all mistakes is very healthy. We should deplore its absence ; for we are sure, because we know instances, that he who intentionally overlooks errors in apparently irrelevant matters, will treat in the same spirit also details of fact which appear to him trivial, which may, however, be of the greatest bearing upon general questions, and, therefore, mislead altogether the generaliser who has to depend on the accuracy of the specialist. (6) If it is proved that the original specimens belong to more than one species (as do, for instance, the originals of Walker’s Macroglosswm_ sitiene, corythus, Nephele viridescens, etc.), systematists have adopted several methods of narrowing down the conglomerate to one species. These methods are as follows :-— (4) First method of restriction: The name of a composite species is to be restricted to that component which is the first to which the name is afterwards applied by the same or some other author. Illustration: Macroglossum corythus A, of 1856 consists of three species { B, Of these B is the first mentioned as C. corythus after 1856; ergo, the name of corythus is restricted to B.—To be certain of the result, it is necessary to know which is the first, and that requires a knowledge of all the books where the name occurs, and, moreover, a knowledge of what is meant therein by the name. These premisses may sometimes easily be got over, but they present more often difficulties which are as intricate as those which the method professes to solve. (6°) Second method of restriction: The name is to be restricted to that component of the composite species which remains after the other components have been subsequently separated under new names. Illustration: Macroglosswm ’ A, corythus +B, As A and B are described as new in 1875, the name corythus C. remains for C.—To arrive at this result, one has to inquire into the descriptions of the new species, in order to find out whether the new names really apply to A and B; the new species will in many cases again be found to be composite. The method, therefore, creates new difficulties in trying to remove the old ones. ( xxii ) (c?) Third method of restriction: As the first and second methods are opposed to one another, differing nearly always in the results attained, we reject them both. The energy spent on the book-research which either method requires is misapplied, reminding one too much of the famous fight against windmills. Nomenclature is not part of nature; it is an auxiliary means invented by the classifier for his own convenience. What in the name of common-sense compels us, then, to turn a convenience into an inconvenience? There is a wide scope for research in nature requiring all the energies of scientists. Why, then, impose upon scientists those unnecessary labours which have only a nomenclatorial, but no scientific result? The method adopted by ourselves is at once logical and very simple, and removes all the difficulties as far as that is possible. Our method of dealing with composite species (and genera) is to narrow all cases down to the case dealt with under (1) by simply applying also here the law of priority recognised by nearly every classifier as the only means of arriving at a stable nomenclature. From the sequence of the localities under a composite species, or from the characters mentioned in the definition, or from the bibliography referred to by the author of a new species, one is able to draw up a sequence of the components of the species. If Macroglossum corythus, as conceived by Walker in 1856, consists of three species, A, B, C, we have :— A= WM. corythus ; Macroglossum corythus\ B = M. corythus ; le = M. corythus. Each of the three components is M/. corythus, according to Walker. According to the law of priority, the same specific name cannot stand twice in the same genus, and the name occurring more than once can be valid only for the species which was first published under that name, or which stands first in the book where the name is defined for the first time. This rule being applied to the above case, it follows that the name corythus can stand for A only. All we have to do, therefore, is to find out the sequence of the components of a composite species. This is mostly easy, especially in the case of geographically separate forms, In the case of composite genera the sequence is given by the names of species mentioned, there being very few genera defined withont reference to one or more species. Strict adherence to the above rule makes the first species mentioned the type of the genus. {One might object that this mechanical application of a rule leaves it entirely to accident which species becomes the type of the genus, or to which particular portion of a composite species the specific name is restricted; and, further, that the author did not intend to give the first species or the first specimens respectively any such pre-eminence, and that the “type” thus fixed may be just the one to which the description applies least. We reply, firstly, that we do not know the intentions of the author, as he did not state them ; and, secondly, that, if the description applies accidentally less well to the species ( xxiii ) or specimens first in the sequence than to the others, this argument holds equally good in the case of the types fixed by any other method of restriction. We invite the reader to find out the types of such genera as Papilio, Sesia, and Zygaena by all three methods. A glance at the original definitions of these genera suffices to fix priamus, tantalus, and filipendulae as the respective types according to the third method. A study of several families is necessary before the types can conscientiously * be ascertained by the first and second methods, since the species originally included in each genus belong to different families. In mammals and birds and several other groups of animals the second method has almost generally been adopted, for genera at least. If the systematists have there really arrived at a stable nomenclature, no change 1s necessary, a stable nomenclature being the main aim of the principles of nomenclature. In Lepidoptera, however, and other insects, the first and second methods, less often the third, have been followed, and that has landed us in such a muddle that there is no question of stability having been attained. This being so, we should have adopted, as a matter of course, the surest method of restriction for the sake of avoiding waste of energy, even if our method was not the logically correct one for all who agree that strict priority has to be adhered to. If the authors of the names for varieties, species, genera, etc., had done from the beginning what we now have to do with their names; if the authors had restricted every name in the way that we now are compelled to restrict it, much time would have been saved. We all agree that a specific (or varietal) name based on one specimen, and a generic name founded on one species, are as valid as names based on more material. Further, if all names were based on one individual or on one species respectively, there would be no composite species and genera; and if the original individuals of each species and variety were preserved, scarcely any difference of opinion would arise among careful workers about the application of the names. We cannot alter what has been published; but our contemporaries and the scientists who come after us have it in their own hands to simplify nomenclature in the way here indicated by making all names monotypical. We do not know who was the first to fix a type (=typus) for the name of a species, variety, or genus. The nomenclatorial term appears already in 1816 (Dalman). The word tyye was perhaps not the best that could have been chosen, as it had already a definite meaning also in science, signifying that which is typical for a group of units. But as we frequently use in science the same word for different conceptions (claw, wing, tarsus, lip, mandibles, tail, tongue, etc.), the philological objection against the term “type” is not of much weight. However, the difference between the meaning of the word as used in ordinary language and the meaning of the nomenclatorial term has occasioned confusion, and hence led to another kind of objection. * Some writers have simplified matters for themselves by ignoring the exotic species altogether! ! (_ xxiv ) Some authors, accepting the word “type” in the ordinary sense implying that the specimens called types are typical individuals, very properly reply that these types are often aberrant specimens, and very seldom the most typical for the group of individuals to which they belong. This confusion of the verbal and the technical meaning of the word “type” misleads those authors to insist further that, there being no “types” in nature, one individual being no more a pre- eminent representative of the species (or variety) than another, the word “type” as a nomenclatorial term has no standing. It is obvious that those authors fall into a deplorable error of confounding the names, which are the product of scientists, with the objects named, which are the product of nature. Certainly there are no types in the nomenclatorial sense in nature, but there are also no names. ‘he type is as such not at all the type of the species, but is the type of the arbitrary name given to the first specimen or specimens, and applied by common consent to all the specimens which belong to the species, of which the type-specimen is only a member, like any other individual. Those who have the stability of nomenclature at heart, and are unwilling, When proposing a new name, to lay an avoidable burden on scientists—and who do not consider themselves infallible—should mark one individual as type (= typus) of the name, and make a clear statement to that effect when publishing the name— and one individual only. Jyery care should be taken to have such individuals preserved. There is neither justification for opposing this usage by which the systematists benefit enormously, nor for employing the purely nomenclatorial term “type” in any other nomenclatorial sense than the one here advocated. As a name is not valid if the animal or plant has already an earlier valid name, we reject also all those uames of composite species and varieties of which one of the components has an earlier valid name, and the names of composite genera and higher categories which comprise the type of an earlier validly named genus or higher category respectively. Temnora brisaeus of Walker (1856) has no standing, because it is a mixture of several species, of which one is Cramer's pylas (1779). Dalman’s Hemaris (1816) is a synonym of Macro- glossum (1777), because it includes the type of Macroglossum. And for the same reason the subfamily name Macroglossinae (1875) becomes a synonym of Sesiinae (1819). Ambulyx of Walker (1856) cannot stand, as it includes Amplypterus of Hiibner (1822). In general terms :— If A is based on a, or on a and 4, which are not cospecific, respectively not congeneric, ete., or on a, b,¢, ” ” ” ” ” ” 2 and B is based on a, or on é and a, _ is * ” » ” ” or on a and 4, s ia . 5) ” ” ” or on ¢ and a, Pe + 5 ” ” ” or on a and ¢, % » ” ” ” ” ” or on 4, a, ¢, or a,b, c, or a, ¢, B, ete. then B is a synonym of A. ( xxv ) This does away with some names proposed in scamped work, of which science would be well rid, especially with those names which are founded on actually the same material. Before a new name is introduced, the author should ascertain to the best of his ability that the material for which the name is meant to stand has no earlier name. ‘This is a demand on a describer, on the fulfilment of which classifiers should rigorously insist. In an ideal nomenclature a name should convey to the systematist the characters of the animal or its place in the system. As long as the classification is not final this is not possible, and final it will most likely never be. One step towards this goal was made by Linné himself when he established it as a rule that an animal or a plant was to be designated by a generic and a specific name. In Lepidoptera he tried to go even further by indicating, in the case of some groups, by means of a certain ending to the specific name (-ed/a, -ana, etc.), to which family the species belonged. But this proved to be a complete failure. The Linnean binominal system of nomenclature had the one great advantage that, when the number of forms became larger and larger, there was no serious objection to employing the same specific name in several genera. In Linné’s time, when so comparatively few animals and plants were known, the generic name was indeed sufficient to tell the scientist the position of the form in the system. This is no longer the case. By far the larger number of generic names do not convey any idea to scientists, except to a few specialists who happen to know them; the family or even the order to which the genus belongs has generally to be added to make clear what is meant. So far students of natural science have adhered to the Linnean binominal system with that tenacity with which human beings generally cling to what they have become accustomed by long usage. However, a great change has taken place in one direction. When the theory of descent threw an entirely new light on the forms of animated nature, the study of variation became an all-important subject. That the species were more or less variable was long known. But now the variability assumed quite a different aspect. It was found that there were different kinds of variation. Whereas formerly the chief object of classificatory research was to separate the individuals into species, and group these in genera, and so on, now there were in addition the several kinds of varieties to be carefully studied. For this purpose a nomenclature of varieties is as necessary and as convenient a help as the Linnean binominal nomenclature is to the student of species and higher categories. Systematists agree that the name of a species, genus, or family must be of the same form, so that one recognises by the name (or rather the formula) which classificatory category of units is meant. A family name must be different in form from a subfamily name, and a genus name from a non-generic name. The name itself must show us whether it designates a species, a genus, a variety, a family, etc. The efficiency of nomenclature would be nil if one could not see from the name that Charaxes castor designates a species, Papilio a genus, Agaristidae a family, according to common agreement, Sphinwx ocellata x Amorpha populi a hybrid, Araschnia levana f. t. prorsa a seasonal form, etc. This is so ( xxvi ) self-evident that we ask ourselves in wonder how it is possible that there are systematists who do not—should we say will not ?