Australian Entomologist, 2012, 39 (4): 231-237 231 COURTENAY NEVILLE SMITHERS: HIS SCIENTIFIC CONTRIBUTIONS TO THE 8SMALL ORDERS9 OF INSECTS T.R. NEW Department of Zoology, La Trobe University, Vic 3086 Abstract Courtenay Smithers9 contributions to the knowledge of Psocoptera and other orders, notably Neuroptera and Mecoptera, are summarised. They extended over some 50 years and established him as a leading authority on these orders. His contributions to systematics, biogeography and biology of Psocoptera, particularly within the Australian region, were accompanied by compilations of global catalogues and bibliographies of immense value to others. His many publications assure him of a persistent place in the history of psocopteran studies. Introduction Courtenay Neville Smithers (1925-2011) was known and respected widely as an excellent naturalist with a keen, life-long sense of curiosity and wonderment that led to a variety of scientific interests, together with an enduring enthusiasm about communicating his passions to others. These personal traits rendered him a highly effective publicist for entomology and led to his writing of several books to introduce insects to the wider public. His 1971 8Australian Insects in Colour9, with Anthony Healy, was a pioneer in all-colour popular publication, while his 8Handbook of Insect Collecting9, first published in 1981, remains a useful and readable resource. His 8Backyard Beekeeping9 has just been republished 25 years after it first appeared in 1987. However, his major formal scientific contributions focused on studying the systematics and biogeography of a commonly ignored insect order, the Psocoptera, augmented by original contributions on two other 8small orders9, the Neuroptera and Mecoptera. His studies on these insects assure him of a lasting place in their documentation. Psocoptera Smithers9 taxonomic interest in Psocoptera (the barklice and booklice) flowed from the 1950s (A. Smithers 2012), with his first papers on these insects (in 1956) the source of a stream that reached more than 150 contributions over more than half a century and established him firmly as a leading world authority for much of that period. When I started to work on British psocids (in 1965), I wrote to Courtenay at the Australian Museum for his advice, and his kind and encouraging response 4 together with a set of his reprints then available 4 marked the start of a long friendship founded in our mutual interests in 8the other orders9. I met him first in England in 1967 and next at the XIIIth International Congress of Entomology in Moscow (August 1968), an occasion that marked perhaps the first major international gathering of psocid workers, with several meeting for the first time (Fig. 1); it was not until the 1980s that such a meeting of psocopterists, always few in number, was held again. Following my arrival in Australia in 1970, we continually exchanged ideas on psocids and lacewings. 232 Australian Entomologist, 2012, 39 (4) | | | i $ i | / i Fig. 1. Gathering of psocopterists outside Moscow University, August 1968. Left to right: A.M. Nadler (USA), T.R. New (UK), A. Badonnel (France), I.W.B. Thornton (Australia), C.N. Smithers (Australia) (Photo: V.N. Vishniakova). While most of his published work deals with systematics, distributions and relationships of psocids, Courtenay was always aware that he was dealing with dynamic living organisms and, somewhat unusually for that era, his taxonomic studies flowed in large part from material collected during his own field studies, rather than from specimens mostly collected by others and submitted for identification. Biological oddities intrigued him greatly. One such was his discovery of the remarkable phragmotic nymphs of Psilopsocus mimulus Smithers (Figs 2-3), which resemble small bark beetles and live in twigs, blocking the entrance to their tunnels with their heavily sclerotised abdomens (Smithers 1995). His knowledge spanned the world fauna. Among his first major self-appointed tasks was to bring together the then very widely scattered literature on Psocoptera into a single, easily available and convenient format: his 8Bibliography of the Psocoptera9 (Smithers 1965, with drafts circulated to most specialists for comment and augmentation) was of incalculable value to tyros such as myself. His comment in the introduction that it was 8an attempt to provide a working tool which will relieve students ... of the drudgery of compiling their own bibliographies9 was indeed pertinent. The Bibliography was followed by a World Catalogue of species (Smithers 1967). The two works together provided, for the first time, a firm basis for later studies; subsequent progress in understanding Australian Entomologist, 2012, 39 (4) 233 Psocoptera over the last half century is largely due to Smithers9 Herculean efforts in producing these accounts. It is worth remembering that these compilations were made long before availability of computers, email and even routine use of photocopiers 4 so that obtaining and checking the numerous obscure references for verification could entail weeks of patience awaiting international postage, or laboriously copying out details by hand during sporadic visits to major institutional libraries. They were followed (Smithers 1972) by publication of his Ph.D. thesis, a global synthesis of psocid classification and phylogeny, with illustrated diagnoses of all genera and families then known and discussions of putative relationships that set a basis for much later consideration. It remains a potent reference source of earlier information and diagnoses. 3 Figs 2-3. The wood-boring psocid, Psilopsocus mimulus Smithers: (2) adult male from Lindfield, NSW (AM Paratype K68224); (3) final instar nymph with abdomen modified to plug burrow, from same series (Photos: D. Britton, Australian Musem) These landmark publications were revised later, with new keys to families and genera (Smithers 1990), a revised Bibliography (with Charles Lienhard: Smithers and Lienhard 1992) and Smithers9 magisterial joint volume (Lienhard and Smithers 2002), as a world catalogue and bibliography, updating these to comprise perhaps the single most significant publication ever issued on the order. The classification used there was essentially that of Smithers (1990) and pre-dated current opinion that the order is not monophyletic but intertwined intricately (as 8Psocodea9) with the true lice, Phthiraptera (Yoshizawa and Johnson 2006). Psocoptera had become much better known between 1967 and 2002, in no small part reflecting the impact of the publications noted above in collectively facilitating the perspective needed for others to study these insects. The known richness of Psocoptera rose from 1605 species (in 197 genera) in 1964 to 4408 species (371 genera) by 2000, together with an increase from 31 to 41 generally recognised families. Smithers was an active contributor to this increase. Alone or with his collaborators (notably the late Ian Thornton and in associations that led to global dominance of Psocoptera 234 Australian Entomologist, 2012, 39 (4) Studies by Australians for some two decades), he described somewhat more than 350 new species of psocids, across about 25 families. Many of these came to his attention though extensive field work and collecting in Australia and 4 with Thornton 4 during a major Australian Research Grants-funded Survey during the 1970s of the various islands, including New Guinea and the Melanesian Arcs, to the north and east of Australia. While his first major regional synopsis dealt with the psocids of Madagascar (Smithers 1964), his first paper from the Australasian region (following his move from South Africa to the Australian Museum in 1960) dealt with Philotarsidae from Macquarie Island (Smithers 1962). However, the account that founded his reputation in the Australasian fauna was a major paper on the New Zealand fauna (Smithers 1969), with 15 new species bringing the country9s total to 43 species. His early African experiences assuredly whetted his appetite for biogeographical puzzles centred in the 8southern continents9 and the regional fauna remained his primary focus for many years. From the early 1960s he published notable species additions and family synopses of the Australian psocids and, by the time he prepared the entry on Psocoptera for the Zoological Catalogue