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Book Reviews Journal of the Lepidopterists' SOCiety 38(4), 1984, 324-327 BOOK REVIEWS Systematische Untersuchungen am Pieris napi-bryoniae-Komplex (Lepidoptera: Pieri­ dae), by Ulf Eitschberger. 1983. Herbipoiiana 1(1), 504 pp.; 1(2), 601 pp. Published by the author and Hartmut Steiniger. Available from the author at Humboldtstrasse 13, D-8671 Marktleuthen, West Germany, and from entomological book dealers. (Due to fluctuations of the Mark against the Dollar, current prices are not available. Early in 1984 the price was 360.- DM + 15.- DM international postage.) The biosystematics of the Pieris napi group remains one of the great intractable problems in the Holarctic butterflies. This is despite the massive and valiant revisionary effort represented by this lavishly-produced monograph, which has truly been a labor of love for its author. Systematische Untersuchungen ("Systematic Investigations"), hereafter referred to as S.U., brings together in one place more morphological and distributional data on the napi group than have ever been assembled before. Eitschberger did an incredible amount of finely detailed morphological work, which is reflected in extraordinary series of pho­ tographs (optical and SEM) and drawings of characters of both adults and immatures. The entire second volume is made up of illustrations, among them 218 color plates averaging over 30 specimens/plate. The photographs are meticulously produced and the colors by and large very true. Within each taxon a range of variability is usually repre­ sented, including seasonal forms and sexual differences; for napi and bryoniae numerous aberrations, rare genetic morphs, and sexual mosaics are also presented. The same spec­ imens are usually shown in upper and lower surfaces on the same plates. For some reason the ventral surfaces are printed slightly smaller than the corresponding dorsals, which is confusing. There is no back-referencing system from the plates to the text, and the forward-referencing system is somewhat clumsy. The first volume of S. U. contains all the text, plus numerous distributional maps. Except for long quotes from the primary literature, which are reproduced from the originals by photo-offset and are thus in their original languages, the text is in German and will not be easy going for readers unskilled in that language. (The most important previous work on the group, the monograph on napi and bryoniae by Muller and Kautz, is also in German and is even more strenuous reading. Moral: If you want to work on the napi group, learn German.) Volume 1 is divided into a fairly brief overview of previous taxonomic work in the group and of the morphological characters deemed to be of value in such work, and a very lengthy taxon-by-taxon treatment which does include "biological," live-bug infor­ mation when available. Twenty-five species are recognized, with a total of 48 subspecies in addition to the nominate ones. The species are grouped into four sets: a Eurasiatic complex of 11 species, including true napi and bryoniae; a North American group of six (to be discussed below); and two Asiatic groups of four species each. There is a tabular summary of character states for the taxa of the first group (pp. 46-51). Even a casual inspection of volume 1 reveals a number of potential problems. (1) Geographic coverage is extremely uneven. This is presumably no fault of the author, who in fact has been remarkably successful in assembling material from odd places. But as one might expect, distributions are mapped in almost infinitesimal detail in western Europe (diminishing rapidly to the east!), moderate (and to this reviewer, rather unsat­ isfactory) detail in North America, and poorly indeed in Asia-where, except for Japan, most taxa are represented by a handful of widely separated, random-looking dots on the map. The inevitable result is that taxonomy is much coarser in some areas than in others. (2) The author is not an ideologue, and does not attempt to force the taxa into the formalisms of cladistics or the quantitative definitional modes of phenetics. He is, how­ ever, apparently not much of an evolutionist or biogeographer either, and he has an old­ fashioned, implicitly typological and explicitly morphological species concept. His work thus most resembles the alpha-taxonomy done on poorly-known groups of bark beetles from Java, and is not at all like what one has come to hope for in the Holarctic butterflies VOLUME 38, NUMBER 4 325 in these sophisticated times. (3) The naming of new taxa has been promiscuous and based on the sort of species concept just described. Many of the new taxa are unlikely to sit well with regional specialists, and many are apt to be ignored or to be treated as junior synonyms of more familiar names, at least until more information about the biology of the animals is available. All these points are relevant to the handling of the Nearctic fauna. Eitschberger recognizes 18 taxa in the Nearctic, of which nine, or 50%, are new. They are (* = Eitschberger