The Australian Olethreutine tortricids

OLETHREUTINE MOTHS OF AUSTRALIA (LEPI-

DOPTERA: TORTRICIDAE). By Marianne Horak,with contributions from Furumi Komai. Monographs onAustralian Lepidoptera, Volume 10. CSIROPublishing. 2006.

522 pp. Hardback. ISBN 0-643-09093-2. AU $160.00.

Marianne Horak begins this tenth volume of the Australian

monograph series with the statement that ‘‘most people’sintroduction to olethreutine moths would have been anunpleasant experience,’’ referring to biting into an apple

infested by the codling moth, one of the many pest species inthe family. To the contrary, this is a relished contribution tothe world lepidopterists, especially those working withTortricidae. The long established quality of this Australian

monograph series, initiated by the late I.F.B. Common andlate E.S. Nielsen, is maintained, if not surpassed, by thislatest volume. The Olethreutine Moths of Australia

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provides a ‘‘top-down’’ approach to define 90 genera andprovide assignments of 249 named species, leaving 200þknown new species to future revisions of the defined genera.Twelve new genera and a selected 16 new species aredescribed. A large number of new generic combinations

are provided for Australian and Oriental species. A remark-able 984 figures of imagos, genitalia, wing venation, andother structures of the body are provided, many of which arescanning electron images, and all are of the highest quality

to be found in current publications. The Grapholitini isjointly treated with Furumi Komai (Osaka University ofArts, Japan), the leading specialist on this tribe in the world.

Chapter 1 of this work provides a phylogeny of Australiangenera, which Horak regarded as preliminary and incon-clusive. Yet, this is the first phylogeny of genera ever

proposed for world Olethreutinae based on a cladistic ana-lysis. The phylogeny and subsequent text notably includesGatesclarkeanaDiakonoff and related genera in Olethreutini,rather than in Gatesclarkeanini, and it strongly supports the

distinctiveness of Enarmoniini as distinct tribe based ona suite of new characters. The phylogeny was based on 126characters that were unordered and unweighed and ana-

lyzed with WinClada and NONA. The strength of thephylogeny is weakened by the lack of weighting of indepen-dent reductions of characters, especially wing venation.

Additionally, some characters involving scent scales andassociated structures of the male are not homologous, e.g.,character 14 with modifications of scales on the male fore-

and mid- tibia should be two characters, rather than onecharacter with three states. Regardless of questionablehomologies of reductions and scent scales, the charactermatrix provides a valuable foundation for future cladistic

analyses of the subfamily.Chapter 2 provides a thorough discussion of olethreutine

morphology, one of the most complete and well illustrated

treatments ever presented for the subfamily and complement-ing a previous treatment by Horak (1991). The head andthorax, along with their appendages, are described in detail,

supplemented by images with a scanning electron micro-scope. In addition to illustrations of genitalia of selected taxa,53 figures of wing venation are provided for genera.

Chapter 3 on biology provides a review of larval hostplants and support for externally feeding larvae beingancestral to internal feeders, based on the presence in allsubfamilies of an anal fork for ejection of feces from the

feeding shelter and its absence in internal feeders. In regardsto the latter, the cited reference ‘‘Horak, 1989’’ is a misprint,and this should be Horak (1998). The author should have

noted that the presence/absence of the anal fork and feedinghabit is not always correlated because it is absent in someexternal feeders, e.g., some species of Olethreutes Hubner,

and present in some internal feeders, e.g., someCrocidosemaZeller (MacKay, 1959). Horak documents examples ofrepeated transitions between leaf rolling and internal feederswithin a tribe or genus. The information on host plants of

Australian Olethreutinae is understated in this chapter asnumerous new host records for species and genera areprovided in Chapter 5.

Chapter 4 presents an analysis of the diversity anddistribution of the Australian fauna relative to faunas in

other geographic regions. The number of genera in Australia(90) exceeds that of Europe (64) and the Nearctic Region(59). This chapter illustrates that the center of diversity of

Australian genera is in the Oriental Region, which has beenpoorly sampled. Diakonoff ’s (1973) treatment of just theOlethreutini in the South Asiatic region included 88 genera –more than twice the number for this tribe than in any other

region of the world. This chapter also correlates the highdiversity of species in the Spilonota-group of genera with theradiation of Myrtaceae in the Indo-Australian Region.