of Australia (Smithers 1996), he had described well over a third of the Australian species then known. These included records of several families new to the country, as well as a number of novel genera that hinted at the considerable complexity and high levels of endemism of the fauna. His successive chapters in the original (1970) and revised (1991) editions of 8The Insects of Australia9 revealed an increase from 120 to 299 species over that period. Regional accounts, such as for South Australia (Smithers 1984) and (jointly) for Norfolk Island (Smithers and Thornton 1974) and Lord Howe Island (Smithers and Thornton 1975), were augmented by substantial family accounts for the Melanesian Arc psocids that collectively set a new perspective for their diversity and evolution within the region. Families such as Philotarsidae, Calopsocidae, Myopsocidae and Psocidae proved to be far more diverse, intricate and biogeographically informative than suspected before these studies began. Elucidating their features did much to enhance understanding of the Australian fauna and how this had developed. Many of the places visited in Melanesia had never been explored properly for psocids, although the few taxa described sporadically from New Guinea and elsewhere suggested the likely richness of those areas. The fieldwork in remote areas was sometimes hazardous, frequently adventurous and innovative. Courtenay recalled some of these adventures in our tribute to Ian Thornton (New and Smithers 2004). However, his dedication and care resulted in the Australian Museum now housing one of the all-time greatest and most representative collections of this order. Smither9s seminal work on psocids has been commemorated by his colleagues by seven species named 8smithersi9. One of these he himself Australian Entomologist, 2012, 39 (4) 235 synonymised, a step that accorded him some amusement. Members of the genus Sigmatoneura Enderlein (Psocidae) are characterised by extensive sexual dimorphism, so that males and females can be associated clearly only by co-incidence. Before this sexual difference was appreciated, I had described a male and female from Nigeria (collected at the. same general locality but a year apart) as separate taxa (New 1973). Courtenay later decided that the more parsimonious approach was to consider them conspecific; unfortunately, the second described was 8smithersi9. In the same paper, though, he also sunk Scaphopsocus Smithers as a new synonym of Sigmatoneura! Two other psocid species are named 8smithersorum9, acknowledging Smila9s continued encouragement and support throughout their long partnership. Courtenay himself named a number of psocids 8alettae9 in appreciation of Smila9s notable contributions. Two patronymic genera also exist: Smithersia Thornton (Myopsocidae) and Smithersiella Badonnel (Caeciliusidae). Neuroptera The psocopteran patronyms are augmented by five patronymic lacewings, marking another of his favourite insect orders. Courtenay9s interest in Neuroptera was also a long one and, although he did not publish as extensively on them, his knowledge was broad and exceptional. He described a number of new Australian species, across four families, and wrote a number of biological notes. Sadly, his long-projected revision of Australia9s spongeflies, Sisyridae 4 a project we talked about at our first meeting, when I was able to show him one of the three British species in the field 4 was not completed, although he described several new species in isolation, including Sisyra pedderensis Smithers, an endemic to Tasmania9s Lake Pedder (Smithers e} al. 2008) and the last insect he described (Figs 4-6). Other Orders His interests in 8the other orders9 also included Zoraptera (the chapter in 8Insects of Australia9) and Mecoptera, for which he described a remarkable endemic genus, 7ytthobittacus, from the Blue Mountains (Smithers 1973) and produced a synopsis of Australian taxa (Smithers 1987). A full bibliography of his publications is given by Britton (2012). In an era in which specialisation is the norm, Courtenay Smithers9 entomological interests remained broad, but never forgoing depth and authority. His commitment to advancing knowledge of Australian insects was coupled with an equally strong commitment to their conservation. His legacy is enduring and few could claim to have changed perspective of any insect order to the extent that his studies on psocids, in particular, have done. His responses to queries were always generous and informative and the encouragement he gave to me, and to other less experienced colleagues, will be remembered with gratitude and affection. 