—recognise the necessity of it. If Fringilla coelebs is accepted as a formula for a species, Sphinx atlanticus is also a designation of a species, and not of a genus or a variety or a subspecies. To speak of “species” Sphinx ocellata and of “ subspecies ” Sphinx atlanticus is a coutradiction unworthy of science. In former times varieties were looked upon as freaks of nature. They were to many a classifier an interesting nuisance, which often threatened to upset the balance of his well-fixed species, and were on that account more often entirely put aside than welcomed as an object for research. Esper, who went perhaps deeper into the phenomena of variation than most of his contemporary entomologists, already distinguished between ordinary varieties (Abweichungen) and abnormal individuals (Ausartungen). However, as long as the principle of evolution underlying these varieties was not recognised, there was no need to study them systematically, and to work ont a system of nomenclature which would bring into order the chaos of varieties, as did Linné’s binominal system the chaotic mass of species. From Linné onwards varieties, if provided with a distinctive name, are recorded in various ways. The following names may serve as illustrations : Papilio iris luteus; Columba oenas 8. domestica; Phasianus gallus B. gallus cristatus ; Phasianus colchicus (B.) Phasianus varius. The word varietas, introduced by Linné as subordinate to species, meant anything deviating obyiously from the normal individuals of a species. The practice of putting the term in an abbreviated form, as varict. or var., before the varietal name does not seem to have sprung up before the beginning of the nineteenth century, and the use of the term aberratio (ab. = aberr.) is still younger. Not rarely the “variety”” was in reality the normal form, while the “species” happened to be described from aberrant specimens. There was no strict rule for the employment of rar. or ab.; some treated well-marked varieties as var. and less obviously different individuals as ad., the distinction between var. and ad. being merely quantitative ; while others employed ad. for abnormal specimens ocenrring singly among the normal ones, and var. for the regularly observed varieties. There are many collectors and classifiers, representing the stagnant element in this department of our science, who look at varieties still from either of these standpoints. Since the middle of the last century, when natural science stepped from childhood into manhood, the study of variation has gradually become more methodical, with a change in the PRINCIPLES OF CLASSIFICATION, and has now attained a height of which our forefathers in science did not dream. New lines of research bring to light new series of facts; and new kinds of facts require a new terminology. It will not do to have the same nomenclatorial formula for a species as for a genus ; and so it will also not ( xxvii ) do to name all kinds of varieties in the same way. It was Staudinger who first separated the varieties of Lepidoptera into two categories: geog raphical and non-geographical varieties. There is indeed a great difference between these two categories, as we shall see later on. Unfortunately Staudinger adopted for the geographical variety the old term varietas (var.), and for the second kind the term aderratio, giving both terms a definite meaning which they did not originally have. He should have invented a new term at least for the geographical variety. As it was, the application of var. and ad. remained in as great a muddle as before. While there was thus some sign that the relation of the varieties towards each other was assuming a clearer aspect, at least to some authors, a curious misunderstanding crept in, which prevented many classifiers from perceiving the true relation of the varieties towards the species. It is self-evident that two or more different-looking animals which are found not to be specifically distinet from one another belong to the same species, 7.2. are components of this species. All the components together are the species. This is a truism. How- ever, systematists became accustomed to look upon that particular component which was first described and named, and of which the name was accepted as the name for the entire species, as being the species, while it was, as a matter of course, only one of the components of the species. It is utterly wrong to say that the first-described form is the species and the later-described forms varieties of it. We know, for instance, that Araschnia levana and prorsa are the same species ; neither lerana nor prorsa is the species, but the pale spring-form levana and the darker summer-form prorsa together are the species, thus :— levana | ; _ -= species. prorsa | It is purely conventional, on grounds of nomenclatorial efficiency, and has nothing to do with the relation of the summer- and spring-broods towards each other, or with the causes and the origin of such horodimorphism, that the first-given name, lecana, is applied to the species. In doing this people forgot that the name Jdevana, originally standing for part of the species, was now employed for the whole as well, and that therefore the true relation between the species and its components is this :— spring-brood levana | P summer-brood prorsa | — Bngtles ena. Or, if we use the term f. ¢. (= forma tempestatis) for horodimorphic forms, we have as formula for the insect :— ; f. t. levana ; Araschnia levana | f. t. prorsa. Instead of employing this formula, classifiers spoke and speak of a species levana, meaning the spring-form, and a variety prorsa, meaning the sammer- form, co-ordinating the one with the other :— Araschnia levana ; Araschnia levana var. prorsd. (_ xxviii ) This is wrong, as the spring-form is no more the species than is the summer-form. What we have said here in regard to seasonal varieties, applies also to geographical and individual varieties. Which of the components of a species is the first-deseribed and -named form depends in nearly every case entirely upon accident. The first-named form may be the most aberrant and the very youngest development of the species, having originated from one of the later- described compounds of the species. To call this accidentally first-named portion of a species the species and the later-named forms varieties of the first, is a Indicrous confusion of facts. And yet, systematic work, from mammals downwards, teems with this glaring misconception. As nomenclature is a convenient auxiliary to classification, as it is sub- servient to science, and must therefore be accommodated to the latter, it should not form a hard-and-fast structure, into the compartments of which the results of classifieatory research have to be squeezed somehow. The distinction between tle scientific part and the accessory nomenclatorial side of classification should never be lost sight of. The aim of scientific research is to discover and elucidate the phenomena of nature. Classification, as part of science, aims at an understanding of the connection between the individuals. To attain this object it relies on facts discovered by two lines of research: firstly, on the facts relating to the body ; and, secondly, on the facts relating to life. And here, as in all scientific research, we find the primary question underlying all investigations to be difference or no difference, because science is always comparative, consciously or unconsciously. Morphology and anatomy provide the classifier with the knowledge of the body. In a vast number of instances there is no other knowledge available than this, to build a classification upon. The corporeal facts of the morphologist and anatomist are, however, no absolutely trustworthy basis for a superstructure. For the primary units of the classifier, the individuals, are always different from one another to a certain extent, and therefore cannot be proved to be classi- ficatorially identical by corporeal comparison alone. As in inanimate nature identity can be established by action and reaction, so also in animated nature. The observed differences and apparent identities in the bodies of the individuals have to pass the higher criticism of the knowledge of the phenomena of life. Two individnals may appear very different to the morphologist ; but the classifier, who knows from observation of the living animals that one is the offspring of the other, cannot establish any other connection between them than that of parent and offspring, however conspicuous the bodily differences may be. The differences between young and adult, male and female, parent and offspring, brothers and sisters, however prominent they are, lose all the classificatory importance which the morphologist and anatomist (and the classifier misled by him) thought they had, when biology establishes the true relationship of such individuals. On the other hand, apparently insignificant corporeal differences, which the morphologist may scarcely deem worth noticing, often turn out to be differences between (xxix ) animals which are entirely independent of one another. The conclusions based upon the facts of anatomical and morphological research must always give way to the direct proofs of biology. The significance of corporeal characters is established by biology. Anatomy and morphology give the quantity, biology determines the quality. It is therefore obvious that the classifier should not let himself be guided in his judgment solely by a consideration of the quantity of bodily characters, but always keep in mind the higher criticism of quality. If he does this, he will not easily fall into the error of treating two groups of individuals as being of the same classificatory category (variety, species, genus, etc.), if ‘piological considerations are against it. If the classifier had no other guide than corporeal similarities and differences, the classification would merely be an artificial arrangement, without regard to the true connection between the animals classified. Such a classification, perhaps very useful for the mere collector, might be likened to an arrangement of minerals according to their external features, without any regard to their chemical composition. Although there are no biological data available of most animals, of which we know nothing but what the dead bodies or portions of them (embryo to adult) tell us, the scientific classifier has nevertheless another guide. This guide is the principle of judging from analogy (the word used in the logical sense). We give two illustrations. If in a certain country the spring- and stmmer-broods of a certain group of species have been proved to be different in all the species examined, we may safely conclude that they differ also in those species of the group which have not yet been examined. Therefore, forms of that group of which it is only known that they differ in the same way as the horodimorphie forms of the better-known species must also be treated as seasonal varieties, and not as distinct species. As we have found that the genital armature, though individually variable, does not exhibit any seasonal differences in those species of Lepidoptera of which forms proved to be seasonal have been examined by us, we are bound to conclude that Lepidoptera which do constantly show differences in those structures are not seasonal forms of one species. The premiss is the better established, and hence the conclusion is the more likely to be correct, the more species have been examined. This deductive reasoning, though logically exact, will never give us certainty. For the animals as we know them are the product of evolution; the result of evolution is dependent on the nature of the animal and of the evolving extraneous factors ; these extraneous factors being independent of the animal, their combination with the internal factors—which combination determines the line of develop- ment—depends on accident; results determined by accident cannot be deduced from an a priori law. That means, we do not a priori know that what holds good in all the cases examined is true also in every case not yet examined. Nearly all the so-called “laws” in biology are nothing but more or less well- formulated rules of probability admitting of exceptions. A rule may*apply to every individual case within a certain group of