name): Pieris venosa venosa; P. oleracea oleracea; P. o. ekisi*; P. marginalis marginalis; P. m. reicheli*; P. m. pallidissima; P. m. mcdunnoughi; P. m. mogollon; P. m. hulda; P. m. meckyae*; P. m. guppyi*; P. m. tremblayi*; P. m. shapiroi*; P. m. brownt*; P. acadica acadica; P. angelika angelika*; P. virginiensis virginiensis; P. v. hyatti*. He is not certain that all the marginalis subspecies are conspecific. There are also brief discussions of several additional marginalis populations he is unwilling to name for lack of good series. Most of the new taxa occur in northwestern North America, from Alaska to British Columbia. (P. angelika, named for Eitschberger's wife, was actually described in 1981 in a paper in the German journal Atalanta, which Eitschberger edits. It is generally unheard of in North American lepidopterological circles. The other new taxa are named and described in S.U. itself. Angelika is described at the species level for reasons which are not terribly clear. It is mostly allopatric with the various marginalis­ taxa, but there is a suggestion of sympatry in a few places. Aside from the co-occurrence of oleracea and virginiensis in a few localities in the northeastern U.S. and perhaps adjacent Canada, this would be the only instance in which it is alleged that members of the napi complex occur sympatrically in the Nearctic.) I have accused Eitschberger of being typological, and I should qualify this by saying that a summary of character-state distributions-raw data only-for selected wing char­ acters is given for most taxa based on the series he examined; and, as noted already, the illustrations portray a good range of variation. Nonetheless, one is left unsatisfied as to the criteria used to recognize and rank taxa; basically, we are being asked to trust the author's judgment. I have discussed this with Eitschberger with specific reference to the northwestern Nearctic taxa, and it is quite plain to me that his weighting criteria are perfectly clear to him. But they are not to me. In fact, I do not consider my own patronymic, shaptrot-which I have never seen alive; oddly, I have apparently worked on the population Eitschberger named angelika-to be well-defined and find it a good candidate for sinking. (I won't miss it.) North Americans tend to bristle at the idea of Europeans working on their fauna from a distance; after all, we are no longer colonials. Such jingoistic reactions should play no role in how we evaluate Eitschberger's treatment of the Nearctic napi. Most Nearctic workers who know our taxa by experience will, however, be properly suspicious of his weighting and grouping. Northern California workers, for example, know that a very complex situation exists in that region in which venosa, marginalis, and pallidissima are all involved; there is no hint of that here. Entities which are considered allospecific by Eitschberger mayor may not be interbreeding in such zones. Such information must ultimately override inferences from morphology. Eitschberger's logical structure would fall apart if interbreeding cuts across his morphological criteria for species status. But does it? The pitfalls, not only of Eitschberger's methods but of their application to this partic­ ular group, are shown by his contribution to the seemingly endless European napi­ bryoniae problem. These two taxa are to European butterfly work what Colias philodice and eurytheme are in North America. Are they one species, or two, or something some­ how inbetween? Given that they appear to interbreed in some places but not others, Z. Lorkovic proposed that they be treated as "semispecies," species in statu nascendi. But "semispecies" is not a taxonomic ranking, and one must decide what to call them. After much soliloquizing, Eitschberger opts to treat them as species and indeed to give bryoniae fifteen (!) subspecies of its own-extending the sense of the name to a large number of poorly-known, hitherto obscure, and very interesting Asiatic populations. Meanwhile, at Bern, Switzerland, Hansjiirg Geiger (1978, Entomol. Zeitschrift 88:229- 235; 1981, J. Res. Lepid. 19:181-195; 1985, Experientia, 41:24-29) has shown that e1ec- 326 JOURNAL OF THE LEPIDOPTERISTS' SOCIETY trophoretically European napi and bryoniae are virtually identical (within-taxon variance sometimes exceeds between-taxon variance). This of course does not prove conspecificity (see below). It is entirely consistent with the hypothesis that the napi-bryoniae distinction is very recent (Holocene) as compared to many other taxic distinctions in Pieris, and it seems inconsistent with the schema developed by Eitschberger. Speaking from the gut, I am willing to bet that electrophoretic data will show Eitschberger's bryoniae-concept to be grossly polyphyletic. Again, time will tell. Why is all this so unsatisfactory? Part of the problem is that Eitschberger's roots are in the German morphological tradition and not in Darwinism, so that we are not all speaking the same theoretical language.
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