Discover millions of ebooks, audiobooks, and so much more with a free trial

Only $11.99/month after trial. Cancel anytime.

Australian Weevils (Coleoptera: Curculionoidea) I: Anthribidae to Attelabidae: The Primitive Weevils
Australian Weevils (Coleoptera: Curculionoidea) I: Anthribidae to Attelabidae: The Primitive Weevils
Australian Weevils (Coleoptera: Curculionoidea) I: Anthribidae to Attelabidae: The Primitive Weevils
Ebook1,227 pages9 hours

Australian Weevils (Coleoptera: Curculionoidea) I: Anthribidae to Attelabidae: The Primitive Weevils

Rating: 0 out of 5 stars

()

Read preview

About this ebook

In Volume I, the primitive weevil families Anthribidae, Belidae, Nemonychidae, Caridae, Rhynchitidae and Attelabidae are treated. One hundred and two genera and 400 species are catalogued. The species are illustrated by about 1035 individual drawings and black and white photographs, in addition to 650 colour photographs relating to primitive weevil families in Volumes V and VI.

Volume I includes a chapter on Nemonychidae by G Kuschel and also an important Postscript detailing some crucial taxonomic changes in several weevil subfamilies that are only dealt with in detail in the later volumes.

LanguageEnglish
Release dateJan 1, 1994
ISBN9780643105607
Australian Weevils (Coleoptera: Curculionoidea) I: Anthribidae to Attelabidae: The Primitive Weevils

Read more from Ec Zimmerman

Related to Australian Weevils (Coleoptera

Related ebooks

Biology For You

View More

Related articles

Reviews for Australian Weevils (Coleoptera

Rating: 0 out of 5 stars
0 ratings

0 ratings0 reviews

What did you think?

Tap to rate

Review must be at least 10 words

    Book preview

    Australian Weevils (Coleoptera - EC Zimmerman

    AUSTRALIAN WEEVILS

    VOLUME I

    ANTHRIBIDAE TO ATTELABIDAE

    AUSTRALIAN WEEVILS

    AUSTRALIAN WEEVILS

    (Coleoptera : Curculionoidea)

    ELWOOD C. ZIMMERMAN

    BSc (Berkeley), DIC, PhD, DSc (London)

    Curator of Weevils, Emeritus

    Australian National Insect Collection

    CSIRO Division of Entomology

    Canberra, Australia

    Honorary Associate, British Museum

    (Natural History)

    VOLUME I

    ORTHOCERI

    ANTHRIBIDAE TO ATTELABIDAE

    THE PRIMITIVE WEEVILS

    Illustrations © CSIRO Australia 1994

    Text © Elwood C. Zimmerman 1994

    Except for private study or research, review or criticism, no part of this work may be reproduced or distributed by any means or stored in any retrieval system or data base without prior written permission from the publisher, CSIRO Australia.

    Volume I published on 28 February, 1994

    National Library of Australia Cataloguing-in-Publication Entry

    Zimmerman, Elwood C. (Elwood Curtin), 1912 –

    Australian weevils. Volume I.

    Includes indexes.

    ISBN 0 643 05145 7 (v. 1).

    ISBN 0 643 05144 9 (set).

    ISBN 0 643 05146 5 (v. 2).

    ISBN 0 643 05147 3 (v. 3).

    ISBN 0 643 05148 1 (v. 4).

    ISBN 0 643 05149 X (v. 5).

    ISBN 0 643 05150 3 (v. 6).

    ISBN 0 643 05144 0 (v. 7).

    ISBN 0 643 05145 9 (v. 8).

    1. Beetles - Australia. 2. Curculionoidea - Australia.

    I. CSIRO. II. Title.

    595.7680994

    Cover Illustration

    Curculio fraudator Zimmerman

    Drawn by Se Kim

    CSIRO Division of Entomology, Canberra

    End Papers

    Agriochaeta crinita Pascoe

    Essolithna villosa Lea

    Drawn by Sybil Monteith, Brisbane

    This Series is available from:

    CSIRO Information Services

    PO Box 89 (314 Albert Street)

    East Melbourne, Victoria 3002

    Australia

    Telephone: (03) 418 7217

    Fax: (03) 419 0459

    Entomological Society of America

    9301 Annapolis Road

    Lanham, Maryland 20706

    USA

    Telephone: (301) 731 4535

    Fax: (301) 731 4538

    Set in 10/12 Stone Serif using a Macintosh Powerbook 180 and Quark Xpress 3.1

    Design and production by CSIRO Information Services, Melbourne

    Printed in Melbourne by Brown Prior Anderson

    IN GRATITUDE

    THIS SERIES OF VOLUMES IS DEDICATED TO

    Douglas F. Waterhouse A.O., C.M.G., F.R.S., F.A.A., D.Sc.

    Chief, Division of Entomology, CSIRO, 1960 – 1981

    under whose direction and encouragement the project was initiated and from whose wise counsel I have profited.

    Whoever as an entomologist looks into the future knows full well that we are steering into a shoreless sea, no matter whether he estimates the total number of insect species at three, ten or fifteen millions. In the near future any beginner will be greyheaded before he has caught up with what is already known.

    Walther Horn

    If any worker believes that he can complete most of an alpha study of the Australian weevil fauna in a full, long lifetime of concentrated labour, he is suffering a delusion. If any institutional administration is convinced that the task can be completed in a generation or two or three, or four, it is being deceived.

    E.C.Z.

    TABLE OF CONTENTS

    Preface

    Acknowledgements

    Catalogue of the Australian Curculionoidea Illustrated and Annotated

    Conspectus of the Families and Genera Treated in this Volume

    Checklist of the Species Listed in this Volume

    Anthribidae

    Anthribinae

    Choraginae

    Belidae

    Pachyurinae

    Belinae

    Nemonychidae

    Caridae

    Rhynchitidae

    Attelabidae

    Appendix on Nemonychidae by G. Kuschel

    Mecomacerini

    Rhinorhynchini

    Postscript

    Index

    Those who know most about this subject

    know best how little they know.

