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Jeyaraney Kathirithamby

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Beschreibung

Comprehensive account of Strepsiptera biology, including their taxonomy, morphology, fine structure, physiology, and behaviour, ranging from genes to phylogeny

Beautifully illustrated in full color throughout, with original up-to-date material from the author’s research collection, Insect from Outer Space: The Biology of Strepsiptera is the first book to provide a comprehensive guide to the unique biology of an intriguing group of parasitoids that, until relatively recently, have remained something of a mystery in terms of both phenotype and genotype.

The author explores the exciting biology and life history of these extraordinary insect parasitoids. In addition, the book provides a history of each family, and an account of the questions surrounding the evolution of Strepsiptera, explaining their position within the phylogeny of insects.

Written by a leading international expert in the field, this book includes information on:

  • Strepsiptera, which are among the least species-rich insect parasitoids, but they parasitise hosts from seven orders and 36 families of insects
  • The two examples of polyphensim that Strepsiptera exhibit are: i) free-living planidia that moult to endoparasitic larvae; and ii) the neotenic females of derived families
  • Adult males are free-living, while females are free-living in primitive families, and are endoparasitic and neotenic in derived families
  • Metamorphosis is a crucial process in the evolution of the sexually dimorphic Strepsiptera, and the neotenic females in derived families undergo a metamorphic transformation in the cephalothoracic region
  • Differences in Strepsiptera life cycles between the suborders Mengenillidia (a more ancestral lineage) and Stylopidia (a recent suborder)

Insect from Outer Space: The Biology of Strepsiptera is an essential reference work for entomologists and biologists alike who seek to unravel the secrets of these extraordinary creatures. The text is also an excellent learning resource for graduate and undergraduate students because the unique biology of Strepsiptera answers a wide range of biological questions.

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Veröffentlichungsjahr: 2025

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Insect from Outer Space

The Biology of Strepsiptera

Jeyaraney Kathirithamby

Department of Biology and

St Hugh’s College

University of Oxford

Oxford, UK

This edition first published 2025© 2025 John Wiley & Sons Ltd

All rights reserved, including rights for text and data mining and training of artificial intelligence technologies or similar technologies. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, except as permitted by law. Advice on how to obtain permission to reuse material from this title is available at http://www.wiley.com/go/permissions.

The right of Jeyaraney Kathirithamby to be identified as the author of this work has been asserted in accordance with law.

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Library of Congress Cataloging‐in‐Publication Data Applied for:

Hardback ISBN: 9781118406519

Cover Design: WileyCover Image: Courtesy of Mike Hrabar

About the cover:Left ‐ Adult male Xenos peckii emerging from a puparium in host Polistes fascatus (Hymenoptera: Vespidae) x15.

Right ‐ Adult male Xenos peckii inseminating a neotenic female stylopised in Polistes fascatus (Hymenoptera: Vespidae) x15.

In memory of the two colleagues who inspired meFiona Proffitt (1966–2013)

and

Hans Henderickx (1961–2016)

The organisation and habits of the tiny Stylops are as much proper subjects of investigation, of wonder and admiration, to the right‐thinking mind, as are those of the Leviathan of the deep, or of the Elephant of the forest; whilst their very diminutiveness and isolation, like all microscopic analyses of organism, or singleness of action, tend greatly to facilitate our investigation of principles, and lead us more easily to understand those on which structure is formed, and function and instinct unfolded.

George Newport (1851). The History and General Anatomy of Meloë, and its affinities, compared with those of the Strepsiptera and Anoplura, with reference to the connexion which exists between Structure, Function, and Instinct. Transactions of the Linnean Society20: 321–356.

‐‐‐‐‐‐‐‐‐‐‐‐‐

But their size increases by the day, especially when they have enough food. Some of them attain their full size in several weeks; others require up to two months. Many push off their skin completely three or four times, just as a person removing a shirt over his head, and change to different colours; some reappear as they were before.

Maria Sibylla Merian (1679). Der Raupen wunderbare Verwandelung und sonderbare Blumen‐nahrung. Nuremberg: M. S. Merian.*

Note

*

The Flowering of Ecology: Maria Sibyilla Merian: Caterpillar Book, ed. Kay Etheridge, English translation by Michale Ritterson. (Brill 2010, p. 142).

Preface

Strepsiptera are an enigmatic group of entomophagous parasitoids that most entomologists have either heard or read about, but very few have actually seen as live specimens. For decades, they remained a neglected group, and the few entomologists who studied them focused on the taxonomy of a few species. I dedicate this book to Fiona Proffitt and Hans Henderickx. The title of the book Insect from Outer Space is from Fiona Proffitt’s article in Science in 2005, which bore the same title. She came to see me while she was a graduate student, since she had become fascinated with what she had read about Strepsiptera and was eager to write an article for Science in order to gain experience as a science writer. Sadly, she passed away in New Zealand in 2013. Hans Henderickx, was a gifted entomologist and natural history photographer.

