59.57,92(7) Article VII.-PHYLOGENY OF CYNIPID GENERA AND BIOLOGICAL CHARACTERISTICS1 BY ALFRED C. KINSEY PLATE XXXII CONTENTS PAGE INTODUCTION ....................................................... 357a DATA CONCERNING EVOLUTION.357b Radial Cell.357c Radial Vein...................................................... 358 Second Abdominal Segment ....................................... 360 Host Plants .................. 361 Gall Structure .................. 365 Reproduction .................. 369 Alternation of Generations ................. 372 Other Considerations ................. 382 PHYLOGENETIC POSITION OF GENERA.......................... 385 Aulacini.......................... 385 Aulacidea ........................... 385 Phanacis ........................... 387 Timaspis ........................... 387 Aylax ........................... 387 Diastrophus........................... 389 Gonaspis .......................... 391 Rhoditini................I ......... 391 Rhodites ........................... 391 Cynipini .......................... 394 Neuroterus.; - 394 Disholcapis.397 SUMMARY.399 BIBLIOGRAPHY ......... 402 INTRODUCTION Up to the present time the relationships of the genera or even species of the Cynipidae (Hymenoptera) have been poorly understood. Species have been catalogued without much attempt to discover an ar- rangement which would indicate evolution, and the genera have been in large part unnatural groups containing large numbers of unassorted, diverse organisms. Moreover, few suggestions have hitherto been made as to the origins of the peculiar biologic phenomena so character- 'Contribution from the Entomological Laboratory of the Bussey Institution, Harvard University No. 165. 357a 357b Bulletin American Museum of Natural History [Vol. XLII istic of the more specialized gall-wasps. Information as to the evolu- tion of some of these phenomena is highly desirable and nothing of the biological characters of such specialized creatures can be adequately learned except as it is studied in connection with the morphological char- acters, careful taxonomic work, and observations of the living insects. The data here assembled is the result of a study of the Cynipidae of the world, actual specimens of almost ninety per cent of which have been examined. Much information has been gained by a study of living material. The conclusions reached, as yet only incompletely applied to the genera of the family, will serve as the basis for a thorough revision of the classification of the Cynipidae which I hope to be able to publish soon. I am under great obligation to the same friends who have aided my other studies of the gall-wasps: Dr. Wm. M. Wheeler, Prof. C. T. Brues, and Prof. Irving W. Bailey of the Bussey Institution of Harvard University; Nathan Banks of the Museum of Comparative Zoology; Charles W. Johnson of the Boston Society of Natural History; and Dr. Frank E. Lutz of The American Museum of Natural History. DATA CONCERNING EVOLUTION RADIAL CELL (Table I) The wing venation of the Cynipidae is remarkably uniform through- out the whole range of species. Only the genus Eschatocerus, with one known species, shows any considerable modification from the type; in that species the first and second cubital cells are closed by the extreme reduction of the second abscissa of the radius and of the second cross- vein. The related Figitidae show a wide range of variation from the complete cynipoid venation to a condition almost as specialized as that found in those Chalcididae which have retained only a single vein. In the Cynipidae the modifications are very slight, and these, e. g., the presence or absence of the areolet, the extent of the cubital vein in particular, the extent and continuity of other veins, and the angles of the veins, are characters mainly of specific or even individual worth, sometimes differing on the two sides of the same specimen. Nevertheless, it is possible to perceive two lines of modification of the venation which seem to indicate something of group relationships: (1) the radial cell, varying from a completely closed to a wide open con- dition, find (2) the first abscissa of the radius, varying from an arcuate to an angulate condition. 1920] Kinsey, Phylogeny of Cynipid Genera and Biological Characteristics 357c It may be considered that the open radial cell is more highly special- ized than the closed cell, for the process.of openiiAg in this instance involves the disappearance of the marginal vein and later of the extremie ties of the subcostal and radial veins. It would be surprising to find that species with the open radial cell had developed the veins necessary to close the cell and thus originated the closed-cell forms. Evidence df evolution based on a consideration of this point alone would be meager enough, but if it parallels the data of other sorts it will lend that much more weight to the final conclusions, and at a couple of points it will bridge gaps in the history. From the table it will be seen that the character of the radial cell is