    PREFACE

    One cannot understand how difficult it is to prepare

    a work of this kind unless one has prepared a work of this kind.

    With the issue of this book, five of the scheduled eight volumes of the Australian Weevils monograph have been published: Volume II, Brentidae, Eurhynchidae, Apionidae and Immature Stages, 1994; Volume III, Rhynchophoridae, Erirhinidae, Curculionidae: Amycterinae, Literature Consulted, 1993; Volume V, colour plates 1–304, 1991; Volume VI, colour plates 305–632, 1992; Volume IV, Curculionidae: Adelognatha, is scheduled for publication in 1995.

    The area covered by this treatise is continental Australia and its adjacent islands and the Pacific island possessions of Lord Howe and Norfolk which share many Australian genera. It does not include the distant subantarctic islands of Macquarie or Heard or the Indian Ocean island possessions of Cocos or Christmas whose faunas are Indomalayan.

    Near this position in a book it is customary to include an Introduction, but to decide what may come first is the last thing that one can do. In a large work of this kind, parts of which remain to be published, it is better to prepare a Postscript after one’s work is done and there has been time for reflection upon what has been written. Hence, the equivalent to an Introduction to this monograph is planned for inclusion in Volume VIII, the end of the series, but in this Preface I have included details that could be part of an Introduction.

    In this Volume I the primitive weevil families Anthribidae, Belidae, Nemonychidae, Caridae, Rhynchitidae and Attelabidae are treated. One hundred and two genera and 400 species are catalogued. The species are illustrated by about 1035 individual drawings and black and white photographs, in addition to 650 photographs in colour which have already been published in Volumes V and VI. The annotated bibliography is included in Volume III. The word weevil is a derivative of the old Anglo-Saxon wifel, a beetle.

    This work is not a definitive monograph; it is a base upon which to build. It is a catalogue embellished with illustrations of almost every described Australian weevil species, annotated with notes, keys and commentary prepared as time and opportunity permitted. The goals that I had set for the text have not all been attained, and, partly because of my advanced age, some work had to be terminated in mid stream. I am the one most aware of the faults and failures.

    Resources were insufficient to allow for laboratory assistance to carry part of the heavy loads of routine and preparatory work. This has delayed or prevented completion of many parts of the text. Moreover, it has been my sincere wish that it would be possible to arrange for a dedicated weevil specialist to replace me on the staff of the Australian National Insect Collection to continue my work, to inherit my material, my extensive library and my manuscripts and to continue to advance Australian weevil research after I am gone. That great hope of my life has not eventuated. Consequently, this monograph, my work and well-being have suffered severely. If we are serious about the urgency of biodiversity studies in the light of catastrophic extinction and the Nation-wide need of well-illustrated catalogues and identification manuals, it is my hope that it will be possible to procure the resources essential to support continuing research on Australian weevils, our largest family-group of living organisms.

    I have been a student of weevils for a long time. I recently chanced upon some specimens in the South Australian Museum that I had collected in California in 1931 and sent to the Museum in exchange. That was the year when I wrote my first descriptive paper on weevils. My entrance upon Pacific studies began in 1930, when, as a grammar school boy, I was employed at 25¢ per hour as a laboratory technician preparing specimens collected in the Marquesas and Society Islands by the Pacific Entomological Survey.

    In 1934, upon my return from the never-to-be-forgotten Mangarevan Expedition sent out by Bishop Museum of Honolulu to explore scientifically many islands of southeastern Polynesia when the area was still relatively unspoiled and primitive (I am the sole survivor of the staff of that expedition, the greatest event of my life; my salary was then $25 per month, food and lodging), I began my study of Australian weevils when I curated the important Richard Helms collection of Australian Coleoptera that had been purchased by Bishop Museum. Thenceforth, my Australian studies continued and expanded in association with my research on the weevils of Pacific islands, especially in preparing reports upon the collections I made while exploring on more than 50 islands from the Tuamotu Archipelago and the Pitcairn Islands in the east to Fiji in the west. For that work it was necessary to examine Australian specimens and literature. My first studies of Australian material were published in 1936. Before the end of the Second World War, I had prepared a detailed catalogue of the Australian weevils. During 1948, the first five volumes of my Insects of Hawaii monograph were published. Between 1949 and 1973,I spent much time at the Natural History Museum in London preparing five more volumes of Insects of Hawaii and studying weevils with the late Sir Guy Marshall (see the biographical notes in the bibliography in Volume III of this series), and I visited collections and specialists at Tring, Oxford and Cambridge and elsewhere in England, and Sweden, Denmark, Holland, Belgium, France and Italy on the Continent. In America I spent several months at the National Museum at Washington and later, over a period of several years, I enjoyed access to the collections, rich library facilities and stimulating staff of the Museum of Comparative Zoology at Harvard University.

    In England in 1972, Douglas Waterhouse, then Chief of the Division of Entomology, CSIRO, with strong support from E.B. Britton, invited me to come to Canberra to prepare this monograph. At that time we did not anticipate that the scope of the work would expand as it has or that more than 20 years of concentrated labour would be required to produce this introducion to the study of the extraordinarily rich and diversified Australian weevil fauna.