animals; but the further away we go from that group, the greater becomes the chance of exceptions turning (xx }) up. It is hasty to generalise from a limited group of animals, though biologists are nowadays in the habit of doing so. Every group first requires independent be true in another group can be pronounced investigation before a rule found to to be true also in the one under consideration. The “laws” of development have all a premiss; they are true only under certain conditions. The scientist must, therefore, always remain aware that the unexpected may happen, and individual cases turn out to be exceptions. For instance, what the classifier considered to be a generalised character may turn out to be a specialisation, and what is a specific difference in one group may only be seasonal in another. Since the classifier is in the same position as the commentator of a book which is written in a language only imperfectly known to him, and from which many words and whole pages are missing, there is necessarily a certain amount of assumption in classificatory work which makes the results proportionately unstable. However, the degree of instability can be lessened to a great extent. Just as we know «@ priori that a certain character which appears generation after generation in the larger number of individuals of a species will finally be present in all (if the conditions of life remain the same), so the probability of the correctness of a conclusion in classification will become certainty, if all the details of the animals and all the facts of their biology render it probable that the conclusion is correct. In order to approach this certainty the classifier has to give his conclusions the widest possible basis. The results of anatomy and morphology must be checked off by biology, and the conclusions derived from one organ by the conclusions based on every other part of the body. If one organ contradicts the other, there is a mistake either in observation or in interpretation. The more such mistakes are discovered and corrected, the safer the superstructure of conclusions will be. Grouping the animals according to the apparent develop- ment of one organ leads to an artificial arrangement; grouping them in accordance with all organs checked off by biology results in a final classification. However, finality, even if the classification is restricted to a small group of beings, entails such au enormous expenditure of energy that it can be approached only gradually in the course of time by continued co-operation between the various lines of research. All classification begins with the discrimination between individuals. Every- body distinguishes between the specimens he observes around him and brings them into groups, thus setting up a rough kind of classification. Experience shows which individuals stand in the possibly closest blood-relationship—that of parent and offspring, and of brothers and sisters. The classifier starting with this experience, which can be corroborated over and over again, has therefore as foundation for classification a solid fact which will remain firm when a flimsy superstructure that a wild fancy may think fit to erect is blown away. Obser- vation of the individnals so closely connected as parents and offspring teaches us that there is always a certain amount of corporeal difference between them, every individual having an individuality. We notice this variability in all organs ( xxxi ) when studying them closely. There is no constancy. How far the variability extends is a primary subject of investigation. Lepidoptera are the most convenient group for the study of variation, as they can be comparatively easily reared and experimented upon. Variability may be fairly equal in all organs, or may be excessive in one and slight in others; variability of one certain organ may always be accompanied (within one group of animals) by variability in certain other organs, or the variability may be quite independent. Variability (= state of being different) is to be accounted for by two kinds of variation (= process of becoming different): firstly, ¢ndividual variation, pertaining to brothers and sisters ; and, secondly, generatory variation, pertaining to parent and offspring. Individual variation is normally such that there is a gradation from one extreme to the other, the number of individuals becoming, however, smaller towards the ends : monomorphism, respectively—chromatism. Or the individuals which are all the offspring of the same parents fall into two or more groups, which may or may not be connected by intergradations : di- or polymorphism, respectively—chromatism. In every case there appear occasionally specimens which stand outside the ordinary range of variation. Such aberrations are generally aberrant in one organ only, and otherwise quite normal. Aberrations should not be confounded with monstrosities, in which the deviation from the normal is due to injury of the early stages. The dimorphism of commonest occurrence is sexual, the purely sexual characters being nearly everywhere in sexually separate animals accompanied by differences in size, colour, and some kind of structure. Marked dimorphism in both sexes is comparatively rare. But where such di- or polymorphism exists, and where the similar individuals of both sexes are referred to under one name, it must not be implied that the similar specimens are biological entities. They constitute like aberrant indi- viduals a morphological or anatomical unit named in contradistinetion to the other individuals. A black ¢ of Papilio machaon is not more nearly related to a black ? than to its own normally coloured brothers and sisters, and the brown 3d of Mimas tiliae are as much the dd of the green ? ? as of the brown ones, in spite of corporeal similaritivs and dissimilarities, and in spite of similarly coloured specimens bearing the same name in classification. In most groups of Lepidoptera the variability of the males is inferior to that of the other sex, there being numerous species in which the male is monomorphic, while the female is polymorphic. In such cases the females are generally protectively coloured or otherwise adapted in several directions, and may occasionally become mono- morphic again or more constant than the males, in consequence of the best- adapted form becoming prevalent and finally appearing alone. The habit of naming varieties has been in vogue among entomologists and conchologists more than among any other systematists on account of the great variability of insects and shells; and as the student of variation requires names for the sake of brevity of reference, there is nothing to be said against the habit. But since every individual deviates in some organ from the mean, there is some danger of the naming turning out indiscriminate. Fortunately, Lepidopterists have so ( xxxii ) far confined themselves to ardent baptism of individual varieties showing some distinction only in colour, pattern, or shape. There is, of course, no reason for such restriction. There is just as much justification in Lepidoptera as there is in beetles for naming structural varieties. If it is not objectionable to baptise sculpture-varieties of Carabus, there can also be no objection against providing with a name, for instance, the numerous individuals of Chalcostidae which are different in nenration. However, it is our opinion that the naming of individual varieties should not go further than is necessary, and that only the student of variation can decide how far it is necessary. Conspicuous individual variability is observed among Sphingidae both in the larvae and the adults, the caterpillars of Mdacroglossum and Cephonodes and the imagines of some Ambulicinae being very variable in colour. The differences between parent and offspring are no less marked in many instances than those of the offspring ¢mfer se. The generations as they follow one another are either practically the same, or there is a cycle of more or less different generations. In Lepidoptera the cycle is generally restricted to two or three broods. As these coincide in time of appearance with the seasons as a rule, this particular kind of “ generatory” variation is called seasonal. Though as a matter of course the individuals of each separate brood are a biological entity varying individually in the way explained above, there is nowadays a great inclination amongst systematists as well as biologists to confound seasonal with individual variation. If two different specimens are offspring of the same female, they are surely not seasonal varieties. At the bottom of the confusion lies the assumption that, since seasonal variation depends on meteorological differences of the seasons, all differences are cansed to appear by such factors. This assumption is certainly incorrect, and therefore the indiscriminate treatment of different - looking specimens as seasonal quite misleading. Everybody may draw conclusions as he pleases, but the facts must be represented as they are; they must not be tampered with. Seasonal variation is observed in several instances among Sphingidae. The most remarkable cases are found in Huemorrhagia and Hyloicus. Though we know that generatory variation is often structural in insects (Aphidae, Physopoda, etc.), we were nevertheless surprised to find a structural difference in the claw-segments of the two forms of Hylotcus perelegans, which forms we presume will turn out to be seasonal. Having studied the differences between the individuals proved by rearing to be parents and offspring, the student will be able to select from the individuals at large those which a comparison of their bodies convinces him to be so nearly identical with the specimens reared that they might very well be brothers and sisters of them. But knowing that corporeal similarity is no absolute proof of biological identity, he will prudently test his conclusion— perhaps gather some female moths which come to his lamp and which seem to him identical to all intents and purposes, get eggs from them, and rear the 5D* larvae. To his astonishment he finds that the apparently identical females have ( xxxili ) produced two very different kinds of caterpillars, from which he obtains in the course of time a series of imagines again very much alike, but presenting now to his searching eye and suspicious mind some slight differences. Continued experiment with the two insects proves to him that he has to do, not with a dimorphic larva, but with two entirely independent beings, which fly together and feed as larvae on the same plant, and which are scarcely distinguishable as adults, and are nevertheless perfectly kept apart in nature. He realises that there is a gap between these two kinds of insects which is utterly different from the gap between the varieties which stand in the relation of parents and offspring ; that there is an effective barrier which lies in the nature of the specimens themselves, separating the two sets of individuals completely, though there is no extraneous barrier between them. And by studying further the life around himself, the student will perceive that the animated world is composed of a multitude of such sets of specimens, of such entities, all separated by that same kind of barrier. The knowledge of the existence of this barrier is essential for the classifier. What the barrier is the student cannot know with certainty. The corporeal differences observed in the individuals are not the barrier, but are only accessory to it. Passing now into a neighbouring country, the scientist will find practically the same composition of the fauna, though some old friends may be missing and some strangers may meet his eye. A good many of the entities will indeed be identical with what he knew before, but others appear in an altered garb. In one the range of variation has remained the same; but the indi- viduals which were in the minority in the first place are here in a majority, the mean of the variability having changed. Another entity, which was known to him as being monomorphic, is dimorphic in the new locality. A third, which was seasonally variable there, does not exhibit seasonal variability here. Among the specimens of a fourth entity there appear individuals different from what the student had hitherto seen, the range of variation having become shifted or widened. In others, again, he finds the proportion of such different individuals to be larger and larger, until the student comes to entities of which all specimens exhibit some distinction from the individuals of the former country. They are the same entities, but with a difference. Now, in which relation to each other stand such geographically separated entities? The two extreme cases are these :— (1) The geographically separated entities are, each taken as a whole, identical, with the same range of variation and the same mean of variability. (2) The geographically separated entities are different from one another in all individuals. Between these two extreme cases there are all intergradations. The difference may be found only in a very small proportion of the specimens, or only in one sex, or in the greater number of individuals, or in nearly all, or in all. The difference may be structural, or chromatic, or both; it may be c ( xxxiv ) conspicuons or scarcely perceptible ; the gap between two geographically separate entities may be small, or large, and it may be completely bridged over by individuals from other countries. A careful observer of any group of sexually separate animals will know many