of generic importance. Although it is true that Aulacidea is separated from Aylax primarily upon this character and that this is the best reason for considering Aulacidea a distinct group from and more primitive than Aylax, nevertheless, in many other instances where the genera arg founded upon other morphological characters, it will be seen that the condition of the radial cell is as good a generic character. Thus, for instance, Disholcaspis is a genus founded on peculiarities of thoraci' sculpture, the size and shape of the second abdominal plate, and the shape and proportions of parts of the head, among other very definite characteristics. The occurrence of the open radial cell in all the known species of that genus suggests that the genus as a unit is descended, rather indirectly, from a group with the closed cell. On the other hand, in Rhodites, a genus maintained likewise on abundant, definite morpho- logical and biological characters, although thirty of the species have the cell entirely closed, six species have the cell more or less open. It seems probable that in this instance the group has originated directly from a.closed-cell genus, and that comparatively recent evolution within Rhodites itself has given rise to the few species which possess the open cell. The order in the accompanying table is that of the apparent order of evolution, as indicated by this sort of evidence only. Again, I ac- knowledge the meager value of these considerations, except they be judged in connection with the other sorts of data present in this paper. 358 Bulletin American Museum of Natural History [VOl. XLII TABLE I. RADIAL CELL Genus No. of Species Nature of Radial Cell Ceroptres Entirely closed Synergus i60 Periclistuws -410 Aulacidea 22 Phanacis 2 Timaspis 7 Closed in 1 sp. Partly closed in 1 sp. Entirely open in 5 sp. D)istrophus 14 Closed in 1 sp. Open in 13 sp. Rhodites 36 Closed in 30 sp. Open in 6 sp. Aylax 26 Entirely open. Neuroterus 54 tclosed in 6 sp. Open in 48 sp. Cynips (in European sense) Entirely open. Disholcaspis 27 Oe us Amphibolips 25 All other Cynipini Open, usually entirely open. RADIAL VEIN Plate XXXII The first abscissa of the radial vein of the wings of the Cynipidae varies from an arcuate to a sharply angulate condition, some species showing a pronounced, vein-like projection into the radial cell at the apex of the angle made by the vein. Every degree of gradation between the extreme forms may be found. Indeed, so remarkably complete is the list of transitional forms that it was this exhibition which first drew my attention to the existence of evidence of the course of evolution in the group, and I have failed to find any other one line of data which offers as complete a story as that presented by these gradations in vein form. A sufficient display of this variation is shown in the wings of the forty-five species which I figure to remove any necessity for an account of the details of conditions in particular species. It is to be remarked that no great variation in the vein is to be found among the species of a single genus, i. e., the form of the vein is a generic character, indicating the same lines of generic limits which were drawn originally after considerations of very diverse morphological characters. In the Figitidae, in Aulacidea, Aylax, Neuroterus, Disholcaspis, Cynips (of European authors), and in Apmhibolips every species which I have seen agrees in details of venation with a pattern typical for its genus. 1920] Kinsey, Phylogeny of Cynipid Genera and Biological Characteristics 359 But in Diastrophus and Rhodites we find both an arcuate and an angulate condition occurring in the same genus, though the differences between two species at the extremes of variation within the genus are not to be compared with the extreme differences which occur between the genera. Apparently the form of the first abscissa of the radius gives evidence not only of group descent but also of the evolution of the species within a genus. Such a complete set of data, interpreted in the light of other evidence, should prove of great value for discovering an arrangement of the species of the family which will indicate the natural order of rela- tionships. The problem with which we are now confronted is to discover where in the series was the starting point, the primitive condition-of deciding in which direction development has proceeded. If, in trying to solve this question, we seek evidence in the related families of Hymenoptera, we find that apparently very little help is to be obtained there. In the Figitidae, which are mainly parasitic species, every species of the hun- dreds I have examined shows an arcuate vein, which isoneof the extremes to be found in the Cynipidae. Shall we assume that the gall-wasps have originated from the more primitive figitids or that the figitids have originated from the most highly specialized of the cynipids? I think that the story might be read with equal justice in either direction if the whole of our information came from the wing-venation.