    Notes on History

    When the baseline for the modern classification of animals was established by the immortal Linnaeus in 1758, the outside world did not know that Australia existed. Linnaeus knew fewer than 100 species of weevils from the entire world. It was not until Captain Cook’s discovery and exploration of the east coast of Australia between April and August 1770, and the return to England of the specimens collected by Joseph Banks and Daniel Solander, naturalists aboard the Endeavour, that the first Australian weevils were made known to the outside world. Some of those specimens were collected in the Sydney area (Botany Bay) of what is now New South Wales, and Bustard Bay, Thristy Sound and Endeavour River (now Cooktown) in what is now Queensland where most specimens were collected during the several weeks that the Endeavour was laid ashore after being holed on a coral reef and almost lost. Most of the species collected were described by Fabricius in 1775, 1781 and 1801. In Volume III of this series I have given biographical notes on Fabricius and have listed the names of the Australian weevils described by him together with their current dispositions. Some of the specimens described by Fabricius were mislabeled and were recorded in error as having come from Australia, two were synonyms, and thus by 1801 only 19 Australian weevils were known to science. Then followed a number of voyages of discovery from several Europen countries, and naturalists aboard such ships as l’Astrolabe (later named La Coquille), Zelee, Novara, Uranie and Beagle collected many weevils that were described by several authors. In 1835, Boisduval added numerous species from the voyage of the Astrolabe. The Schoenherrean monographs of 1823, 1825, 1826 and the great eight volume Genera et Species Curculionidum of 1832–1845 introduced a new era of research on weevils. At the end of that work, Schoenherr listed 237 species then known from Australia, including 10 from Tasmania, but he had overlooked Erichson’s 1842 paper, Beitrag zur Insectenfauna von Vandiemensland, in which eight new genera and 46 new species were added. In the concluding volume of Schoenherr’s monograph in 1845, he listed 7,141 species of weevils as all then known in the world. In 1826 (1827) William Sharp Macleay added a few genera and species from the west coast. Between 1848 and 1865, Boheman, Germar and Jekel added new species to the Australian list.

    The insects collected by Charles Darwin in 1836 in Tasmania (Hobart area, 5–7 February), New South Wales (Sydney to Bathurst, 12–30 January), Albany area of southwestern Australia (6–14 March) appear not to have been prepared expeditiously, and most of the weevils lay unstudied for many years. In view of the favourable impact that the great Schoenherrean monograph made on the scientific world, one would have expected that the specimens collected by Darwin would have been sent to Schoenherr for study. In 1921, Gilbert Arrow sent to Arthur Lea all of the Darwin weevils that could then be found in the British Museum, and in 1926 Lea published a paper entitled On Some Australian Coleoptera Collected by Charles Darwin During the Voyage of the Beagle (Transactions of the Entomological Society of London for 1926, Part II:279 – 288). Lea wrote (page 279) "The British Museum having at various times sent to me for identification specimens taken by Darwin mixed with others, I suggested to Mr Arrow that it appeared desirable to identify all the remaining Australian beetles taken by the great naturalist and deposited in that institution. To this he agreed, and they were sent to Adelaide in 1921 .... The specimens were all small, and ... Mr Arrow wrote:

    Darwin did not give his collection to the Museum, but allowed different individuals to take particular groups which interested them, and the unsorted mass of minute specimens was given to G.R. Waterhouse, only coming here in 1887.

    From this last assemblage of Darwin material, Lea recorded 28 species, including six new to science.

    By 1860, Blanchard, Boisduval, Chevrolat, Donovan, Duponchel, Germar, Guerin-Meneville, Hope, Jekel, Kirby, Olivier, Redtenbacher, Shuchard and Waterhouse had added more than 200 species to the Australian weevil list.

    In 1863 and 1866, that remarkable Belgian coleopterist, Jean Theodore Lacordaire included the then known Australian weevil genera in his monumental Genera des Coléoptères and laid a base to be built on by all following workers.

    The English amateur coleopterist, Francis Polkinghorne Pascoe (see the bibliography in Volume III for biographical notes) described his first Australian weevil in 1859, and from then until 1887 he was very active in describing Indo-Pacific and Australian weevils. He was sent large consignments of Australian specimens, especially by George Masters and F.H. duBoulay (see Volume III, page 717). Pascoe described about 180 genera and 630 species from Australia. He described more Australian weevil genera than any one other author, and his Australian studies extended over a greater number of years (28) than any other worker. Many of the genera and species he described have remained unrecognised in Australian collections (and because of the nature of his descriptions, most are unrecognisable from the literature). I have illustrated specimens of many of his species, but much remains to be done to make them better known to science. His collection is well preserved in the British Museum, and I have studied it.

    In 1865, William John MacLeay published his major work on the Australian ground weevils entitled The Genera and Species of Amycteridae and, although superficial and inadequate, this laid the foundation for the study of the remarkable and multitudinous group of characteristic Australian Curculionidae.

    In 1873, T. Vernon Wollaston published his monograph On the Genera of the Cossonidae in which he described about 20 genera that contain Australian cossonids as well as a small number of Australian species.

    In 1882, the Reverend (later Canon) Thomas Blackburn, of the Church of England, arrived in Australia after several years serving the Church in Hawaii where he was the pioneer coleopterist. It was his work in Hawaii that stimulated David Sharp, then at Cambridge University, to plan and direct to completion the classic Fauna Hawaiiensis exploration and publication. Blackburn was an able and dedicated amateur coleopterist, and in spite of his heavy official church duties, he devoted a large amount of time to the collection and description of Australian Coleoptera. He described more than 3 000 Australian Coleoptera, and that was a remarkable accomplishment. For reasons unknown to me, it was not until six years after he arrived in Australia that Blackburn began to publish on the Australian fauna. He described about 32 genera and 257 species of Australian weevils between 1888 and 1906. Most of his types are in the British Museum, and some are in the Lea-South Australian Museum collection.

    The most active and productive student of Australian weevils was the Australian-born Arthur Mills Lea. He began publishing on Australian Coleoptera in 1894 while Blackburn was still working actively. Lea took great interest in the weevils early in his career, and Blackburn, I believe, cutailed his own studies of weevils in deference to the activities of Lea.

