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...