instances referable to the Various Cases alluded to. We will call such geographically separate animals which are different, geographical representatives. In order to see clearly what the difference means, it is necessary to know the meaning of identity in geographically separate individuals. If two geographically separated specimens prove on comparison to be identical, or are considered identical, the student has to conclude that they are not separated by that barrier which we have above characterised as effectively keeping the entities of the same country (= synoecic entities) apart. As each of the two individuals is a portion of an entity in its native country, these two respective entities are also not separated by that barrier. That means, the two entities together are only one single entity separated from all the others by the above barrier. We cannot help adding, that it appears very strange to us to see some indi- viduals pronounced identical, and to find them, nevertheless, treated by the same author as belonging to two “species.” * It is clear, from what we have just said, that geographically separate animals which are identical in some of the individuals, or, in other words, of which the range of variation overlaps (A! varying from 1 to: 5, A* from 4 to 12, A® from 7 to 19, At from 18 to 21, ete.), are one biological entity ; that is to say, the geographically separate different specimens are the product of geographical variation of the same animal. It is a difficult task to find out the limits of variation. The material which comes into the hands of the student being insignificant compared with the number of individuals at large, it depends upon chance that the extreme individnals become known. Experiments, especially with Lepidoptera, have proved that the range of variation can be increased artificially. That is to say, the capability to vary is greater than we generally observe it to be in the material collected, and therefore we must expect that the range of variation is in many cases actually more extended than we see it, and that, consequently, very often also those geographically separated different animals overlap in characters which differ constantly in the specimens contained in collections. If the number of individuals is small (we know frequently only one or a few Specimens, often only one sex), it is the merest assumption to say that all the specimens of the respective locality are different from all the individuals of a certain other locality. Aud an author who makes such a statement is no less liasty or superficial in his judgment than the one who waves the distinctions aside as being of no importance. However, if there is any reason for expecting the characters to overlap, it follows from what we said before, that the individnals from the two places together belong to one entity. The accom- panying diagram (Fig. 1) demonstrates perhaps more clearly the correctness of * Bee Nov. Zool. TX. p, 459 462 (1902). (exxxy™) our argument than any words. The diagram illustrates an extreme hypothetical case. The curves A! and A? represent the variation* of two geographical representatives. The extent of variation, @ c, is the same in both repre- sentatives, but the means are different. The small number of specimens, a 4, belonging to A? are identical with the majority of the individuals of A, while the small proportion 4 ¢ of the specimens of A! are the same as the majority of A®. It is obvious that there is little chance of the student getting any of the specimens a 6 of A? and J ¢ of A’, if he has not a really large material at disposal. To him A! and A? would appear to be separated by a gap; they would appear to be constantly different, though they are, each taken as a whole, morphologically identical. Two sexually or otherwise di- and polymorphic animals do not. differ in all specimens in the same way; one sex or one set of individuals may be distinguishable by colour, the other by shape; one form may be paler, another darker, than the respective forms of the animal with which they are compared. To find ont the differences between two geographical representatives which are A! Fie. 1. A? di- or polymorphic, it is necessary to compare the corresponding forms with one another, just as one has to compare sex with sex, larva with larva of the same stage, pupa with pupa. There are also monomorphic geographical representatives which differ from one another in a similar way, some individuals differing in shape, others in colour, others again in pattern or in structure. The difference between the two representatives is also often constituted by the ensemble of the characters in each individual. Such cases lead over to those in which the geographical representatives are completely separated by a gap in their morphology, either in one or in several organs. If At, A®, A®, At, etc., are the representatives of one certain type of animal, inhabiting, for instance, each one particular island, and B', B’, B%, B*, ete., those of another type of animal found on the same islands respectively, there is a corporeal gap between A! and A’, etc., and between B! and Bb’, etc., which do not live together, as well as between A! and B', A* and B’, ete., which live together. And now the question arises, Are the gaps between the various A’s * The ordinate gives the number of individuals, the abscissa the degree of difference. ( xxxvi ) and those between the various B’s biologically equal to the gaps between the synoecic A’s and B’s? Does the barrier which we know to exist between the aynoecic Al and B!, which oceur together, exist also between A’ and A’, which are geographically separate? There are many scientists who say Yes, and many who say No. Let us consider first the arguments advanced for the correctness of an affirmative answer. (1) The geographical representatives A’ and A? are constantly different ; ergo, there is a constant barrier between them, as in the case of A? and Bl.— Firstly, this is begging the question. Secondly, we have seen above (p. xxxiii) that corporeal differences as such do not constitute the barrier existing between A' and B' Thirdly, in a vast number of cases it is mere guess-work to maintain that A! and A2 are constantly different ; all we know of them is that they are different under the special conditions under which they are living, similarly as seasonal varieties may be constantly different if the conditions are constant. If the constancy of the special conditions falls, the constancy of the corporeal difference between A! and A? will certainly or perhaps—we cannot tell a priori which is correct—also break down. (2) Al and A® are geographically isolated. They form therefore separate biological entities which do not interbreed and fuse—We reply, firstly, that this is again a restatement of the question; secondly, that, as there are numerous cases of identical individuals (as far as there is identity in individuals) being geographically separated from one another, geographical isolation as such is no criterion whatever ; and, thirdly, that the facts of A’ and A? being geographically separate, and A! and B! living together, constitute certainly not an agreement, but, on the contrary, a difference in the relation between A! and A?®, and A! and B', respectively. Our arguments for geographical representatives not having a prior? the same biological standing as the synoecic animals which are separated by the barrier before characterised (p. xxxiii), are as follows :— (1) The geographically separated and morphologically distinguishable representatives A’, A®, A%*, etc., are morphologically and anatomically more similar to one another than to B!, B?, B, ete. The A’s are modifications of one and the same type, the B’s of another, the C’s of a third, etc., and each type as a whole stands in contradistinction to the other. This is a statement of fact, not of opinion. Now, since the existing modifications are the result of evolution from the ancestral types, it follows that phylogenetically A’. A®, A’, ete., stand in closer relation towards each other than do the synoecic animals A’, B', Cl, D!, ete., the A’s forming one branch, the B’s another, the C’s a third, ete., of the phylogenetic tree. This differenee in the degree of blood-relationship between the geographical representatives (= geographically separate components) of one type and between synoecic types is very conspicuous in all classes of animals where adequate material has been conscientiously studied. From the point of view of a morphologist alone, all the animals which are clearly geographical representatives of one ( xxxvii ) another have a different standing in classification from the non-geographical entities. (2) If a small proportion of the specimens of A’ and A® are the same, and A! and A? (or one of them) are brought under conditions which favour the appearance of the characters of this small proportion in each case, bionomics teach us that A? and A? will in the course of time become more and more similar, overlap wider and wider, and become finally identical. That is to say, A' and A? cannot live together without fusion. However, if A‘ and A®, which differ, say, in 99-9 per cent. of the individuals, cannot exist together as separate entities, A? and A®*, which are connected by intergradation in the intermediate countries, or A? and A‘, of which the ranges of variation are contiguous, or At and A®, which are separated by a small gap, have no chance of remaining separate entities, if by accident brought under the same con- ditions of life. As there is a gradation in the morphological difference between geographical representatives (see Fig. 2, in which the curves D', D?, D?, etc., are Fie, 2. the geographical representatives), some being slightly, others more distinctly, and others again widely different, it is obvious that the chance of the representatives not overlapping in characters, under those altered conditions of life above referred to, becomes larger and larger, the wider the morphological gap is between them. From this point of view it depends, therefore, entirely upon the characters of the geographical representatives whether these can exist together or not, 7.e. whether there is a similar gap between A’ and A? to that between A! and B'. In other words, considering the gradation in the morphological differences, there are morphologically different geographical representatives which can certainly not exist together without fusing to one entity, and there are others which may be so far settled in their characters that a fusion will not take place. From what we have said it follows that it is wrong to treat all geographical corporeally different forms indiscriminately as being separated by that same barrier which prevents synoecic animals from fusing. As science does not knowingly perpetuate errors, the contention which we have demonstrated to be erroneous should be abandoned by those who claim their ( xxxviii ) work to be wholly scientific; and nomenclature, the language of classification, should therefore have a different formula for the two different conceptions, if classification is meant to be precise. And the language recording in formulas the results of classification must be precise, because science has specially invented the formulas to prevent confusion arising. However, before going further, let us consider another side of the question. It might be argued, with some degree of justification, that nomenclature is a practical invention for the convenience of the classifier (and those who make use of classification), and, as we said above, must not be turned into an inconvenience. Now, if the classifier is compelled to distinguish between geographically isolated forms which are equal to distinct synoecic animals, and such which are not, and again between geographical and non-geographical corporeally distinct animals, an amount of work is thrown on him which he could easily avoid by treating all definable forms in the same way, as being all co-ordinate. Those of our readers who follow classificatory literature * are aware that we are not stating an imaginary case, with the object—as so often happens in science and elsewhere—of demolishing an opinion which nobody entertains. The procedure above advocated is certainly a great simplification of work. But we may be allowed to state in response that a far greater simplification can be attained by putting all geographical modifications of the same type down as identical, nomenclatorially and classificatorially. If the splitter who indiscriminately calls everything definable a “species” claims the right to live, ourselves as impartial onlookers must concede the same right to the Inmper who treats everything not conspicuously different as being identical. These two opponents are almost hopelessly at loggerheads. We say almost; for we perceive some glimmer of hope of a good ending in the fact that, as in the struggle between two nations the victor becomes generally influenced by those details of civilisation in which the vanquished excels, both the lumper and splitter will profit from one another in the course of time, both remaining victorious in the struggle till the end, the one knocking over what the other considers distinct, and the other separating again what the former has put together as the same. Do not let it be truly ad infinitum. We sympathise with both; for we believe that we understand the cause of the struggle and the reason’ for its unreasonable continnance. When Linné invented his nomenclature (binominal, except most Lepidoptera) for his classification, he started with the conception that the animated world was composed of a great number of forms which, though variable to a certain—mostly small—extent, were fixed entities. These entities he defined as species, and designated each with a name. The introduction of a precise discrimination (or what was meant to be precise) between the species, and their fixation in classification by a name, gave a great impulse to collecting and studying the forms of animated creation. During the last century the material increased at such a rate that the number of species known to Linné became soon insignificant as compared with what * See, for instance, Verh. V. Intern. Zool. longress p. 910 (1902). ( xxxix?) more extensive collecting brought to light.