    Blackburn and Lea made extensive use of umbrellas to beat vegetation when they collected. In that way they obtained many weevils, often in large series, that have not, or have rarely been taken since.

    Much of Lea’s work on weevils was not well done, and, because of the many genera and species he described, we have inherited many problems. He used a hand-lens for most of his work, his preparatory techniques were generally not of good quality, and many of his observations have been proved erroneous. His type series often contain more than one species or even specimens of different genera. One type series was found to contain three different species. There are many specimens in collections bearing Lea’s type or other labels that represent manuscript names. Many of the specimens were studied many years before his death, but his manuscripts, if any, were not published. Not infrequently he mistook a protruding ovipositor for an aedeagus and thus confused the sexes. Lea had the bad habit of labeling most of his paratypes with only small labels reading Co-type with no species name. Hence, if such a Co-type becomes separated from the labeled Type (holotype) in the collection, its identity may be lost or confusion result, as frequently has happened. To enable him to see various features better, Lea removed the heads of many specimens, including types, and glued them separately to the cards holding the specimens. Some of them have since been damaged or lost. See further discussion under Cryptorhynchinae in Volume VII.

    Lea began his entomological career as an applied or economic entomologist. It is therefore difficult to understand why he rarely cited the name of a plant from which he collected specimens, he rarely mentioned the dates of his captures, and the localities of many of his collecting sites are imprecise. He rarely cited a type, type data or type locality.

    Lea’s collection was in such a confused state of curation that much work was required to prepare it for convenient use. It will take many years for workers to clear the confusion left by Lea who was seriously infected by mihi-itch. His enthusiasm and dedication were great, and it is a pity that he was not a more careful worker. He named about 5 500 species of Coleoptera, including about 145 genera and approximately 2 200 species of Australasian weevils. Lea described about half of the known species of Australian weevils.

    Lea was for a long time engaged in state governmental service in New South Wales, Western Australia and Tasmania, and he travelled and collected extensively in many parts of the country. No person has collected so many specimens or species of Australian weevils over so much of Australia as did Arthur Lea. In 1911 he joined the South Australian Museum in Adelaide and worked there with absolute dedication for nearly 21 years building that institution’s Coleoptera collection. The weevil collection alone is by far the most complete and valuable collection of Australian weevils. It incorporates most of Lea’s types and his personal collection.

    Eustace William Ferguson, a microbiologist with the Department of Public Health, Sydney, became interested in the Amycterini, and in 1909 he published Part I of his Revision of the Amycteridae. Then followed 14 substantial papers on the group, including Parts II to VIII of his Revision, the last part of which was published in 1923. The Ferguson series cleared many of the faults of the early Macleay studies, and it has remained the basic contribution available on the amycterine weevils. It is, however, an unsatisfactory attempt to monograph the difficult-to-understand group. I have found much of it unusable.

    Following the death of Arthur Lea in 1932, there were only scattered small publications issued on Australian weevils, such as those of Marshall, Uther Baker, Schedl and Zimmerman, until Richard Thompson published his large Revision of the Genus Catasarcus Schonherr in 1968. More than 40 years elapsed without any professional worker being assigned by any Australian institution to study the remarkable Australian weevil fauna. When assistance was required, especially for identification or description of Australian weevils of economic importance, Australian institutions begged help from overseas workers, and the bulk of the requests fell upon the busy shoulders of Sir Guy Marshall of the Imperial, later Commonwealth Institute of Entomology and the British Museum (Natural History). It distresses me greatly that my career is now ending without any provision being made for a successor to continue the study of the extraordinary Australian weevil fauna. How long will it be before an Australian institution will again engage a worthy authority to study systematically this great group of animals, many of which are of major economic importance? In view of the national need of the research, I hope that responsible authorities will recognise the problem and soon find a solution for it.

    Study of this huge complex is exceedingly difficult even for specialists with many years training and experience. For non-specialists, including agricultural and forestry workers, the situation has been almost impossibly difficult. The published taxonomy of the Australian weevil fauna is chaotic. Not only is this fauna naturally so complex as to render its study difficult, but the difficulties have been compounded by the poor quality of much of the previous work done upon it, much of it so poor or inadequate that it would have been better had some of it not been published. Much of my research time since coming to Australia has been devoted to the tedious tasks of clearing away the confusion in literature and preparing a base for advancement. There is also much confusion in Australian collections. Many specimens bear false names. For example, a series of eight specimens named Essolithna enchymys Pascoe in the Lea collection contained five different species, and none is enchymys.

    The preparation of specimens, including those continuing to be submitted today, is mostly so inadequate that one must waste hours cleaning and remounting inadequately prepared specimens before they can be studied. Workers should be ashamed to submit their poorly prepared specimens to overworked specialists and expect prompt, free identifications in return. It is legitimate for systematists to refuse to accept for study inadequately prepared specimens.

    Notes on the Size of the Fauna

    The Australian weevil fauna is one of the world’s largest, most diverse, most remarkable, most distinctive of the major faunas. In this monograph is included information on about 675 or more genera and 4 200 described species, but those numbers may represent only about half the fauna. The number of existing species cannot be estimated with any degree of certainty, but the knowledge that many remarkable new genera and numerous new species, some of them the most distinctive of any yet seen, are already in collections awaiting description, and that collectors have briefly and superficially sampled only small areas of this large continent and that the fauna really remains inadequately collected and poorly known, it is probable that there are here more than 1 000 genera and perhaps 6 000 to 8 000 species.

    The Australian weevil fauna contains more species than all of the Australian mammals, birds, reptiles, fishes and amphibians combined plus various invertebrate groups. There are many authorities working on those groups in Australia.

