Cynips

“ We have already shown that the short-winged species represent more specialized developments of the long-winged stocks. One short-winged and one long-winged species of Cynips is known from central Mexico, but our knowledge of the gall makers of that country is still insufficient to make it certain that our genus did not originate somewhere in northern Mexico. ”

“ Of all the areas occupied by Cynipidae in the United States, this Northeastern area, the Eastern mountain country, and areas immediately adjacent to these are the only ones which had a Pleistocene history that would have provided the opportunity for the multiplication of hybrid individuals and which would have offered a subsequent isolation sufficient for the origin of new species. The apparent restriction of the hybrid Cynips to those areas seems confirmation of our explanation. ”

“ We may repeat that we know no single character, morphologic, physiologic, or biologic, by which a Cynips may invariably be recognized. A hairy thorax, complete parapsidal grooves, undivided foveal groove, hypopygial spine which is broadened nearer the tip; a monothalamous, fundamentally spherical, separable leaf gall, occurring on a white oak; and the maturing of the adult early in the fall with emergence delayed until the winter—this is a combination of characters that will distinguish most of the agamic forms. ”

“ Some will consider that individuals are so variable and interbreeding Mendelian races so abundant in nature that taxonomic classifications can be nothing but contrivances without biologic significance. And yet, when the Colorado of the West first cut into the Colorado Plateau the specific stocks of Cynips were in existence, and thruout the years that the Canyon has been cutting, even down onto the present, these complexes which we call species have maintained their identity. ”

“ In nearly every one of the species the agamic form has the thorax hairy, altho there are few hairs on the very small, nearly wingless insects which are the most northern varieties of many of the European and American species in Cynips. In every species without exception the hypopygial spine is broadest posteriorly, and the structure terminates in a well-developed tuft of hairs. ”

“ This widespread confusion in the interpretation of Cynips has not been wholly consequent on the difficulty of interpreting the relatively uniform structures of gall wasp species. Felt (Journ. Econ. Ent. 19:672) considers the situation due to the complex life cycle of our insects and to the failure of a sufficient number of economic entomologists to turn to cynipid taxonomy as an avocation. ”

“ Following the suggestion of Lacaze-Duthiers, all these workers have found four fundamental zones of tissue in most cynipid galls. These zones have been called the nutritive, protective, parenchyma, and epidermal layers, and in the degree and character of the development of each of these the European students have seen an essential uniformity of structure among the European species of Cynips. ”

“ The original concept of Acraspis was based on the well-known, short-winged species pezomachoides and erinacei (fig. 3) of the eastern United States. All of the close relatives of pezomachoides are similarly short-winged insects in the agamic generation; but ever since Triggerson's studies (1914) of the life history of Cynips pezomachoides erinacei we should have known what the long-winged ancestors of the group looked like. The experimentally proved bisexual generation of erinacei is an insect with fully developed wings (figs. 1–2) , and other characters typical of long-winged Cynips. ”

“ For this reason, in the publication of new specific names in Cynips, it will be wise to avoid all names that have ever been used in the family Cynipidae. Names applied to galls only, or such combinations with Cynips as antedate the description of the insect itself, have not been considered to have nomenclatorial standing and are not included in this list. Thruout these references, Dalla Torre, 1893, Catalogus Hymenopterorum, volume 2 (Cynipidae) , is abbreviated to D.T. 93: ———. Dalla Torre and Kieffer, 1910, Das Tierreich 24:, is abbreviated to D.T. & K. 10: ———. ”

“ About fifty years ago Hartig reared large numbers of certain species of gall-flies from their galls, obtaining from 28,000 galls of Cynips disticha about 10,000 flies, and from galls of C. folii 3000 or 4000 examples of this species; he found that all the individuals were females. His observations were subsequently abundantly confirmed by other naturalists, among whom we may mention Frederick Smith in our own country, who made in vain repeated attempts to obtain males of the species of the genus Cynips. ”

“ He found, for instance, that some gall-flies of the genus Aphilothrix produced a series of generations the individuals of which were similar to one another, and were all females and parthenogenetic. In some species of the old genus Cynips no males are even yet known to occur. A very curious observation was made by the American, Walsh, viz. that of galls gathered by him quite similar to one another, some produced speedily a number of both sexes of Cynips spongifica, while much later on in the season the remainder of the galls gave rise to females only of an Insect called Cynips aciculata. ”

“ Because of the poor distinction of several of the genera of the oak gall makers, it seems unwise to employ the generic titles Cynips, Callirhytis, Philonix, etc., until a careful revision of the genera of the family has been made. For that reason, I have used the name Andricus, the most meaningless of all these titles, with a number of the forms.
To distinguish the alternate generations of a species, I have had to adopt a new form of naming. It is highly unreasonable to continue to use the names given independently to each generation before the connections of the two forms were understood. ”

“ Geoffroy, a contemporary of Linnaeus, made a better distinction (1762) between gall makers and parasites, altho calling the parasites Cynips and the gall makers Diplolepis. During the next century several attempts were made to fix the type of Cynips, but the only designation acceptable under the present International Rules was made by Westwood, in 1840, who named folii, probably the best known of all European gall wasps, as the type of the genus. Detailed discussion of this designation is given in the introduction to the European subgenus in this study. ”

“ In another paper, on the phylogeny of the Cynipidæ, I am discussing this question in more detail and offering data which may be used to draw lines for natural genera. I hope to be able in the near future to revise the genera for the family. Until that time, it is surely wise not to maintain, in the description of new species, the several “genera” known as Cynips, Dryophanta, Callirhytis, Andricus, Biorhiza, Philonix, et al., species of which groups might all be described under the designation of "Andricus." ”

