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Volume 45, Number 3
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more of the population, it may have more error (e.g., counting individuals multiple times). When the sampling time is limited, the method of Dowes is recommended because it is simpler and its results are similar to those of the "sides" and Pollard methods. Future work should (1) test the effect of non-independence of methods applied simultaneously and (2) use a population of known size to evaluate the accuracy of each method.
We thank T. C. Emmel, J. Feltwell, P. Hanson, L. F. Jiron, and one anonymous reviewer for comments on the manuscript.
Vanessa Nielsen, Escuela de Biologia, Universidad de Costa Rica, San Jose, Costa Rica, and Julian Monge-Najera, Museo de Zoologia, Universidad de Costa Rica, San Jose, Costa Rica.
Received for publication 18 March 1991; revised and accepted 28 June 1991.
BOOK REVIEW
Journal of the Lepidopterists' Society 45(3), 1991, 243-244
Primitive Ghost Moths. Morphology and Taxonomy of the Australian Genus Fraus Walker (Lepidoptera: Hepialidaes. lat.) (Monographs on Australian Lepidoptera, Volume 1), by Ebbe S. Nielsen and Niels P. Kristensen. 1989. CSIRO, East Melbourne, Victoria, Australia. Distributed by Apollo Books, Lundbyvej 36, DK-5700 Svendborg, Denmark. xii + 206 pp. with 435 figures. Hard cover, 17.5 x 25.5 cm, ISBN-0-643-04999-1; DKK 378 (about. $57 U.S.)
The Hepialidae are phylogenetically and biologically distinct outliers that sit near the very base of the evolutionary tree that has at its tips some 150,000 species of butterflies and moths. Hepialids are arguably the most interesting and popular of the ancient lep-idopteran lineages, being noteworthy for their diversity, often beautiful coloration, and comparatively enormous size (at least one of these "microleps" has a wingspan that may exceed 250 mm!). Their peculiar flight and courtship habits—from which their common name the "ghost moths" obtains—also have garnered them a great deal of attention from a range of naturalists. Further, they stand as biological record holders in being among the most fecund non-social herbivores as well as candidates for the most polyphagous (even omnivorous!) Lepidoptera.
Primitive Ghost Moths is the first volume in a new monographic series on the Australian Lepidoptera recently initiated by E. S. Nielsen. In this work Nielsen and Kristensen revise what is believed to be the most primitive hepialid genus, Fraus Walker (hence the book's title), a genus of 25 species endemic to Australia and Tasmania. The authors review thoroughly the morphology and taxonomy of the genus and survey all available biological data.
The first 100 pages are given to a morphological review that is exceptionally detailed and will be of use to all entomologists concerned with lepidopteran anatomy. Emphasis is placed on the skeletal structure, visceral anatomy, and musculature of the head, thoracoabdominal articulation, and genitalia segments. This first chapter is generously supplied with 222 line drawings and photomicrographs. The tissue sections are very clearly stained. Even more impressive are the electron scanning micrographs of sections that have secondarily had the embedding material (paraplast) dissolved away. The numerous scanning electron micrographs of the external anatomy are used effectively throughout. The first 85 pages treat the structure of adults, with the remainder given to the egg, larval, and pupal stages. One cannot help but be struck by the fact that there is perhaps more information on moth morphology here than in John L. Eaton's mistitled book Lepidop-
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Journal of the Lepidopterists' Society
teran Anatomy (1988, John Wiley & Sons, New York, 257 pp.; reviewed in J. Lepid Soc. 43:338-339)—a work limited to the morphology of Manduca sexta.
The larval and pupal descriptions are based on examination of a single species, Fraus simulans, an occasional pest of turf in southeastern Australia and Tasmania. Although the larval SEM's are rather dirty (perhaps an effort might have been made to collect fresher material or to sonicate more thoroughly the material used), there is more detail in these figures than in any previous publication on the immature stages of this family. Figures 216 and 217 showing the branched microtrichea in the larval spiracles are especially striking. The authors claim that the larval antenna is two-segmented, yet the normal condition within the Lepidoptera is for the antennae to be interpreted as being three-segmented. No SEM's of the pupal stage are given, although detailed line drawings are presented. One comment made by the authors with which I fully concur is that the pupal stage promises to be a rich (and presently underexplored) source of characters for defining phylogenetic interrelationships within the family.
One of the most significant findings of the anatomical studies is a reinterpretation of the tegumen. Rather than being derived from dorsum IX as in other Lepidoptera, the authors make the case that the hepialoid "tegumen" is a derivative of the lateroventral wall of segment X. In addition, Nielsen and Kristsensen note that the ventral nerve cord of Fraus is strikingly different from the condition reported for other (higher) Hepialidae. The second chapter on biology is short and adds little to previously published literature on the genus. Much of the discussion is taken from a paper by R. J. Hardy (1973, The biology of Fraus simulans Walker (Lepidoptera: Hepialidae), J. Aust. Entomol. Soc. 30: 113-120). Chapter 3 on diversity and distribution is also brief, mostly given to listings of species found in various regions and habitats about Australia and to ten photographs of Fraus localities in Australia and Tasmania. There is little discussion or synthesis likely to be of interest to biogeographers working outside Australia. The fourth chapter examines the phylogenetic position of Fraus within the Exoporia and the interspecific relationships within the genus.
The final chapter is a thorough taxonomic revision of the 25 members of the genus. Seventeen new species are described; keys to both males and females are provided, along with black and white photomicrographs of the adult male and female, male genitalia, and female genitalia and subanal plate for each species. The photographs of adults appear a little overexposed and might have benefited from a higher contrast printing, whereas those of the genitalic structures are outstanding and provide for ready interpretation. Label data and distribution maps are also given.
Technical and production aspects of the book are very good. The print is clear and figures have reproduced well onto the semi-gloss white paper. The work is handsomely supplied with 435 figures, all of which are clearly captioned or labeled, almost none of which are redundant. The text is generally well written, but I found many of the descriptive sections tedious, especially passages where eight or more sentences in a paragraph began with "The . . ." (e.g., p. 21). I ran across very few typos or editorial problems. The cardboard used for the cover is prone to warpage, such as in both my copies.
Even after a quick glance through this monograph, there will be little doubt as to why Nielsen and Kristsensen are so highly regarded. They have set a new standard for those preparing revisionary works. Although the book's audience will certainly be limited because of its focus on a single rather obscure and uncolorful genus of Australian moths, the detail provided in the morphological section will make the book a must for all interested in anatomy of Lepidoptera or in the evolution of the order. And because Fraus represents a basal lepidopteran lineage, those studying ordinal relationships among insects may also want to own a copy.
With Nielsen as the editor-in-chief of Monographs on Australian Lepidoptera, all can expect subsequent contributions to be outstanding. I am already looking forward to forthcoming publications in this series.
David L. Wagner, Ecology and Evolutionary Biology, U-Box 43, University of Connecticut, Storrs, Connecticut 06269-3042.
Date of Issue (Vol. 45, No. 3): 5 December 1991