    Incalculable numbers of precinctive species surely have been exterminated by deforestation, extensive burning, intensive grazing and overgrazing by millions of sheep and cattle and human activities. Extermination continues, in some places at an alarming rate, but not enough is being done to collect and to preserve for study by future generations representative samples of these vanishing marvels of evolution.

    The Australian weevil fauna is larger than that of the Palaearctic and the Nearctic where there are many workers studying weevils. In Canada and the United States of America there are about 2 500 species of weevils in about 375 genera, comparatively small numbers considering the vast area involved, but the last Ice-age impoverished much of north of that area. The only comprehensive monograph (and not illustrated) of the weevils of that region was published more than 100 years ago.

    There are only about 475 species of weevils known in Great Britain, and the entire coleopterous fauna amounts only to about 3 750 species. There are many more species of weevils in Australia than the total number of all beetles in Great Britain. A modern, illustrated monograph of the weevils of Great Britain remains to be written.

    [May I digress from my main theme to consider the question of why it is that such areas as the United States and the United Kingdom, where there are many able systematists, have no modern, well-illustrated manuals to aid in the identification and understanding of their weevils? Why is it that so many of their workers keep rushing off to foreign lands to dabble in exotic places, as interesting and valuable as some of that activity is, and in some instances necessary for monographic work, while the faunas of their own fields and forests are largely ignored by monographers, and they continue for generations without adequate texts being written upon them? Is it because the difficulties of preparing such texts are too great for most workers? Is it that few are those who can accept being kinds of long-term prisoners of such projects? Is it lack of interest, finance or institutional support? Or is it that the insidious mihi-itch is so contageous and widespread? A friend suggests that in some respects Australian Weevils has certain parallels with Biologia Centrali-Americana - both have been privately funded. It is a sad state of affairs when to get major monographs prepared and published individual initiative and personal finance must replace much of that which institutions should accomplish.

    Of course, such work takes time, money, and hard working, dedicated staff, but too many administrations are in too much of a rush to boast of results to justify long-term funding, politicians are mostly too ignorant of biology to comprehend our activities (Just what do you taxonomists do at CSIRO? one has asked), and there is too much pressure on staff to multiply titles in their personal bibliographies, no matter how trivial the contents, to justify salary increments. Most institutional workers are overburdened by so many meetings, conferences, report writing, grant applications and other paper work that they have little time left for pure research and certainly little or no opportunity to prepare large works such as this. How few are the monographs produced by the underfinanced, underequipped, understaffed museums, most of whose workers are frustrated for want of support and reduced to little more than caretakers, and then unfairly criticised for being so. But there must be something more that governs large monograph preparation.

    The Australian weevil fauna is ten times as large as that of the United Kingdom and more than double that of America north of Mexico, and preparation of guidebooks similar to this would be infinitely easier to prepare on those limited, well-studied faunas than one on the little-known, complex, difficult-to-understand Australian fauna. But skilled authors must be found who have intense interest in the subjects and with inspired, creative desire to produce the books and be prepared to make the many personal, family, financial and social sacrifices that such monographic work demands, and there must be sympathetic, supportive, understanding, institutional administrations. Thomas Edison said that Genius is 1% inspiration and 99% perspiration. So it is. Too often it is found that those with the will and inspiration lack the finance, and those with the finance lack the inspiration. I have been fortunate to have been able to arrange to be left mostly alone during the preparation of my books, although not without some criticism. I have published so little separately while writing these texts that had I been in some institutions I may have had my advancement curtailed or may have lost my job. In the light of my previous work, especially on the Hawaiian fauna, I was engaged by Douglas Waterhouse specifically to prepare these books, told to get on with the work and left mostly alone to do it. The results speak for themselves, and Australia now has a guide to its weevil fauna unlike that of any other country. Most workers who have the desire to prepare such books are, however, not as free as I have been to do the work. The writing was only part of the struggle. Manuscripts resting unpublished on authors’ shelves do not serve the purposes for which they are written. To get this monograph published we have had not only to sell the family jewels, but we had to sell the family farm as well. I have not taken a holiday away from the work for more than 20 years. I have served without respite for more than the past 10 years without salary or institutional pension, and (although the statement was later withdrawn when the extent and values of our work were emphasised) it hurt when a former executive stated that I am one of those who is indulging in his retirement hobby at taxpayers’ expense because I continue to use the Organisation’s facilities to produce an urgently-needed monograph of biological, ecological and economical importance. My books are far better than a tombstone, and they will serve and remind generations of workers that I left something worthwhile as I passed this way. That is satisfying payment for effort expended, and I can feel content that I have given something worthwhile to the Nation.]

    In 1845, Schoenherr recorded 7 141 weevil species then known from the entire world. In 1871 – 72, Gemminger and Harold (in the Munich Catalogue) listed the known weevil fauna of the world at about 11 600 species. The recent catalogue of the Central and North American fauna by O’Brien and Wibmer (1982) lists about 850 genera and 7 000 species, and their catalogue for South America (1986) lists about 9 000 species in about 1 000 genera with partial duplicating overlap on the North American catalogue, but those two catalogues unfortunately do not include the Anthribidae, Brentidae, Scolytinae and Platypodidae. There are no catalogues of the weevil faunas of Africa, the Oriental Region or Indonesia.

    Before this text was prepared there was only one species of Nemonychidae (Rhinomaceridae) recorded from Australia, but herein below Kuschel adds seven new genera and lists 14 species, 12 of them new discoveries. There were only three species of Bagous recorded from Australia, but O’Brien (1992) has reported 27. There were 15 genera and 22 species of Anthribidae listed in the previous major study, but here I list 72 genera and 174 species, but there are many more. There were six genera of the primitive Belidae previously known from Australia, but here I report 18, and this number will probably be increased when more specialised collecting is done. In Volume II I have given details of 75 species of Apionidae, including many undescribed species, four new genera, two new tribes, two new subfamilies, noted that none of the numerous species previously reported in the genus Apion belong to that genus and suggested that the actual number of Australian species probably exceeds 100, and they represent several new genera.