“ Finally, the limits of variation of any character prove to be strikingly uniform thruout the great populations which we propose to call species. Whenever we have taken a reasonably large sample from any point over the usually considerable range of a species, the biometric data have not proved fundamentally different from data for any other fair sample from any other point in the range. The case of Cynips erinacei will serve to illustrate our experience. ”

“ I have got a notice of your views about the female Cynips inserted in the "Natural History Review" (183/3. "Nat. Hist. Review," January 1865, page 139. A notice by "J.L." (probably Lord Avebury) on Walsh's paper "On Dimorphism in the Hymenopterous Genus Cynips," in the "Proc. Entomolog. Soc. of Philadelphia," March, 1864.) : whether the notice will be favourable, I do not know, but anyhow it will call attention to your views...
As you allude in your paper to the believers in change of species, you will be glad to hear that very many of the very best men are coming round in Germany. ”

“ Wings always 1.50 times the body length, cells always clear (except fora trace of a blotch at base of cubitus) ; hypopygial spine rather broad, rather drawn out ventrally. Entirely Old World species.Subgenus Cynips, 4 Wings from 0.14 to 1.60 times the body length, wing cells clear or spotted, spines various. Entirely American species.2 2. Hypopygial spine rather broad, but drawn out ventrally (see figs. 175-190) ; wings normally 1.60 times the body length, or much reduced in length; with a heavy patch and sometimes spots in cubital cell. ”

“ The gall is a small, thin, and flattened disc bearing no resemblance to any Cynips gall except that of guadaloupensis. The agamic insects emerge early in the spring, several months after most Cynips.
Dubiosa Fullaway, 1911, Ann. Ent. Soc. Amer. 4:339. Diplolepis in orig. publ. and later authors. I saw the Stanford types some years ago, and now have large series of collected material. ”

“ In Acraspis all five layers are developed among the species centering about Cynips mellea, but in the other specific stocks in Acraspis the protective and parenchyma zones seem absent and the collenchyma and epidermal layers are unusually well developed.
The gall-producing stimulus, whatever its origin, is evidently selective in its effects upon particular plant tissues; and, since the gall characters are usually of subgeneric significance, it is apparent that these peculiarities of the gall-producing stimuli are of as ancient standing as any of the morphologic structures of the insects. ”

“ It is to an increased knowledge of the physicochemical nature of protoplasm and of the gene, and to such experimental work as that of Muller and others on mutations affected by the introduction of measurable amounts of energy, that we must look for the explanation of the first step in the process of evolution. But granting that mutations do on occasion occur, we may present a body of new data to show that these laboratory mutants are precisely the materials which have differentiated many of the species of our genus Cynips. ”

“ Dr. B. J. Vos, of the Department of German at Indiana University, who has verified the German translations published in this study.
Dr. R. H. Coon, of the Department of Latin at Indiana University, who has verified the Latin translations published in this study.
Dr. Paul Weatherwax, of the Department of Botany at Indiana University, who has helped in interpretations of some botanical data bearing on this genus.
Dr. C. L. Malott and Dr. J. W. Beede, of the Department of Geology at Indiana University, who have criticized the geologic data used in the phylogenetic interpretations of Cynips. ”

“ A more detailed genetic interpretation of our material must for the most part be postponed for a later paper; but we may point out that the graded series of individuals, obtained when short-winged species hybridize with long-winged stocks as with Cynips bifurca and several species in other genera on which we shall publish later, indicate that wing characters in these insects may be dependent on multiple factors or perhaps on more than one group of such factors. ”

“ The Gall Wasp Genus Cynips
PART I. THE ORIGIN OF SPECIES Among the diverse aspects of the so-called species problem, there is none that has received more unsatisfactory treatment than the study of species. The data of morphology and paleontology have given us information on the origin of orders and classes and phyla, and the geneticists and cytologists have contributed a brilliant interpretation of the mechanism which accounts for the similarities and, ipso facto, for the differences between individuals of successive generations ”

“ We may allow details of relationships to be worked out by other lines of evidence, but we should attach some importance to the very primitive character of all the galls of Aulacidea (without a single exception) , to the less primitive but still simple qualities of the galls of Aylax, Diastrophus, and Neuroterus, and to the highly developed characters of the galls of Cynips (European sense) , of Disholcaspis, and of Amphibolips. ”

“ Items of general biologic interest, and distributional data listed by states and under each species of Quercus, are included in this index. Names assigned to Cynips and the principal page references are in bold face; synonyms in Cynips are in italics. ”

“ In many cases we have bred the same species for several winters, and in the case of Cynips fulvicollis (detailed later) we have had to keep transplanted material for two and three years outside our laboratory windows before the insects matured. Nevertheless, in all of this work, we have secured emergence at dates that would have been normal in the native habitats of the species. The season of emergence has been shown to be a specific quality which is not dependent upon the responses of the individual insects to immediate environmental conditions. ”

“ Galls are produced by insects of various orders, but the greater number by species of Cynips. It is impossible to read M. Lacaze-Duthiers’ discussion and doubt that the poisonous secretion of the insect causes the growth of the gall; and every one knows how virulent is the poison secreted by wasps and bees, which belong to the same group with Cynips. Galls grow with extraordinary rapidity, and it is said that they attain their full size in a few days ”

“ It is the familiar story of evolution being conceivable as a function of the remote past, but unacceptable as a matter of moment in the affairs of the present. It is in essence implied that some great cataclysm once upon a time wrought one short-winged cynipid from which all the others have inherited, directly or thru more devious generic paths, all of their unusual characteristics. ”