236 Australian Entomologist, 2012, 39 (4) Figs 4-6. Adult, host and habitat of the spongefly, Sisyra pedderensis Smithers from Lake Pedder, Tasmania: (4) adult female (Photo: G.N. Forteath); (5) encrusting freshwater sponge, Radiospongilla pedderensis Osborn, Forteath and Stanisic, larval food of Sisyra pedderensis (Photo: A.W. Osborn); (6) Lake Pedder, Tasmania, type locality of both species (Photo: A.W. Osborn). Acknowledgements I am very grateful to Dr David Britton and Professors Nigel Forteath and Andrew Osborn for use of their photographs. References BRITTON, D.R. 2012. Dr Courtenay Neville Smithers 1925-2011. A tribute. General and Applied Entomology 40: 25-34. LIENHARD, C. and SMITHERS, C.N. 2002. Psocoptera. World catalogue and bibliography. Instrumenta Biodiversitatis V. Muséum d9histoire naturelle, Genéve; 745 pp. NEW, T.R. 1973. A collection of Psocoptera from Nigeria. Occasional Papers of the Entomological Society of Nigeria, No 10: 22 pp. Australian Entomologist, 2012, 39 (4) 237 NEW, T.R. and SMITHERS, C.N. 2004. Ian W.B. Thornton - notes on his life and work, with list of publications. Pp 9-30, in: Garcia Aldrete, A.N., Lienhard, C. and Mockford, E.L. (eds), Thorntoniana, a commemorative volume for Ian W.B. Thornton (1926-2002). Publicaciones Especiales del Instituto de Biologia 20, Universidad Autonoma de México, Mexico; 205 pp. SMITHERS, A. 2012. Courtenay Smithers 4 before Australia. Australian Entomologist 39(4): 211-220. SMITHERS, C.N. 1962. Insects of Macquarie Island: Psocoptera: Philotarsidae. Pacific Insects 4: 929-932. SMITHERS, C.N. 1964. On the Psocoptera of Madagascar. Revue de Zoologie et de Botanique africaines 70: 209-294. SMITHERS, C.N. 1965. A bibliography of the Psocoptera (Insecta). Australian Zoologist 13: 137-209. SMITHERS, C.N. 1967. A catalogue of the Psocoptera of the world. Australian Zoologist 14: 1- 145. SMITHERS, C.N. 1969. The Psocoptera of New Zealand. Records of the Canterbury Museum 8: 259-344. SMITHERS, C.N. 1972. The classification and phylogeny of the Psocoptera. Australian Museum Memoirs 14: 1-349. SMITHERS, C.N. 1973. New species and records of Australian Bittacidae (Mecoptera). Journal of the Australian Entomological Society 12: 296-300. SMITHERS, C.N. 1984. The Psocoptera (Insecta) of South Australia. Records of the South Australian Museum 18: 453-491. SMITHERS, C.N. 1995. Psilopsocus mimulus Smithers (Psocoptera: Psilopsocidae), the first known wood-boring Psocoptera. Journal of the Australian Entomological Society 34: 117-120. SMITHERS, C.N. 1987. Synopsis of Australian scorpion flies (Mecoptera). General and Applied Entomology 19: 31-44. SMITHERS, C.N. 1990. Keys to the families and genera of Psocoptera (Arthropoda: Insecta). Technical Reports of the Australian Museum 2: 1-82. SMITHERS, C.N. 1996. Psocoptera. Pp 1-79, 333-335, 363-372, in: Wells, A. (ed.), Zoological catalogue of Australia. Vol. 26. Psocoptera, Phthiraptera, Thysanoptera. CSIRO Publishing, Melbourne. SMITHERS, C.N. and LIENHARD, C. 1992. A revised bibliography of the Psocoptera (Arthropoda: Insecta). Technical Reports of the Australian Museum 6: 1-86. SMITHERS, C.N. and THORNTON, I.W.B. 1974. The Psocoptera (Insecta) of Norfolk Island. Records of the Australian Museum 29: 209-234. SMITHERS, C.N. and THORNTON, I.W.B. 1975. The Psocoptera (Insecta) of Lord Howe Island. Records of the Australian Museum 29: 453-471. SMITHERS, C.N., FORTEATH, G.N. and OSBORN, A. 2008. A new species of Sisyra Burmeister (Insecta: Neuroptera: Sisyridae) from Lake Pedder, Tasmania. Australian Journal of Entomology 47: 77-79. f YOSHIZAWA, K. and JOHNSON, K.P. 2006. Morphology of male genitalia in lice and their relatives and phylogenetic implications. Systematic Entomology 31: 350-361.