    The most recent catalogue of the weevils of Australia is the second edition of George Masters’ Catalogue of the Described Coleoptera of Australia, issued in 1886, more than 100 years ago! He listed the names of 1 263 species of weevils.

    Estimates of the numbers of described weevils are as variable as are the opinions of those making the estimates. O’Brien and Wibmer, 1978:89, upon a review of catalogues, concluded that there had been 671 genera and 5 166 species of weevils recorded from Australia. The same authors (The Use of Trend Curves of Rates of Species Descriptions: Examples from the Curculionidae [Coleoptera], The Coleopterists Bulletin 33(2):156 - 166, 1979) [unfortunately, this title was accidentally omitted from the bibliography in Volume III], using statistical methodology, estimated that the total number of described Australian Curculionidae to be 6 618. My more accurate count approximates 4 200, or about one-third less than the O’Brien and Wibmer estimate. They also estimate that there may be about 85 000 described weevils in the world, but if their other calculations are as far from reality as their estimates for the Australian fauna, their total figure is an inflated one. The late Sir Guy Marshall, the most experienced authority in this century, whose knowledge covered the world’s faunas, in 1956 estimated that there may be from 200 000 to 250 000, mostly undescribed, existing weevil species.

    On Taxonomy

    There have been many attempts to prepare an acceptable taxonomy for the Curculionoidea over the past 160 or more years. All have failed. Any author who believes that one can construct a definitive classification of the weevils in this or the next generation is, I believe, mistaken. We remain in the infancy of our work.

    Taxonomists too frequently are inhibited by tradition and too often attempt to fit organisms into existing taxonomic systems instead of revising the systems to fit the organisms. Taxonomic systems in use today in many parts of the world are remarkably poor and unworkable. However, the situation is now changing rapidly. The suprageneric classification is in a state of flux and uncertainty, and some new, untested taxonomies are being advanced. No general classification of the Australian fauna exists, and a satisfactory one will be long in coming.

    The earliest great advance in weevil taxonomy was that of Schoenherr in 1823, 1825, 1826, 1832 - 1845, and many of the conclusions reached by him remain sound. The next great advance was made by Lacordaire in his outstanding volumes in 1863 and 1866, and those volumes have guided all of us since. Then came the greatly used work of LeConte and Horn in America in 1876 as well as some lesser works in other areas. Various European provincial works have not stood the tests of time. The greatest advance in this century has been the stimulating, revolutionary work of Roy Crowson in 1955, and his publications launched a new era in the study of the classification of the Coleoptera. Following Crowson, Katsura Morimoto, the able, highly productive worker who has been misleading or unworkable. How, for example, can one use keys with such dichotomies as Size small as opposed to Size larger, or Rostrum short as opposed to Rostrum longer, or Rostrum much longer in male than female (and in this example the author confused the sexes) when one has only one sex?

    Funds to produce this Volume

    were given by

    Elwood and Hannah Zimmerman

    Derivation and Character of the Fauna

    The Australian weevils have two basic geographical sources: Gondwanaland and Indonesia. The Gondwanaland element may be divided into two parts. One of these contains autochthonous derivatives of southern origin whose allies appear to have become extinct elsewhere, if they ever occurred outside of Australia. Many of these probably arose after the break-up of Gondwanaland. The long, slow, isolated drift of the continent from Antarctica to near the Equator allowed ample time for much evolution and biotal change. The other element, if it can be considered such, shares close allies in New Zealand and/or southern South America and to a considerably less extent in southern Africa. I have found no evidence of any old Indian association. The Indonesian element is a large, diverse, late Tertiary and Recent invasion from the north which has resulted in a multitude of Indo-Pacific introductions and a few Ethiopian or Asiatic genera that have passed through the Oriental Region. The Indonesian invasion continues very active today, and also is the invasion of Indonesia by numerous Australian genera. Thus, northern Australia is now inhabited by so many Indonesian elements and eastern Indonesia is inhabited by so many Australian elements that one must consider the areas a blend. New Guinea reveals its geological youth with an island biota in an exuberant state of explosive speciation. Those who expect to discover ancient or unusual peculiar elements in that fauna will be disappointed. The Indonesian tropical element has penetrated most of the more humid east of Australia where it is well mixed with the truly Australian fauna. It is in the drier zones and the south and west where the Australian forms dominate. The southwest supports an extraordinary rich truly Australian flora, and associated with the several thousand endemic plants there are many endemic genera of weevils, but they have been inadequately collected and are poorly known. Because of specialised habits and the brief times when they are in their adult stages, it is not possible for many of them to be found by the occasional collector during hasty fieldtrips. Many of the most remarkable of all Australian weevils occur in that part of the Continent, but they are under great threat of extinction by unwise, unthinking land-clearing and widespread habitat destruction.

    At various times over the ages, Australia has been extensively invaded by the sea and broken into islands. Some of those isolated ancient lands developed their own distinctive weevil faunas, and these facts are reflected in the present character and distribution of the weevils.

    The alterations from warm and moist, to cold, to warm, to hot and dry, from rainforest to desert, from continent to archipelago to continent has brought great biotal change, extinction and evolution. Land open for colonisation is conducive to speciation. A biotal mozaic, often difficult to understand, has resulted.

    I regret that I can write nothing about Australian fossil weevils; I have never seen one. Because of the ubiquitous nature of the group over eons of time, it is surprising that none have been recorded in Australia, especially in view of the abundance of well-preserved plant fossils. Mesozoic fossils are especially desired.