While revising chapters during the COVID‐19 lockdown, I read an article by Stephen Asma in The New York Times (18 April 2020), which rightly pointed out that the coronavirus is a zoonotic ‘spillover’, like parasitism and predation, which are not sent as punishments but represent ‘business as usual’. Coronavirus was simply passing over genes to the next generation – a Darwinian evolution. Stephen Asma gives an example of Rhizocephala barnacles that invade as seeds in their crustacean hosts and spread throughout the host organism, castrating it, stopping it from moulting, but keeping it alive for years. Another example he cites is the tarantula hawk wasp, which lays an egg in the host tarantula after paralysing it, while any emerging larva feeds off the host but avoids its vital organs. As Stephen Asma says, ‘Not even the most inventive Hollywood writers can spin tales of this fantasy, yet it is the bread and butter of every day biology’.

In this book, I give another example of a fantasy in nature: the entomophagous parasitoid Strepsiptera. This is a parasitoid that exploits its host in many different ways in order to pass on its genes to the next generation. Most zoonotic organisms, such as the coronavirus or parasites and predators, have a narrow host range to complete their mission in life. Strepsiptera, however, have 36 families of insect hosts (apter‐, endo‐ and exopterygota), which they stylopise. They have coevolved with their various host life cycles in order to pass on their genes. In this book, I outline the ways they can castrate the external and internal genitalia of their hosts, change their behaviour, halt their moulting, while the host remains alive and mobile so that the strepsipteran can complete its life cycle. When I relate the life cycle of Strepsiptera, many have suggested that I am writing a science fiction story, since the life cycle is a ‘tale of fantasy’. But in reality, Strepsiptera are simply carrying out the ‘bread and butter’ functions of everyday life.

My own lifelong obsession with this group began when I ‘stumbled’ onto stylopised rice leaf‐ and plant‐hoppers in Malaysia, where I was born. Many of the leaf‐ and planthoppers I collected were found to be stylopised by Strepsiptera, and from that time onward, I became mesmerised by these parasitoids. When I came to work with Professor George Varley as Rhodes Visiting Fellow at St Hugh’s College, Oxford, I found that, except for its taxonomy, very little else was known about this enchanting parasitoid. Numerous questions arose: How many endoparasitic larval instars are there? What range of hosts do they parasitise? What is their distribution? What physiological damage do they do to their hosts? What is their sex determining mechanism? I must confess that some of these questions still remain unanswered. Although many museums have a collection of Strepsiptera (usually small), I decided early on that the best way to study them was to collect the material myself and to aim at a study not only of the strepsipteran parasitoid but also of its effects on the 36 families of insect hosts.

I made contacts with various colleagues and also went to the field to collect, which has taken me to Australia, Brazil, France, Germany, India, Italy, Japan, Malaysia, Mexico, Panama, Papua New Guinea, Spain and United States. I am also fortunate in that many collaborators have sent me, and continue to send me, material. At the start, my father used to accompany me to field trips in Malaysia. Some of my exciting experiences in various sites, such as Papua New Guinea, include a native man with a bow and arrow standing guard at one of the guest houses; a very pleasant view of the wonderful tropical rain forests when I was invited by a pilot to sit with him and his copilot in the cockpit on a flight from Popondetta to Port Moresby, a pleasure initially marred by my fear that it had been discovered I was carrying live grasshoppers; and a ride in a ‘trash boat’ from Barrow Colorado Island to Panama City and back, with waves from the canal that were so high I thought we were likely to drown.

I owe my understanding of the best way to collect Strepsiptera to the late George Varley, who had a study plot in his garden at the end of Apsley Road, in North Oxford, which had planthoppers stylopised by Javesella dubia. George would often join me on a hot summer’s day to ‘poot’ the stylopised specimens. At the end of these sessions, he would offer me a glass of potent home‐made beer, but I preferred his delicious home‐made ginger beer. These are moments I still treasure.

I was fortunate enough to be invited in 1983 to the CSIRO in Canberra to study the collection there. I worked on the taxonomy of the group and wrote descriptions of species of four families from Australia (Elenchidae, Halictophagidae, Corioxenidae and Myrmecolacidae). The late Edgar Riek left copious notes on the Strepsiptera that he had collected in Australia, which are referred to in this book. The late Mary Carver was a great friend and colleague while I was in Canberra. The late Ian Gauld (then editor of Systematic Entomology) encouraged me to write a review on Strepsiptera, which was published in 1989, while another review of the group by me appeared in 2009 in the Annual Review of Entomology.

