The text below is grayed out because it is not intended to be read. It is a necessarily imperfect OCR of the original and is only used by a search engine.
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. Herbipoliana 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 photographs (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 represented, including seasonal forms and sexual differences; for napi and bryoniae numerous aberrations, rare genetic morphs, and sexual mosaics are also presented. The same specimens 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 Miiller 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 information 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 unsatisfactory) 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, however, 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. browni*; 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 characters 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, shapiroi—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 may or 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 particular 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 somehow 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 elec-
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. But that is only part of the problem; the other part derives from the animals themselves. The taxonomic characters in the napi-bryoniae group (if it is a monophyletic group) are mostly wing-color and -pattern things—things we know are commonly determined by a handful of loci. The correspondence between these characters and reproductive isolation is not good, as Lorkovic has shown us with his sibling species balcana. Most re visionary work in Lepidoptera involves morphological characters, especially genetalic ones. But in this group these characters are close to worthless. The differences are so slight—as the hundreds of drawings in vol. 2 show us— that Eitschberger is forced to seize on trivia and to weight them heavily in order to be able to define taxa at all. The results are, not unsurprisingly, not very satisfactory when we compare the napi complex to other groups in which nature has been kinder to the taxonomist.
And that is not all; such differences as exist are often overshadowed by phenotypic plasticity in the napi group. Most of the taxa are polyphenic; seasonal phenotypic differences within taxa are normally greater than differences in the corresponding seasonal brood between taxa; and pupal morphology varies depending on whether or not the individual is in diapause. All of this spells a mess of overwhelming proportions.
I maintain that in many cases, only reproductive-compatibility and/or electrophoretic data will permit proper species assignments in this group. Yet we can now see that these two types of data may conflict with each other! Geiger finds that, electrophoretically, balcana is no more different from napi than bryoniae is—but balcana and napi are highly intersterile (Lorkovic) while bryoniae and napi are more or less highly interfertile, with some exceptions. There is a hint in all of this of an infective or transposable genetic element responsible for sterility, something akin to the "hybrid dysgenesis" factors in Drosophila. If that should be the case, it kicks all conventional species concepts into a cocked hat.
Whatever the basis of sterility in hybrid crosses, it is certain that character reversals, parallelisms, and the like are lurking everywhere in the napi group. The European group of taxa clustering electrophoretically with napi, and probably the Nearctic ones grouped in marginalis and angelika by Eitschberger, seem to represent rapidly-evolving complexes consequent on Pleistocene and post-Pleistocene events. In northwestern North America there is reason to think one is dealing with two, perhaps three, invasions across Beringia—the oldest perhaps Tertiary, the youngest very recent. Preliminary electrophoretic data from Geiger, not available to Eitschberger, roughly support the latter's gross clustering of Nearctic taxa.
No grand synthesis is possible without genetics or without historical-biogeographical analysis or without cladistic reasoning (if not cladistic formalisms). If and when all this is done—and we intend to try it for the northwestern Nearctic—I suspect Eitschberger's judgments will be shown better than might have been forecast. But, I do not intend to start using his nomenclature until then.
Lionel Higgins has given this book a favorable review in the British journal Entomologist's Gazette (1984, 35:174-175). I cannot be so sanguine. (Higgins, by the way, seems to have gotten lost in this Teutonic tome; he asks what the structures shown in vol. 2, pp. 317-321, might be. They are the distal ends of the tongue-cases and antennae of pupae, as should be self-evident to anyone who has seen pierine pupae but at any rate is explained in vol. 1, p. 15.) Eitschberger, who is by profession a pharmacist, has gone to incredible trouble and almost unimaginable personal expense to provide us with this huge work. As I noted at the beginning, it is the biggest compilation of data on these
Volume 38, Number 4
327
animals ever assembled, and therein lies its primary value. The data cry out for other sorts of interpretation than Eitschberger has given them. Anyone interested in this most exasperating of groups, and who reads German, must have access to this book. If you are not willing to buy it, have your institutional library do so. Otherwise, it will become another Miiller and Kautz. Possibly the only academic library copy of Muller and Kautz in the United States is at Yale, which prohibits photocopying of interlibrary loan materials. The only way to get hold of the book is to go to New Haven or to buy one through an antiquarian. Will S.U. disappear in similar fashion?
Eitschberger told me over a stein of beer that he hopes other people will take up and expand his work. That is good, for it must—and will—be done. Systematische Unter-suchungen . . . could be read superficially (and apparently was, by Higgins) as the definitive resolution of the napi problem. It isn't. It is a beginning.
Arthur M. Shapiro, Department of Zoology, University of California, Davis, California 95616.
Journal of the Lepidopterists' Society 38(4), 1984, 327-328
Dear Lord Rothschild (Birds, Butterflies & History). Miriam Rothschild. 1983. Hutchinson Publishing Group, 17-21 Conway St., London, W1P5HL. Format 7" x 9%" 398 pp., including index & appendices. 90 pp. of B/W photographs. 12 pp. of color plates. Cloth bound. 14.95 (British pounds.)
One usually thinks of Lord Rothschild in connection with Karl Jordan or Ernst Hartert, both of whom were among his co-authors. In contrast, few people know the history of the Tring Museum, nor the other aspects of his life which took place beyond the boundaries of Tring. This book, written by his niece, is a revelation. It is a story of one life, liberally embellished with ancestors and heavily endowed with wealth. It is the story of the workings of Parliament, the education of the young, of action on the high seas, and of wild creatures, both alive and dead, which previously had never been known to the world.
"It is not easy to be born. The average man is squeezed out into the world with blood to lubricate his passage and wild shrieks of anguish to speed him on his way."
So begins the biography of Lionel Walter Rothschild, 2nd Baron of Tring.
At the age of seven, at tea time in the nursery, Walter suddenly stood and made the following announcement: "Mama, Papa, I am going to make a museum, and Mr. Minal is going to help me look after it." This prophesy came true.
Walter was the classic example of a child who shows little scholastic promise, but at some point becomes fired with enthusiasm in one particular field of endeavor to the extent that he becomes expert in that field to the exclusion of all else. A psychologist might have altered this, if such a person had existed in England at that time. He was born to a mother who was strict and sensorious on one hand—overprotective and indulgent on the other. His father was never able to understand either his love for animals or his failure in finances. From the beginning he had a speech defect which resulted in crippling shyness. He was tutored at home and rarely played with boys of his own age. All of this, added to the astronomical wealth of his family, contributed to his enigmatic personality.
He began his collections at the age of seven with one butterfly. By the time he was 19 he had collected 5000 birds (2000 of which he had already mounted) and 38,000 Lepi-doptera. Two years later, his family built him a museum as a 21st birthday present.
This book mirrors the life-long curiosity of one man to discover and collect all the