* Among the new materials the systematist found a multitude of forms connecting by all kinds of intermediate grades a great number of hitherto well-separated “species.” In dealing with these intermediates systematists adopted three methods. The one class of describers, disregarding variability, thought to carry out in the correct way Linné’s method of classification and nomenclature by applying Linné’s term “species” to every group of individuals which they found to be definable. As every individual differs to some extent from the other, every specimen was naturally a trap for these anthors, who continually considered individual differences to be specific, and hence described an army of “ species” which had no standing at all. Another class of systematists, noticing the links connecting what otherwise appeared to be distinct “species,” were inclined to regard everything similar as being the same. They forgot that the first object of the describer is to distin- guish between what is distinguishable, and they were still further carried away by the reaction against the indiscriminate creation of species which necessarily set in. There was, therefore, a kind of excuse for the one as well as for the other class of systematists; the one student erring in being too zealous in applying throughout what he thought to be the generally adopted Linnean method of dealing with the animated world, and the other falling into mistakes by over-zealously correcting the errors (or what appeared to him as such) of the former. The right path lies, as it mostly does, in the middle between those followed by the extremists. A third class of students, keeping their mind unbiassed, were led along this middle path by their own power of discrimination. They learnt from the investigation of the mass of material in museums and private collections, and from observations ‘on living specimens, that neither everything similar is identical, nor everything dissimilar is specifically distinct. It is the lack of discrimination which prevents either extremist from finding the right path. However, the work of the splitter has a great advantage over that of the lumper. The differences which he points out between tne * The numbers of species described by Linné in Syst. Wat. ed. x. 1758 are as follows ‘~~ Mammalia . A ; é . a 6 : 6 + 184 Aves. F ‘ < 5 b 6 0 0 F - . 554 Amphibia. fF 5 5 ‘ . c é ‘ . 218 Pisces . A “ : c . . . 0 fs F ee cits) Coleoptera. 5 5 . . ‘ 5 . . 665 Hemiptera. c 4 “ 5 5 . . raed i Lepidoptera . A . . . 5 . A . 542 Insecta { Neuroptera . ty : . E 5 ° 7 60 Hymenoptera . 4 . : . a A 2 . 229 Diptera . 5 : é 3 : 4 5 5 LSS Aptera . . . : . . . . . - 229 Vermes . 927 Total . ; . . 4371 (Gals) animals are there. In the statements of fact he is correct; but he errs in the interpretation of these differences. His sight is keen, his reasoning—less so. The lumper, on the other hand, does not perceive the differences, or he perceives them only in a hazy way; he puts them down as insignificant and passes on, halting only if there is a conspicuous quantity of difference which impresses itself on his mind. His eye and reasoning are on the same level, and his work is generally scamped. As he depends in his judgment on the conspicuousness of the characters, he naturally falls constantly into the error of treating as the same what differs in non-contrasting colours or in vaguely perceived structures, and as distinct what differs in contrasting colours, in shape, size, and other easily noticed characters. It may appear ludicrous to the non-initiated, but it is nevertheless true, that in one and the same group of animals—for instance, Lepidoptera—the same kind of difference is considered by the same authors as being of no significance in the case of small forms, where it requires careful research to find the distinctions (Zpiplemidae, Geometridae, Thyrididae, ete.) ; while it is treated as being specific in the case of large ones, where even a dull eye cannot fail to perceive the difference (Papilio and other butterflies). Sombre- coloured animals (some Sphingidae, Noctuidae), small forms (Hpiplemidae), and such with a uniform pattern (Micronia, for instance), are generally great stumbling-blocks for the Iumper. ‘The author who treats everything definable as distinct has at least method in his errors; the author who depends entirely on what appears to him to be a sufficient quantity of difference is quite arbitrary in his judgment. The errors of an author arising from physical shortcomings, lack of training, and a certain flightiness in reasoning may leniently be passed over, if the mistakes are not persisted in when they have clearly been pointed ont. However, the direct misstatements as to variation and distribution, which the lumper is more liable to make than the splitter, are scarcely pardonable. If it is the object of the systematist to elucidate facts, and not to conceal them or to misstate them so as to mislead, it is wrong for an author to suppress distinctions which he has noticed; to refer to differences which are geographical, as if they were individual ; to consider characters as seasonal which he knows not to be seasonal ; to pronounce a form constant, if he knows only one or a few specimens ; to treat another as being individually variable, though he knows only single individuals from different countries. In short, it is wrong—and may become objectionable, because bordering ou charlatanism, if the misrepresentation is made deliberately— to represent anything to be what it is known not to be, or what on a little consideration it would have been known not to be. Neither the author who considers every form definable as being specifically distinct, nor his opponent who treats as identical everything of which the distinguishing characters do not appear to him to be conspicuous enough, fulfils one of the primary demands on the conscientious classifier—namely, to discriminate carefully between the differences presented by the various forms of animals he has to deal with, instead of lumping and separating indiscriminately. (xii) We are aware that a good many systematists, both among amateurs and professionals, have no higher object than naming and arranging the material in their collections, and maybe issuing books to enable others also to name and arrange the specimens, the knowledge aimed at being a knowledge of some distinguishing characters, and especially the name of the “species.” However, the knowledge of the alphabet does not carry with it the knowledge of the language, and he who knows the words and speaks a language is not yet a philologist. So there is also in classification a higher object to be attained than merely describing, baptising, and arranging in some arbitrary order the forms of animated nature. This higher object is to understand the phylogenetic relation between the forms, and on this understanding the scientific classifier bases his system. In order to comprehend the connection between the forms, it is necessary to know what it is that separates them. It was the one kind of difference to which we have referred before, separating the animals which exist side by side, the one effective barrier consisting of differences in the organisation of the animals themselves, which was the keynote to the Linnean Reformation of Natural Science and to the Darwinian Revolution. The individuals within the barrier form an entity which has an existence independent of all the other entities. Hach entity was in the Linnean classification understood to be a special creation, and the effective barrier to be intended to prevent fusion of the entities. This was the conception to which Linné applied the term species. And this was again the conception which formed the subject of Darwin’s Origin of Species. The great, mystery which the theory of descent sought to explain was the fact of the co-existence of such innumerable independent species, all separated by that gap which we know to keep the species apart. How did this specific barrier come into existence, if not erected by special creation? That is the fundamental question which is before scientists. The question is not solved by looking it straight in the face and then shelving it by applying the term “species ” to something else than what it originally meant. No friend of true research should let pass unchallenged what so many classifiers nowadays try to do—namely, to substitute for true species the geographical form. We know that a specific barrier exists between synoecic animals; we know that there is a morphological distinction between geographical representatives. Can the one barrier which we know to be specific honestly be replaced by the geographical difference which at the highest may be assumed to be specific, and of which we know that it is mot always specific? We shall oppose any such attempt at underhand shifting of the meaning of the term “ species,’ which would misguide the public and prevent the student himself from seeing clearly the question at issue. If the specific barrier is the result of the evolution of the organic world by natural causes; if, further, the multitude of species is the outcome of the divergent development of species into a greater number of species,—then we have to search for the rudiment (= beginning) of the specific barrier among ( xlii ) differences found within a species between the various component varieties and individuals. Therefore it is necessary, in order to understand the origin of the specific barrier, to study the varietal differences, and find out among which varieties there is a rudimentary specific barrier, and hence which varieties are rudimentary (= incipient) species. It has been shown by one of us* (and therefore we do not again fully enter into the same question) that the development of gamogenetie species into two or several species is not possible without an effective extraneous barrier between the varieties, which barrier prevents the fusion of the varieties, as does the specific barrier the fusion of the species, and, further, that this extraneous barrier is provided by geographical separation. Isolation of one or more mutating factors is the cause of the portion of a species subjected to them becoming different from the other components which stand under other influences. All our researches confirm this conclusion based on the facts of variation, and all attempts to demonstrate the possibility of the separation of a species into several without some kind of local isolation are fallacious in reasoning. Geographical variation leads to a multiplication of the species ; non-geographical variation at the highest to polymorphism. Geographical variation is, therefore, of another kind than non-geographical variation, and therefore geographical varieties have a different standing in the evolution of the organic world from the individual and generatory varieties. Geographical varieties as incipient species are the next classificatory category below species, just as subfamily is a degree lower than family, and no better term could have been invented for them than subspecies. With subspecies we designate, therefore, nothing else but the geographically separated different components of one and the same type, which components represent together a species. The criterion of a subspecies is not a certain amount of difference, but bodily difference and geographical separation. Synoecie varieties—i.e. varieties from the same locality—are never subspecies. We have to emphasise this distinction, as many authors constantly confound subspecies with non-geographical varieties. There are comparatively very few species which do not vary geographically. It was an ardent opponent of Darwin—Wiegand—who put forward as an argument against the theory of evolution that geographical variation was a conditio sine qua non for the correctness of the theory of descent, and that there was no such general basis for evolution. Systematists have proved by their minute research that geographical variation is the rule and not the exception, and they may be justly proud of this result of their untiring labours. Curiously enough, non-systematists do not generally seem to be aware of this result, nor to fully comprehend its bearing on the theory of descent. A species which has not developed into subspecies (= geographical varieties = geographical races or forms) may be individually or seasonally di- or poly- morphic, and similarly the individuals of a subspecies may all fall into seasonal and these into individual varieties. As the species of a genus are co-ordinate * See “Mechanical Selection” in Nov. Zool. iii. p. 426 (1896); “ Reproductive Divergence, etc., in Natural Science xii. p. 45 (1898), ( xliii ) with one another and subordinate to the genus, so are the subspecies co-ordinate with one another and subordinate to the species of which they are the components. Since Linné applied the term varvetas to the forms which are not specifically different, we do not see any reason against the use of this very convenient word in the same sense for all the components of a species which differ from one another. We understand, therefore, under variety not a particular category of the components of a species, but employ the term for all the different members of a species indiscriminately. The different categories of varieties must receive special terms in a precise classification, and special formulae must be employed for them in a precise nomenclature. We distinguish three categories of varieties, namely :— I. Individual variety.