    The primitive weevil families (see the table on page 5) Anthribidae, Nemonychidae, Belidae and Caridae, are well represented in Australia. There are many Australian Anthribidae of old and new derivation. The nemonychid fauna is either the world’s largest or second only to that of South America. There are no known Nemonychidae in Africa. Australia has the largest, most complex fauna of Belidae. Excepting a small invasion of Australian Rhinotia in New Guinea and neighbouring islands, the Belidae are confined to Australia, New Zealand and South America. In view of the rich Australian belid fauna, it appears surprising that the allied primitive Oxycorynidae, well-known as fossils in the Holarctic, whose living representatives now occur in America and Africa, with one African genus of palm pollen eaters extending into Indonesia, do not occur in Australia. Also, the Aglycyderidae, which are notably widely distributed in the Pacific from New Zealand and New Caledonia to the Marquesas Islands and Hawaii, and which are allies of the Belidae and Oxycorynidae, have no Australian representatives. Only four genera of Caridae are known: two in Australia and two in Chile. See this volume below for discussions of the fore-mentioned families.

    The Rhynchitidae are represented by the largest known complex of Auletobius species, a genus of almost world-wide distribution, but the rich Indonesian rhynchitid fauna has not crossed into Australia (unless the single specimen of Involvulus recently captured represents a native insect). Euops is the only genus of the Attelabidae to invade Australia from Indonesia where the family is multiform. Brentidae, strangely, evidently are represented in the endemic fauna by only three genera of myrmecophiles. This is in sharp contrast to the rich brentid faunas of South America, Africa and Indonesia. Where was Australia when those areas were seeded with Brentidae? See Volume II.

    The remarkable Eurhynchidae are purely Australian with a recent extension into New Guinea. There are no living relatives elsewhere. See Volume II.

    The Australian apionid fauna is now rather rich, although mostly undescribed, and it has been derived geologically recently from Indonesia. It shows no affinity to South America or Africa. See Volume II.

    One of the surprising features of the fauna is the poor representation of Rhynchophoridae, a group noted for its abundance in America, Africa, the Oriental Region and Indonesia. Only one monotypic genus is considered to be an Australian endemic. See Volume III.

    Aonychus, an isolated endemic genus which I have placed temporarily in association with Tadius in the Erirhinidae, in the newly revised sense, is an enigma about whose origin I cannot now speculate. See Volume II.

    In the Gonatoceri, or advanced weevil groups, the Amycterinae are usually considered the most singularly characteristic of the Australian weevil fauna. They appear as strange to some overseas observers as are the kangaroos. Their only living relatives are in South Africa. Among the more than 400 kinds are species as remarkable in appearance as are the homed dinosaurs. Many are remarkably adapted for life in the vast arid lands of the continent. They may be progenitors of the Adelognatha. See Volume III, and see Volumes V and VI for coloured illustrations.

    The Adelognatha, or broad-nosed weevils, are numerous and multiform in Australia. The fauna is a large mixture of old and derived forms from Gondwanaland, some of them especially unusual, and Tertiary and Recent descendants of Indonesian introductions (Myllocerus, for example). The Entimini, with the large complexes of Leptopius, Mandalotus and associates, is dominant. Interesting developments are the species of several unrelated genera that independently have evolved tanyrhynchiniform rostra. See Volume IV.

    The Aterpinae (including the Rhadinosominae) are best represented in Australia where there are numerous distinctive genera. There are a few allies in New Zealand and South America, but, unless there are some genera unrecognised as Aterpinae, there are none in Africa.

    Numerous Bagous of the Bagoinae are recent Indonesian derivatives.

    There are many Baridinae, especially in the north, which also are of geologically recent Indonesian origin.

    The Campyloscelinae and Ceutorhynchinae contain only a few endemics of Indonesian origin.

    There are numerous Cossoninae. Many of these have New Zealand affinities and others are of unknown derivation.

    The most diverse, multitudinous and ubiquitous group is the Cryptorhynchinae which contains about 160 described genera and 1 000 species, but with many genera and species undescribed. Many of these are unusually distinctive, and they form complexes of ancient and modern forms. Some of the genera are shared with New Zealand, others with Indonesia and most evidently are Gondwanaland derivatives with extensive autochthonous evolution and radiation.

    The Curculioninae, in a broad, newly revised sense, which contains the flower weevils, is so complex and inadequately known that little can be said about them now. The Derelomini are of Indonesian affinity, the Eugnomini have close relatives in New Zealand, adjacent islands and southern South America, and some extend into New Guinea.

    The distinctive Haplonychini are characteristically Australian with extensions into some southern Polynesian islands, but they are unknown elsewhere.

    The Storeini include the dominant, multiform Storeus, Emplesis, Melanterius and allied genera of uncertain geographical affinity, but some South American genera may be allied.

    The endemic Ramphini are of recent Indonesian origin.

    The numerous tychiine-like and allied genera are too poorly known for comment, but many are probably of old southern origin with considerable evolution and radiation in association with the remarkable diversification of flowering plants in Australia.

    The Gonipterinae form another of Australia’s distinctive groups of weevils of unknown affinity whose larvae feed externally on the foliage of Eucalyptus and some other Myrtaceae.

    The Hyperinae contain several endemic genera of undetermined affinity and others that have come from the north.

    The Cleoninae have only a few endemic species of Lixus of recent Indonesian origin.

    The Magdalininae, known elsewhere in the Holarctic and Nearctic, have the greatest proliferation of genera and species in Australia. Their nearest relatives are in South America. Strangely, nearby New Zealand lacks the subfamily.

    The Molytinae are well represented by numerous endemic genera, including peculiar forms. Here belong the unusual cycad-eating Tranes complex of genera of undetermined affinity. The Phrynixini form a cluster of genera found in New Zealand and Australia.