The final chapter 5, the body of this work, includes atreatment of each of the 90 genera with generic synonymies,diagnosis, excellent descriptions of superficial characters,

including wing pattern and venation, and male and femalegenitalia, along with general distributions in Australia andother regions, biology, remarks, and constituent species.Illustrations of heads showing palpi, wing patterns, and

genitalia are provided for one or more species in each genus.These figures are the first to illustrate many genera andspecies that were described by O.B. Lower, E. Meyrick, and

A.J. Turner.The literature cited and appendices of this volume will be

valuable resources for future research on systematics of

Tortricidae. Appendix 1 includes the character matrix forthe phylogenetic analysis. Appendix 2 that lists host plants isunduplicated elsewhere and will provide a reference for

anyone working on Olethreutinae at the world level.This work is very thorough and well written, and

criticisms becoming a matter of ‘‘nit-picking,’’ as indicatedby the following. 1) The history of tortricid systematics in

Australia is not treated, and a short discussion of thecontributions and fieldwork of Alex Diakonoff, A. J.Turner, O. B. Lower, and E. Meyrick relative to Australia

would have have added value. 2) A list of genera andconstituent species is not provided, and this would haveaided retrieval of names. 3) In regards to terminology for

wing patterns in Chapter 2, the author correctly definesstrigulae as pairs of pale marks separated by dark spacers onthe forewing costa throughout Tortricidae. Because pale

strigulae are not conspicuous in species with pale wings, sherefers to the dark spacers as strigulae for ease of description,but this confuses homology in analyses of wing patterns.The Olethreutine Moths of Australia is the culmination of

many years of research by Marianne Horak, and it is anexclamation mark to her previous contributions on tortricidsystematics. Although this work concerns the Australian

fauna, it has the greatest impact on systematics of thesubfamily worldwide since the works by Heinrich (1923,1926) on the Nearctic fauna. The importance of this work

for the Oriental Region is demonstrated by a recent doctoralthesis on Olethreutinae in Thailand (Pinkaew, 2006). Thisthesis included many genera with uncertain tribal associa-tion and a large number of generic morphotypes. The

available this volume by Horak made it possible to assignnames to nine of these generic morphotypes and resolvetribal and generic assignments of many taxa.

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The library of every Lepidoptera systematist and everyuniversity involved in biological and systematic research

should have a complete set of the Monographs on AustralianLepidoptera. But if funds are available for purchase of onlyone volume, Horak’s volume 10 is recommended as the first

choice because of its systematic value, its application toa worldwide fauna, and its treatment of an economicallyimportant group of moths. Marianne Horak’s OlethreutineMoths of Australia will provide the foundation for future

systematic research on Tortricidae.

RICHARD L. BROWN

Mississippi Entomological MuseumMississippi State University

References

Diakonoff, A. (1973) The South Asiatic Olethreutini (Lepidoptera,

Tortricidae). Zoologische Monographieen van het Rijksmuseum

van Natuurlijke Historie, 1, 1–699.

MacKay, M. (1959) Larvae of the North American Olethreutidae

(Lepidoptera). Canadian Entomologist, Supplement 10, 1–338.

Heinrich, C. (1923) Revision of the North American moths of the

subfamily Eucosminae of the family Olethreutidae. Bulletin of the

United States National Museum, 123, 1–298.

Heinrich, C. (1926) Revision of the North American moths of the

subfamilies Laspeyresiinae and Olethreutinae. Bulletin of the

United States National Museum, 132, 1–216.

Horak, M. (1991) Chapter 1.1. Morphology. Tortricoid Pests. Their

Biology, Natural Enemies and Control (ed. By L. van der Geest

and H. Evenhuis). Elsevier, Amsterdam.

Horak, M. (1998) The Tortricoidea. Lepidoptera, Moths and

Butterflies. Volume 1: Evolution, Systematics, and Biogeography.

Handbuch der Zoologie/Handbook of Zoology. Vol. IV Arthropoda:

Insecta Part 35 (ed. N. Kristensen). Walter de Gruyter, Berlin.

Pinkaew, N. (2006) Taxonomy of Olethreutinae (Lepidoptera:

Tortricidae) of Thong Pha Phum National Park, Kanchanaburi

Province, Thailand. Thesis. Graduate School, Kasetsart Univer-

sity, Bangkok.

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