—The following terms are employed by us: (1) ad. = aberratio for individuals which stand outside the normal range of variation. (2) f. = forma in the case of di- and polymorphism. If a form occurs rarely, it may be termed /. aé., in contradistiuction to je norm. (3) 2-f. or d-f,, if the respective form belongs to one sex only. (4) f. loc. = forma alicuius loci, if, in the case of polymorphism, a form is restricted to one portion of the range of the respective variety or species. II. Generatory variety.—This variety is seasonal in Lepidoptera, and is designated as (0) f. t. = forma tempestatis. Il. Geographical variety or subspecies.—This is the highest category of varieties. As the term varietas includes also other varieties, it cannot be employed as such for the geographical variety except in a precise nomenclature; either a specifying attribute must be added (var. geogr.), or an abbreviation of another term chosen (subsp.). But we do not see that it is at all necessary to put any such abbreviation of a term before the subspecific name. We can do without the encumbrance of the abbreviation—what we can do without is unnecessary : and what is an unnecessary encumbrance in nomenclature, common-sense compels us to drop—by (6) Simply mutually agreeing that a subspecies is designated by its name added to that of the species without any abbreviation before the subspecific name. This means simplification of nomenclature, nothing else. The following diagram illustrates the nomenclatorial relation towards each ( xliv ) other of the various categories of classificatory units from the individual variety upwards to the genus :— fe faites subsp. ab. 18 Ue é. pete g—f. 9—f. subsp. 3o-f. | jet species 16 Ais 9—f, | of ab. ab. subsp. ab. ab. f. norm. eae subsp. species | be foat: BPEciee f. norm. f. ab. species f species £ There seems to us to be some confusion existing in the minds of some systematists with regard to the degree of phylogenetic relationship of the auimals classified and the nomenclatorial position of the units towards each other. Although it is the highest object of the researches of the systematist to elacidate the phylogenetic connection between the classificatory units from the individuals upwards to the family, order, etc., the linear arrangement of the units in the system gives but a very scanty elucidation of their evolution. If a genus is a development from another, it may be put behind the older one; but this method at once breaks down if there are several genera derived from one, especially if one of the derivatives has again given rise to a series of genera. And nomenclatorially the classifier can do even Jess. In nomenclature all the units of one category are co-ordinate units ; all the species, all the subspecies, all the genera, ete., are respectively co-ordinate with one another. The classifier cannot make any nomenclatorial difference whatever between phylogenetically younger and older genera, between the parent- and daughter-species, between the generalised and specialised subspecies. We have the same nomenclatorial formula for every genus (Papilio, Fringilla), for every species (Papilio priamus ( xlv ) Fringilla coelebs), for every subspecies (Papilio priamus poseidon, Papilio priamus priamus, Papilio priamus coelestis). If the classifier wishes to represent the probable phylogenetic origin of the units which stand co-ordinated in his system and nomenclature, he has to take recourse to the figurative tree, or must give a kind of pedigree, as we have done in the present work. Fringilla coelebs does not mean that coelebs is a derivation from Fringilla, but that it forms part of the genus Fringilla; and Papilio priamus poseidon must not be interpreted as signifying that poseidon is derived from priamus, but that it is one of the several components which together form the species Papilio priamus. This confusion of ideas has been occasioned by the unfortunate habit, which many classifiers cannot shake off, of regarding the first-described component as the typical form of a species, as the “Stammart,’* as the phylogenitically oldest portion of the species, or in the case of a genus as the phylogenetically oldest species of the genus, while it is merely the accidentally first-baptised form. Is the distinction between what is phylogenetically and what is nomenclatorially the oldest really difficult to perceive and to comprehend? The classification of the lower categories from individual variety to species is in a different position to that of the higher categories (from genus upwards). While the species and varieties are realities which can be tested by observa- tion of the live specimens and by experiment, there is no such test possible in the case of genera, tribes, families, etc. These higher categories are definable groups of allied species. The criterion of their being realities, or, as one is used to say, of their being natural, is threefold :— (1) The group must be definable—.e. must not so intergrade with another that there is no line of division, or that the line of division is arbitrary. (2) The contents of each group must be homogeneous. Elements of different origin, though perhaps similar in consequence of convergent development, must not be brought together. (3) Each higher category must have a separate definition based on other characters than those referred to in the definitions of the respective lower categories. The characters which make an animal specifically distinct do not make it also generically distinct, nor can the same character upon which a genus is based be employed to characterise a tribe or a family. To define genera and higher units is not always an easy matter. In order to render a definition precise, a close study is necessary of the forms which come under the unit defined, as well as of the forms of the allied units. The difficulties encountered have induced many authors, especially in Ornithology and Entomology, to propose names for genera, subfamilies, and families without attempting a definition. The naked names thus introduced are a fit testi- monium paupertatis for their authors. In our opinion, the thoroughness of the researches in systematic work can best be estimated from the degree of exactness * The word “Stammart"’ is much employed by German writers in this erroneous and misleading sense, See Reichenow, in Verh. V. Intern. Zool. Congress p. 911 (1902). ( xlvi ) of the definitions of genera and higher units. For the degree of exactness depends here (1) on the more or less intrinsic study of a larger number of forms than in the case of species and varieties, and (2) on the correctness of the author’s inductive reasoning. The classifieatory category coming in our system of classification next to the species is the genus. Some authors interpolate between genus and species a category they call subgenus. From a comparison of a number of subgenera which have been defined, we are able to state that they had either no standing, being groups of species quite arbitrarily put together, or they represented well- defined homogeneous groups—é.e. were equal to a genus. We do not see any possibility of distinguishing between a defined genus and a defined subgenus. To call one defined group a genus and another defined group a subgenus is quite arbitrary. As there is a gap between every two species, and mostly between the varieties as well, and, further, as the species fall into groups different in extent or different in composition, according to this or that organ being taken as the basis of the grouping, the limitation of the genera would be entirely left to the personal opinion and ability of each individual classifier, if there was no general definition of what kind of classificatory unit a genus is meant to be. Systematists differ, indeed, very much in the extent given to genera, some authors adhering to large unwieldy groups of the Linnean type, and others erecting a genus for almost every species, and sometimes even for subspecies. In order to check arbitrariness, to escape uncertainty as far as possible, and to make generic classification more stable, we think it advisable to define a genus as a classificatory unit one category higher than species comprising one definable group of species. In many cases the group contains only one species by the other members having become extinct, or by there being as yet only one species known, the other species being still undiscovered. In the present Revision of the Sphingidae it has been our special endeavour to give a solid foundation to the genera, supplementing and rectifying the vague or faulty definitions with which the workers in this group of insects have contented themselves. Though many genera hitherto considered to be valid haye been shown by us to have no standing, the number of genera of Sphingidae has been much enlarged, owing to the closer examination of the insects proving many groups of apparently similar species to be heterogeneous. We have laid special stress upon the genera as conceived by us representing stages in the evolution of the Sphingidae. There is nowadays a tendency among British Lepidopterists to imitate some American leading spirits in Lepidopterology in shifting the term “ family” (familia; designated by the ending -idae according to common consent) to a lower category than that to which it was originally applied. We do not see what good it serves to call, for instance, all the Hawk Moths together a superfamily, and the next divisions of it families. It is an entirely superfluous innovation, and only leads to confusion, like all shifting of ( slvii ) terms. We divide the Sphingidae into the following eight classificatory categories :— I. Individual variety. II. Generatory variety = seasonal variety. III. Geographical variety = subspecies. IV. Species. V. Genus. VI. Tribe. VII. Subfamily. VIII. Family. Before we proceed to state some of the general results of our study of the Sphingidae, we give a summary of the morphology of these insects, which will enable the reader to more fully comprehend some of the conclusions bearing on phylogeny and distribution. MORPHOLOGY OF SPHINGIDAE. The dorsal skeleton of the head (PJ. LXII. f. 6) is divided by two transverse sutures into clypeus (c/), epicranium (ecr), and occiput (occ). The last is always a narrow transverse plate more or less distinctly placed at an angle to the convex epicranium ; it is as a matter of course longer transversely in those species in which the eyes have a more oblique position than in those which have less obliquely placed eyes. Compare Pl. LXI. f. 6 and Pl. LXII. f. 6. Its anterior edge is generally faintly incrassate in the middle. The epicranium forms laterally the sockets for the antennae, which stand nearer the eye in some Hawk Moths than in others; the suture between epicranium and elypeus is just in front of the antennae, ending in the antennal grooves. The clypeus is the largest plate of the three; it is more or less strongly convex, especially mesially. It bears at the anterior margin the labrum (i, Pl. LXI. f. 6. 8. 9. 10. 11, also Pl. LXII.). The two are almost merged into one, the suture being mostly not distinct. The labrum is in most instances raised to a large, transverse, cariniform tubercle, which is generally vertical in front. It projects sometimes frontad over the base of the tongue, concealing the mesial part of the epistome (ep), and is occasionally very small (Pl. LXII. f. 3). The epistome is a transverse plate of variable dimensions situated in front of the labrum, with which it is so completely fused that it is not easy to say where the one begins and the other ends. It is especially large in a number of Ambulicinae with otherwise reduced mouth-parts (ey, Pl. LXL. via NOG UL FI GROUT Gig 20 tsp) This epistome covers the base of the tongue. When normal, it has a thin mesial lobe and a large process at each side. The mesial lobe varies in size and somewhat in shape, and is rarely absent (Pl. LXI. f. 10; Pl. LXII. f. 1), this occurring when also the other mouth-parts exhibit a high degree of reduction. ( xlviii ) The lateral processes (p, Pl. LXI. f. 6—11; Pl. LXII. f. 1—5) are designated “pilifer” by Kellogg *), and are often erroneously considered to be homologous to the mandibles. The normal pilifer is a curved obtuse process, concave and flattened on the innerside, and is beset on the inner surface with a great number of long stiff bristles which project over the base of the tongue, which they touch. The pilifer and its bristles undergo various modifications. The bristles become modified into scales, either partly or all (Pl. LXI. f. 11), or they become fewer in number and disappear finally nearly completely (Pl. LXI. f. 10). At the same time the pilifer may become shortened and lose the appearance of a process, being represented in the most reduced state known to us by a broad obtuse projection (Pl. LXI. f. 10). Two other modifications are represented by Pl. LXII. f. 4. 