    The Rhythirrininae, a compound group as presently constituted, contains some genera with apparent South African affinity and others allied to New Zealand or South American genera.

    The Zygopinae are all of geologically recent derivation from Indonesia.

    Strangely, although there are numerous Scolytinae and Platypodididae in Australia, most of them are introductions. Only a few are proven endemics of undetermined affinities.

    An unusual feature of the Australian weevil fauna is the large number of species with basally fused tarsal claws, species with single claws, others which lack claws and others which have lost the entire terminal tarsal segments. Such species are found in many genera of Australian Adelognatha, Baridinae, Curculioninae, Diabathrariinae, Erirhininae and Magdalininae. The reasons for these reductions are unknown. They occur on weevils attached to hostplants with foliage as distinct as the broad leaves of Eucalyptus and other plants and as narrow as the needle-like foliage of Casuarinaceae.

    Nothofagus, southern beech, is an ancient genus of plants with a long fossil record in Australia and formerly much more widely distributed than it is today. Only a few weevils have been found to feed upon the genus in Australia. In New Zealand and South America, it is an important host for weevils. Why not in Australia?

    Weevils, Weevils, Weevils Everywhere

    Weevils inhabit all the earth where there is terrestrial vegetation. They are among the most successful and multidinous forms of terrestrial life. They constitute the largest family group of the animal Kingdom. They may be found from the Arctic to the subantarctic, and no habitable island, no matter how remote, escapes their colonisation. No desert that supports vegetation is too dry for them, but it is in the humid tropics where they reach their greatest diversity, and there great numbers of species and individuals occur. Some live along the seashore in or under driftwood or algae or among cryptogams on rocks where they may be wet by sea water, but none is known to invade the sea. Many are aquatic and their larvae may be internal or external feeders on fresh water plants. Many live in the soil in both larval and adult stages; many are blind troglodytes. They feed upon lichens, fungi, mosses, and some transplant and cultivate fungi for food for their larvae. Many feed upon ferns, others on cycads or primitive Equisetum horsetails. Multitudes feed upon grasses and conifers. Probably no flowering plant escapes their notice, and many species of higher plants have many species obligately attached to them. Many higher plants depend upon weevils for pollination. All parts of plants are eaten by weevils from roots, bark, sapwood, heartwood, stems, twigs, buds, flowers, pollen, fruits, seeds and sick, dying, dead and decaying plant material. Many are leaf-miners, others make galls on plants and others inhabit plant galls made by other organisms. The larvae of one Australian species develops in marsupial dung. A few are predaceous on other insects. Many species such as the grain weevils, cotton boll weevils, various stem, fruit and nut weevils, palm, sugarcane and banana weevils and numerous defoliators are pests of major economic importance. Some with soil-dwelling larvae destroy the nodules of nitrogen-fixing bacteria and roots and are serious pests of pastures. Some native species may become pests when crops are planted in their areas, and many foreign pest species that may be accidentally introduced are a constant threat to Australian agriculture. A number are being used in the successful biological control of weeds.

    Most weevils are able flyers, but many have reduced wings and are flightless. Many are parthenogenetic with males unknown or rarely produced. The eggs of many weevils are deposited in the soil, many are laid on the surfaces of the hostplants, others are laid between glued-together leaves, some are inserted into holes or cavities prepared in or on the hosts by use of the ovipositor or the specially formed caudal abdominal tergite, but perhaps the majority of eggs are laid in holes drilled in the hosts by the mandibles at the ends of what are often extraordinarily long rostra of the females. Some larvae are exposed feeders on foliage, and some of those conceal themselves by piling excrement on their backs, but the greater number are internal borers living in tunnels eaten in various parts of their hosts. Some form silken net cocoons in which to pupate, others pupate on or within tunnels in their hostplants and many pupate in the soil. Development from egg to adulthood varies from days to several years. Some wet forest species support vegetable gardens of various kinds of primitive plants that grow on their backs. Many are nocturnal and are rarely seen. Some are myrmecophiles. Many adults are ravenous defoliators, others feed upon pollen. Great numbers are inhabitants of forest-floor litter and are valuable aids in the decomposition of dead vegetable matter. Weevils, weevils everywhere!

    Since the advent of transoceanic jet-aircraft transport, Australia is now constantly threatened by invasion from all continents and many islands by a multitude of weevils of economic importance many of which may cause great damage to agriculture and forestry if they became established. The introduction of the cotton boll weevil from America would devastate the cotton industry. The introduction of lucerne and clover weevils would cause graziers great losses. The West Indian sweetpotato weevil, which is now established on numerous nearby islands, may invade Australia at any time. There are weevils that may be introduced by an overnight aircraft flight from California that could seriously damage our extensive pine plantations. Other exotic weevils may be introduced that may damage or devastate fruit and nut crops, sugarcane, rice, pastures, vegetables, orchids and other flowers, shrubs, timber trees - any grass, grain, herb, shrub, tree, fruit, berry or seed is vulnerable. Australia requires specialist workers who can recognise friend from foe and to be able to advise appropriate authorities when dangerous situations arise and to be prepared to supply them promptly with advice and references to pertinent literature concerning life histories, hostplants, distribution, predators and parasites, and to be able to give advice regarding agents that may be useful in the biological control of noxious weeds.

    It has now fallen to me to contribute what I can to bring from obscurity into the light of science knowledge of our complex, compound, remarkably evolved, multitudinous weevil fauna. I have used illustrations to convey more understandable information to those who use these volumes than had I written additional thousands of pages of text. Although about 10 000 illustrations have been prepared for this work, I consider it incompletely illustrated to attain my goal.

    It has already been found that my admonishments in Volumes V and VI to note that only some of the Australian weevils are illustrated on the colour plates has

    Enjoying the preview?
    Page 1 of 1