I am indebted to now absent friends: George Varley, Richard Southwood, Doug Waterhouse, Max Whitten, Bill Hamilton and Nadia Waloff, who encouraged me early on in my career to work on this group. More recently, Peter Holland has been very supportive, and Chris Graham helped me during times when I most needed advice. I am grateful to Malcolm Davies for translations of various European articles.

St Hugh’s College has been my second home in Oxford since I came there as Rhodes Visiting Fellow. I truly treasure friends and colleagues who have been supportive to me. The Principal, Lady Elish Angiolini, gave me rooms in St Hugh’s when the Zoology Tinbergen Building was abruptly closed in 2017, a critical time for me, since I was checking the proofs of my co‐authored book, ‘Maria Sibylla Merian: Artist, Scientist, Adventurer’ (Getty 2018). For this generous gesture, I am immensely grateful.

The following kindly gave me comments on the relevant chapters: Xavier Belles, Elke Buschbeck, Jerry Cook, Michael Engel, Silvio Erler, Deniz Erezyilmaz, Raghavendra Gadagkar, Mike Hrabar, Yoshitaka Kamimura, Conrad Labandeira, Yuta Nakase, Jim Nardi, Jordi Paps, Robert Poulin, Markus Riegler and Amy Toth.

Kathleen Diston and Danielle Czerkaszyn of the Oxford University Museum of Natural History helped me with several papers, while Amanda Ingham gave me much needed aid with scanning of the figures. Barbara Young, Mike Pitts and Rob Hoare patiently answered numerous questions regarding IT, and Alan Tod gave immense help with the maintenance of my microscopes.

The following collaborated with me during my studies of this extraordinary group of insects: Zoltán Ács, Ricardo Andreazze, Gawain Antell, Richard Askew, Zach Balzer, Juan Barrera, Laura Beani, Giovanni Benelli, Bastien Boussau, Alexandra Bryson, Marcella Carcupino, David Martin Casacuberta, Silvia Chafino, Christian Cocquempot, Jennifer Coggon, Katherine Child, Francesco Collantes, Liam Crowley, Danielle Czerkaszyn, Loic Dahan, Romano Dallai, Juan Delgado, Stefan Dötterl, Alexander Fateryga, Camilo Flórez, Dominique Fresneau, Gerard Gries, Jaime Gomes, David Grimaldi, Natallie Halbert, Justin Hatting, Alexander Hayward, Hans Hendrickx, Marc Hoffmann, James Hogan, Peter Holland, John Huelsenbeck, David Hughes, James Hunt, Marisano James, Spencer Johnston, Teiji Kifune, Regnard Kinzelbach, Helmut Kovac, Dino McMahon, Nora Khayi, Ragnar Kinzelbach, Helmut Kovac, Conrad Lebandeira, Gary Lechner, Meri Lähteenaro, Alain Livory, Yasudo Maeta, Shun'ichi Makino, Darren Mann, Shamir Marabet, Massimo Mazzini, Dino McMahon, Rebecca Millena, Yuta Nakase, Virgina Ocampo, Massimo Olmi, Hans Pohl, Jean‐Yves Rasplus, Gustavo Moya‐Raygoza, David Rentz, Evgeny Perkovsky, Jordi Paps, Robert Poulin, Jean‐Yves Rasplus, Hermish Robertson, Antonis Rokas, Jay Rosenheim, Rebecca Saff, Ruchira Sen, Karel Schoonvaere, Leo Shapiro, Mike Shaw, Ben Sheldon, Souto‐Vilaros, Zoe Simmons, John Smit, Takis Solulu, Steve Simpson, Amoret Spooner, Makino Shun’ichi, You Ning Su, María Camila Tocora, Stefano Turillazzi, Steve Taylor, Viviana de Oliveira Torres, Nicholas Vereecken, John Vile, Bo Wan Wang, Donald Windsor, Nick Wood and Cor Zonneveld. To all of them I extend my heartfelt thanks.

Last but not least, Juan Antonio Delgado, Department of Biology, University of Murcia, Spain, gave me advice on the figures and indeed executed many of them himself. To him I am most grateful.Finally, Xavier Belles and Malcolm Davies deserve many thanks for helping me check the proofs of this book.

Jeyaraney Kathirithamby

Oxford, UK

About the Companion Website

This book is complemented by a companion website.

www.wiley.com/go/kathirithamby/strepsiptera1e 

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