5. In fig. 5 the pilifer (py) is almost stalked, and projects far beyond the mesial lobe, which is large. In fig. 4 the whole epistome is enlarged together, produced forward, the pilifers being close together and very broad, and projecting little beyond the mesial lobe, which is very small. Quite different from these modifications is that observed in all the species of Choerocampinae, and only in this subfamily. Pl. LXII. f. 2 represents a species of Celerio: the pilifer (y) is long, somewhat twisted, and the apical portion is clothed inwardly with short bristles, while the proximal portion bears the ordinary long bristles, the two kinds of bristles contrasting very strongly with one another. “This character is of surprising constancy; its significance in classification will be understood, when the characters of the palpus and antenna are taken into consideration at the same time. Between pilifer and eye, supporting the former laterally, there is a more or less triangular projection, which is an enlargement of the brim which separates the eye from the large labial cavity of the underside of the head. We term this projection “genal process” (gp of PJ. LXI. and LXIL). The upper portion of this genal process is often distinctly separated by a suture, and corresponds to the mandible (md, Pl. LXII. f. 4), as pointed out by Kellogg, 1. c. The genal process is very large in Macroglossum (Pl. LXI. f. 9), Sesia, and allied genera, reaching often to the tip of the pilifer. In the Choero- campinae (in all of them) it is smaller than anywhere else (Pl. LXII. f. 2), not showing in a lateral view the distinctly triangular form observed in all other Sphingidae with well-developed tongue. The suture between genal process on the one side and labrum and clypeus on the other ends often (many Ambulicinae) in a deep groove (f, Pl. LXI. f. 10. 11). If we examine the underside of the head after the removal of the labial palpi (Pl. LXI. f. 7; Jip is the groove in which the palpus is inserted), we find again the pilifer and the genal process between eye and tongue, and observe below the pilifer close to the tongue on each side a short process, dilated apically in the species figured, which process is the remnant of the maxillary palpus (map). It is in most cases densely clothed with loug white scales, which project beyond the pilifer and genal process, being visible also in dorsal and lateral aspects of * Amer. Naturalist xxix.\p. 546 (1895). ( xlix ) the head (Pl. LXI. f. 6. 8). The size of the vestigial maxillary palpns is not constant in the family, nor has the palpus always the same shape. The transverse arched stripe of chitin between the Tabial palpi is the mentum; in front of it we find often a vestigial, very feebly chitinised submentam. The before-mentioned mouth-parts of Lepidoptera have attracted much attention on the part of scientific entomologists since Walter’s now famous paper on the mouth-parts of Micropteryr.* The distinctions exhibited by them within the families of Frenata have, however, not been made use of in classificatory work. The parts are covered by the labial palpi as a rule, and are not visible without pushing the palpus away from the head. A drop of benzine, or, better, a drop of alcohol, applied to the base, is generally sufficient to make the palpus so flexible that it is movable, and allows the genal process and pilifer to be studied without injury to the specimen. The two parts of the caputal appendices which remain to be discussed, tongue and labial palpus, are better known to the classifier, though the descriptions given of them go seldom beyond length of the former and ontline and general aspect of the latter, The length to which the tongue has developed in the family Sphingidae is an exceedingly striking character. Here we find the longest tongue of all insects. But what is far more interesting for the student of comparative morphology as well as the classifier is the fact that the length of the tongue yaries in this family to such an enormous extent as it does, the extremes being represented by Cocytivs, in which the tongue is sometimes little short of 25 cm., and Polyptychus, where we find species with a tongue represented by two tubercles barely longer than 2 mm. A comparative study of the tongue (glossa) of Lepidoptera is a desideratum. It is formed by the first pair of mavxillae, and consists, as is well known, of two halves closely applied to each other (Pl. LXII. f. 2). Each half is concaye on the inner side, and bears at the upper inner edge a very dense fringe of ciliae. The trans-section is in Sphingidae short kidney-shaped, or nearly circular, apart from the inner concave portion. Laterally at the base the tongue has very often a patch of minute hairs; in a few cases hairs are found all over the dorsal surface. Within the cavity of each half we find, in dry specimens, a large trachea and the residue of the dried-up muscles, nerves, etc. The sucking-tube itself (PI. LXII. f. 2, tv) formed by the two halves of the glossa is closed above by the fringe, the ciliae of which are soldered together to form a membrane, which is often quite smooth, showing no trace of transverse striation indicating the ciliae. When the tongue becomes reduced, the two halves are less firmly applied to ove another, and the transverse striation of the closing membrane of the tube becomes distinct, till with the further reduction of the glossa the two halves separate and the closing membrane assumes the form of a fringe of separate ciliae (Pl. LXI. f. 11; Pl LXII. f. 4. 5), this fringe finally disappearing (Pl. LXI. f. 10; Pl. LXIL. * Jen. Zeitschr, Naturm. v, 5, p. 75i (1885). d Cis) f. 1). The fanctionless tongue loses the transverse annulation ; it is very feebly chitinised, and varies individually in length. It not rarely bears scales, and is occasionally tuberculated. The longest tongue oceurs in the tribe Sphingicae, which tribe contains, however, also species with a very short and functionless tongue, and one species with just a vestige of it (llenbechia). A short tongue is frequently found among the Ambulicinae, in which subfamily it is always shorter than the body, but often strong, and in the latter case not showing any sign of reduction. In the Sphingulicae the tongue is also very weak and short in most species. In the other groups of Sphingidae the tongue is never excessively long and never very short, always preserving the ordinary structure and rigidity. It is scarcely necessary to point out that the functionless glossa is a derivation from a tongue efficient as a sucking-tube. As varied as the other mouth-parts are also the labial palpi, shortly designated as “palpi” in Lepidoptera. Though this designation is, strictly speaking, not correct, it is very convenient and quite precise enough, as there ean be no doubt that the labial palpi, and not the vestigial maxillary ones, are meant, when speaking of the palpi. The palpus, if not reduced, is large, broad in lateral aspect, closely contiguous to the head, and has a short third segment. A palpus like this does not occur outside the family. However, where the palpus is reduced, the general aspect is insufficient to recognise it as a Sphingid palpus. The most slender and at the same time longest palpus is found in Z7nostoma and in the 3 of Cressonia, in which latter genus the ¢ palpi are strongly divergent. A very large and rounded palpus is met with in Pachylia, Eurypteryx, Protoparce, and other genera. Reduced palpi oecur abundantly among Ambulicinae and Acherontiinae, while the palpi of the other Sphingidae are mostly of medium size or large, very seldom small. The size of the palpus depends on the width of the segments and upon the scaling ; a broad segment covered by short scales appearing much slenderer than a narrower one with long erect scaling (Pl. LIX. f. 26. 27). It is, as a rule, also the scaling which gives the terminal portion of the palpus its particular shape. The triangularly pointed palpus and the broadly rounded one, as described in systematic work, may have similar segments when denuded. If one speaks of the shape of the palpus, one means the palpus inclusive of the scaling as it appears in a perfect specimen. There is distinct sexnal dimorphism in the palpus among Ambulicinae, the palpus of the male being often larger than that of the female. There are always three segments; the third is, however, nearly always very short and concealed in the scaling of the second, projecting as a little knob. In the few cases where the third segment is more distinct it is conical, sometimes naked and horn-like (Cocytius ; Xanthopan); it is never long, slender, and rod-like. In narrow palpi it is often as broad at the base as the second segment is at the end. The first segment is the longest as a rule, but there are many exceptions (hh) in which the first is not longer or is shorter than the second. The segment is curved, lying along the eye. The inner surface (Pl. LIX. f. 26. 27; Pl. LX. f. 1) is more or less regularly annulated or wrinkled, flattened, or slightly convex, or somewhat concave. It is naked, except the edges, with some long hair-like scales ; or it is more or less loosely scaled for the greater part. The scaling at the apex of the first segment (Pl. LIX. and LX.) is either short, or long and rough, or long and quite regular, affording in several cases conspicuous characters of taxonomic value—as, for instance, in the subfamily Choerocampinae, where the two Neotropical genera and the cosmopolitan genus Celerio have the scaling always rongh and irregular, while all the allied Old World genera with the exception of Pergesa and Mhodafra, derivations from Celerio, have it regular (PI. LIX. f. 19—21. 26. 27). A character of the greatest importance in the classification of the Hawk Moths is found at the base of the first segment. That is a patch of variable size of short (and doubtless sensory) hairs, which is always present in one section of the family (a, Pl. LIX. f. 26. 27), except a few reduced forms, and eqnally constantly absent from the other section (Pl. LX. f. 1). The trustworthiness of the distinctive character was discovered after we had separated the Acherontiinae (= Acherontiicae + Sphingicae + Sphingulicae) and Ambulicinae from the rest of the family on other grounds. This basal patch was found in Butterflies and treated upon at some length by Renuter.* It is of wide occurrence in Moths. Its absence from Acherontiinae and Ambulicinae has nothing to do with the reduction of the palpus, as it 1s absent from the large palpi of Protoparce and other Sphingicae, while it is found in the other subfamilies on the relatively small palpus of Oryba, Berutana, and other genera. The basal patch is present in Geometridae, Notodontidae, Agaristidae, Noctuidae, Pyralidae, etc., etc., and assumes sometimes an obviously distinetive form. We have not noticed it in Satwrnitdae and allied groups, nor among Lasiocampidae, Bombycidae, Eupterotidae, and some other groups. How far this organ can be made use of in the classification of these families more complete research must show; but we are justified in maintaining that the basal patch will prove itself elsewhere an equally good distinguishing character as we find it to be in Sphingidae. The first segment of the palpus exhibits also on the outer side characters of taxonomic value. There is a kind of transverse crest near the eye in Basiothia, Aleuron, and Unzela, and the apex of the segment is strongly convex externally or angulate in these genera; the crest is also found in Gurelca and Sphingo- naepiopsis. A peculiar modification is found in one genus of Acherontiinae (Megacorma) and in a great number of Choerocampinae. It is illustrated on Pls. LIX. and LX. There is at the apex of the segment, ventro-laterally, a space devoid of the ordinary scaling, being either quite naked or clothed with a few long hair-like but flat scales. The scaling around this naked space, * Acta Soe. Se. Fenn, xxii. 1 (1896).—Reuter says that the basal patch of Heterocera is never elevated as in Rhopalocera. We find that it is sometimes raised to a conspicuous ridge, for instance in Pyralidae. (in) which is often somewhat concave, is more or less regular, especially ventrally, and, surrounding the naked space, forms a kind of cavity (gr, Pl. LIX. f, 13. 16. 17. 18. 25; Pl. LX. f. 3). The naked membrane is doubtless sensory, but we could not find any external sensory organs in the dry specimens. The long hair-like scales situated in many species on this naked membrane are not of a sensory nature. There is every intergradation between a rudimentary cavity and a large regular one. This specialisation is found only among Oriental and Aethiopian SpAingidae, not in American ones. The joint between the first and second segments is exposed in the species which exhibit the cavity. A naked and exposed joint is also met with among Améulicinae, but no cavity. The second segment undergoes many modifications in shape; it may be subeylindrical, quadrangular, triangular, ovate; it may be longer or shorter than broad, or square. In Tinostoma it is three times as long as broad. It is angulate at the upper outer corner in Alewron and Unzela. The scaling of the two palpi generally covers the base of the tongue; but there are instances where the mouth-parts are exposed. This is the case when the scaling of the second segment is very short, as in Pl. LIX. f. 10. 11. 26. The inner surface of the second segment exhibits some remarkable specialisa- tions. It is normally scaled all over, but we find the scales very small and rather dispersed in Huchloron (Pl. LIX. f. 24), so that the membrane is partly naked. In all the other species of Choerocampinae these small rounded scales have all, or nearly all, disappeared, leaving the segment bare except at the edges (Pl. LIX. f. 19—21. 26. 27). The character is quite constant, and oceurs only in those species which agree also in certain characters of the pilifer and of the antennal end-segment, and form the subfamily Choerocampinae. The scales at the upper apical angle of the segment with naked inner surface are either short or form a conspicuous tuft, which projects ventrad (Pl. LIX. f. 19—21. 26. 27). This difference is of importance, the absence of the tuft being characteristical for the fifty species of the purely American genus Xylophanes, which has, unlike the Hastern representatives of the subfamily Choerocampinae (Rhodafra and Pergesa excepted), the scaling at the apex of the first segment irregular, as stated above. We have not ascertained the trne nature of the naked membrane, but assume that it serves as an organ of sense. A specialisation reminding one of that just described is found in the genus Psilogramma of the subfamily Acherontiinae. Here the segment bears a naked longitudinal stripe not far from the upper edge, the stripe appearing as a prolongation of the naked part of the first segment. The modification does not occur in the otherwise very closely allied genus Lewcomonia. A third modification peeuliar to the Acherontiicae, but vestigial also in one genus of Sphingicae, is represented by figs. 1 and 2 of Pl. LX. Here the second segment is concave on the inner surface; the scales at the edge of the cavity (c) project over it, forming a kind of roof. The concave part is either practically naked or scaled. The transition from the vestigial groove of i ( liii ) Xanthopan to the deep and naked cavity of Acherontia and Herse is found in Megacorma and Coelonia. The third segment bears at the end a small and deep cavity of a sensory nature. We have not studied this organ. The great variety in the structure of the antennae” of the Heterocera is repeated to a lower degree in most families of larger extent, and this makes it generally impossible to give of these organs a short family diagnosis which is true of all the species of the respective groups. The stereotyped description of the Sphingid antenna handed down from Linnean times applies only to a portion of this family. The prismatic, clubbed, and hooked antenna is by no means common to all the Sphingidae, a large proportion of the Hawk Moths deviating widely from this type. However, comparing each type of antenna of the present family with the corresponding types of other families, we shall always find some difference, and very often some striking difference, revealed by the closer examination of the special structures. The generalised type of antenna of Lepidoptera is, in our opinion, that in which the dorsal surface is wholly scaled, while the ventral surface is scaleless, being clothed instead with a dense covering of fine hairs. This type is derived from an antenna entirely ciliated. Besides the protective scaling and the sensory ciliation there are other organs present, such as sense-bristles and -cones, both of which are rather easily discernible and of considerable taxonomic value. The scaling of the butterfly antenna is reduced apically and in a number of groups altogether absent, except the first segments (Danainae, Papilios allied to sarpedon, etc.). In Heterocera we find the scaling extending to the last or last but one segment, but meet also with antennae which have lost the scaling (Saturniidae and close allies ; Pelochyta). The arrangement of scales in two recular transverse rows on each segment which is prevalent among Butterflies, and is found also in a very great number of Moths, does not obtain among Hawk Moths except on the distal segments in a number of species, the scales being comparatively small and very numerous. Where among Butterflies and Moths the area covered by the fine sensory ciliae is reduced, the ventral surface becomes often partly scaled, the loss of the sensory function being followed by, or going hand in hand with, the appearance of scales, in Lepidoptera the normal clothing of the epidermis where no organs with special function are required. The proximal segments of the antennae of Sphingidae are occasionally nearly or totally scaled—namely, where the antenna is strongly clubbed. That means, where the sensory function is more concentrated distally, the proximal segments, which are reduced in width and function, have acquired an ordinary covering of scaling on the underside. This is most evident in Rhopalopsyche, a genus with also otherwise remarkable antennae. The appearance of scales as a sign of loss of function, or of weakness, is strikingly illustrated by other organs. We have seen above that the reduced tongue becomes scaled in some instances, and that the bristles of the pilifer are replaced in many reduced species by * Compare Bodine, rans. Amer. Ent. Soc. xxiii. p. 1 (1896); Jordan, Nov. Zool, vy. p. 374 (1898), ( liv ) scales. A further confirmation of the view just expressed, that the ventral sealing of the antenna is a relatively young character, we find in the fact that there are sometimes scales on the underside of the antennae of hybrids where there should not be any, these scales being doubtless an expression of weakened vitality. We predict that by breeding in and in specimens will result which show a more extended antennal scaling than the normal individuals. The antennae of the Aegeriidae and Castniidae, which resemble in general appear- ance somewhat a clabbed Sphingid antenna, differ essentially from the latter in having the greater part of the ventral surface scaled, agreeing in this respect with the antennae of a great many Zineidac. Among the latter family (or group of families, perhaps), and among the Limacodidae, we find antennae which are almost entirely covered with scales. Before following the somewhat complicated development of the sensory surface, we shall shortly refer to the sense-cones and sense-bristles. The sense- cones discovered by Bodine do not occur in Rhopalocera, but are widely distributed in Heterocera, They are ventral, mesial, and apical in Sphingidae, one on each segment, except the end-segment and the basal one or ones (PI. LX. f. 4—29, ec), from which they are absent. The cones of the distal segments are generally more prominent than those of the proximal segments. They do not always stand exactly at the apical edge of the segment, being not rarely removed somewhat basad (compare, for instance, Herse convolvuli). However, they are never absent and never abandon the mesial position in this family. They are wanting in the Castniidae, Aegeriidae, and Zygaenidae ; their absence is a distinctive character not difficult to recognise. They are present on the clubbed antennae of Agaristidae and Cocytia. The sense-bristles are stiff hairs of varying length. The Lepidoptera most primitive in respect to these organs have a complete belt of such bristles on each segment. Among Rhopalocera we find such a belt preserved in Lycaenidae and Hesperiidae; we meet with it again among the Jugata and many Hetero- cerous Frenata. Some of the dorsal bristles are, however, generally reduced and covered by the scaling. This is the case also in Sphingidae. The normal number of the bristles found in the Hawk Moths on the non-scaled surface, if we except the end-segment, is two on each side (Pl. LX. f. 14), one being dorso- lateral, the other ventro-lateral and basal. The number is sometimes doubled ; but there is never a complete transverse series, and the bristles are never apical, the Sphingid antenna differing therefore obviously from that of of the clasper. The processes are either firmly connected with the sheath, of which they are an outgrowth or a prolongation (Pl. XXX. f. 22. 48), or they are more or less movable against the sheath, the base of the process at the juncture with the sheath being partly membranaceous, forming a kind of joint (Pl. XXX. f. 31; Pl. LIV. f. 1—12). The long and slender whip-like process found in Perigonia, Sesia, and some allies bears two sensory bristles at the end. We can distinguish two kinds of processes: (1) the process is a prolongation of the apical edge of the sheath, and is accordingly flat or concave on the innerside (P]. XXIX. f. 9. 12); or (2) the process is an outgrowth from the surface of the sheath, an enlarged tooth, and accordingly more or less conical and hollow, at least at the base (PJ. XXIX. f. 47; Pl. XXX. f. 38). The processes are short or long, single or double, simple or dentate. The most remarkable is that of 2. sagra (Pl. LIV. f. 18). The teeth on the processes and on the sheath are small or large, single or in patches, irregularly distributed or seriated, forming serrate ridges. They are in most cases solidly connected with the sheath, not breaking off easily. There is, however, a second kind of tooth, which is easily detached from the sheath. This tooth, which has a central and three or four lateral branches, is found in Theretra latreillei and allies (Pl. LVIIL. f. 4—7). The spine-like teeth of Xylophanes nechus (Pl. LVII. f, 18. 19) also break off easily. The same obtains in centroctena ruther fordi. Within the penis-sheath we find the membranaceous penis proper, the duct of the sperma. This duct can be pushed out, and enters the vaginal orifice with the help of the accessory appliances. The duct has in most cases an armature of its own. It is either partly beset with small teeth (Pl. XXIX. f. 34. 41. 42; Pl. XXX. f. 12), or the teeth are enlarged to stilettos (Pl. XXIX. f. 40; Pl XXX. f. 44). In other cases we find two or three slightly more strongly chitinised denticulate processes of various lengths and shapes (Pl. XXIX. f. 7. 8. 9. 21; Pl. XXX. f. 16—19). Or the love-daggers are strongly chitinised, acute or club-shaped (PI. LY. f. 44; Pl. LVI. f. 41). The armature of the penis-sheath and penis serves a double purpose. The firm processes and teeth of the penis-sheath are grasping-organs like the harpe. But the processes are often far too long for that purpose alone. Now, the fact that the peculiar dentition of Theretra latreillei and allies is easily detached, and the detached teeth are fonnd in the vaginal cavity of the females, makes it evident that the armature of the penis and penis-sheath is a means to stimulate the female, analogous to the apparatus found, for instance, in some Rodents. : The sexual apparatus is of great taxonomic. value, and we have taken special care to dissect as many species as we could. Ina number of cases the apparatus is the only safe guide in the recognition of species. From this point of view the Sphingidae can be divided according to the sexual armature into the following categories :— (1) Species which are not different from their nearest relatives in these organs. —Nephele is an example of this kind. Some Choerocampinae allied to gallii ( Ixxxiii_ ) and euphorbiae also do not show any apparent differences. Pyotoparce florestan and Chlaenogramma jasminearum, though generically distinct, have the same sexual armature. This kind of identity should not be confounded with similarity arising out of reduction. It happens that members of different subfamilies become similar in one or the other organ in consequence of the loss of special structures. Such species are, however, never the same in all parts of the copulatory apparatus, as is the case in the insects mentioned before, the list of which is by no means exhausted by the few species used as illustrations to show that it is quite erroneous to maintain that all specifically distinct Lepidoptera exhibit differences in the copulatory organs. (2) Species which are different from the nearest relatives, but do not show any marked geographical variation in the sexnal armature.—Here belongs by far the greater proportion of the Hawk Moths. The specific differences are very slight or very conspicuous, or intermediate in degree. They may be found in one single organ, or in more, or in all. The differences may be slight in some species, and great in others of the same genus. Two species may differ strongly in external features, and little in the sexual armature, or the reverse may be the case. There is every conceivable gradation in this respect. There is always some individual variation, as a matter of course. When examining the armature of only one individual, one is sometimes induced to consider a certain outline or structure as specific, while it is in fact only an individual character ; if possible, a number of specimens from different localities should be examined. Individual variation is most obvious in species with complex structures, but it is here far less easily misleading than in the case of simple structures, because the specific differences are as a rule more conspicuous there than here. (3) Species which differ in the copulatory organ from the allies and vary in themselves geographically.—Geographical variation is most often met with and is most conspicuous in those forms which are sedentary in habits. Sluggish species with functionless mouth-parts and reduced power of flight, species of which the sole function as imagines is propagation, are especially liable to develop into geographical races with differences in the sexual armature. The phenomenon occurs often among