Niels Peder Kristensen, Honorary Member and former president of SEL, passed away on Saturday December 6^th 2014 in Copenhagen. While his death was not unexpected, its timing came earlier than we had thought or hoped. His loss is felt widely and intensely.

Born on March 2^nd 1943, Niels was the second child of Thorkil and Ellen Christine Kristensen (nee Nielsen). His father was an academic, politician and thinker who served as Minster of Finance in two different government cabinets, and later as General Secretary of the OECD. Growing up in such an environment undoubtedly had a profound influence on Niels’ own world view, one which was powerfully international in its expression, yet retaining a strong interest and deep concern for Danish issues – local and national.

Niels developed an interest in entomology and lepidopterology in particular at an early age, and once told TJS about the first time, when eight years old, he visited the Entomology Department at the Zoological Museum in Copenhagen (ZMUC) ‘clutching his father’s hand’. After completing high school at Birkerød Statsskole in 1961, Niels enrolled as a biology student at the University of Copenhagen, and quickly became a regular visitor to the Entomology Department of the Museum, where he had already started as a volunteer during his last years at high school. In 1965, while still a student, he published his first paper, which was on the faunistics of Danish cicadas. From the very start of his scientific career, one of Niels’ abiding interests was the evolution (particularly evolutionary morphology) of primitive Lepidoptera. Indeed, the work for his Mag. Scient. degree was on the comparative morphology of the primitive glossatan family, Eriocraniidae. During this study, Niels spent the academic year 1966–67 at the University of Bristol, working with the eminent and extremely knowledgeable British entomologist Howard E. Hinton, who was at that time pioneering the use of scanning electron microscopy in entomology. It was while working with Hinton that Niels came to appreciate the value of SEM in comparative morphology. Through it, he made the surprising discovery (published in 1970) that the most primitive Lepidoptera have flat, solid wing scales (i.e. lacking an internal lumen), a condition contrasting strongly with the hollow wing scales generally found in Lepidoptera.

After obtaining his Mag. Scient. degree in 1968, Niels was offered a tenure-track position at the ZMUC as Amanuensis (Assistant Professor). He was promoted to Associate Professor in 1972 and appointed as Full Professor of Entomology in 1995. In 1970, Niels visited one of Europe’s foremost comparative invertebrate morphologists, Jean Chaudonneret, at the Université de Dijon where he enhanced considerably his skills in insect histology and semi-thin sectioning. Working with Hinton, Chaudonneret and Karl G. Wingstrand, the professor of comparative morphology at the University of Copenhagen, unquestionably had a profound influence on Niels’ development as a scientist. He often referred to the effect these three mentors had on his career. Niels was also deeply interested in the analytical methods used in evolutionary research. Together with fellow entomologist Nils Møller Andersen and the palaeontologist Niels Bonde, he was a pioneer in Denmark, and more widely in Scandinavia, of Hennig’s phylogenetic systematics, and his cladistic analyses of the higher-level relationships of butterflies in 1976 remained the standard work on the subject until the study by de Jong et al. (1996) 20 years later.

From the very start of his career, Niels was deeply interested in the morphology and phylogeny of the higher insects. In 1975 he published (Z. zool. Syst. Evolut.-forsch. 13, pp. 1–44) one of his most influential papers: “The phylogeny of hexapod ‘orders’. A critical review of recent accounts”. Thirty years later, Grimaldi and Engel (2005, p. 144) referred to this work as “perhaps the single most important paper in systematic entomology”. This publication formed the basis of his five updated reviews of the subject. The last of these was published in Eur. J. Ent. in 1999, while perhaps the most notable of them is the 1991 text-book chapter “Phylogeny of extant hexapods” in “The Insects of Australia”, which should be mandatory reading for all students of systematic entomology. Over the years Niels authored or co-authored a number of papers on higher Hexapod relationships especially on the lower Hexapod orders, Trichoptera, the enigmatic New Zealand mecopteran family Nannochoristidae, Neuroptera, and of course Mantophasmatodea—the first new insect order to be described for 90 years, the description of which he co-authored in 2004.

It was, however, the Lepidoptera that remained Niels’ main interest, and the majority of his publications are on that order. They range in scope from nomenclatural and faunistic notes to higher-level phylogenetics and to the exceptionally detailed, comparative morphological studies of primitive Lepidoptera. More than anything else, these exquisite studies became his professional hallmark. His early enthusiasm for scanning electron microscopy and histology were combined with transmission electron microscopy and became methodological cornerstones in his work throughout his working life. Much of his productivity, particularly in the first half of his career, led to highly detailed studies of little-understood structures and organ systems of primitive Lepidoptera, including overall head and neck anatomy, mouthpart morphology, anatomy of the alimentary canal, structure of the trachaea system, comparative morphology and anatomy of male and female genitalia, and wing scales and vestiture. Niels’ work on primitive Lepidoptera morphology and anatomy was always embedded in the context of higher Lepidoptera evolution, and his ultimate goal was to establish the early evolutionary patterns within the order, thereby creating a sound basis for further studies higher up the lepidopteran tree. In 1978 and 1979 he also described two new families of primitive Lepidoptera, the basal hepialoid family Neotheoridae and the non-ditrysian family Heterobathmiidae (the latter in collaboration with the late Ebbe S. Nielsen). His work on primitive Lepidoptera phylogeny and comparative morphology culminated in his Dr Scient. dissertation “Studies on the morphology and systematics of primitive Lepidoptera” published in Steenstrupia in 1984. Until the modifications introduced by the very recent advent of phylogenomic studies and especially the surprising discovery of a new primitive moth family from Australia, this remained the standard work on the evolution of the homoneurous Lepidoptera.

In the early 1990s Niels was appointed the editor-in-chief of the two Lepidoptera volumes of the Handbook of Zoology. This immense undertaking was to dominate his professional life for the following decade. The two volumes, which were published in 1998 and 2003, defined the latter part of his career as much as his work on higher Hexapod phylogeny and comparative Lepidoptera morphology had shaped his early and mid career, although he continued his work on these topics until illness forced him to stop just weeks before his death. Niels had anticipated writing or co-authoring a substantive part of the first volume. He did not, however, expect to have made a similar input to the second volume, which was on morphology and physiology. Having to do so resulted in a much greater effort on his part than he had intended: moreover, it required him to write about subjects on which he did not consider himself an expert. The result, nevertheless, stands as a landmark publication and a tribute to Niels’ capacity and breadth of knowledge. The Handbook would have been more than enough of a mega-project for most of us, so it is remarkable that Niels also spent much time and effort during his last years editing a book on the insects of Greenland instead of completing some of his own research projects. While he certainly believed in the value of the Greenland work, his resolve was propelled by that innate sense of responsibility and conscientiousness that were so evident in his personal makeup.

After the mammoth task of completing the Handbook, Niels returned to his work on primitive Lepidoptera — at least as much as his administrative duties permitted. At the time of his death, he was involved in long-term studies of several groups: Micropterigidae with G. W. Gibbs and D. L. Lees; Mnesarchaeidae with G. W. Gibbs; Hepialoidea with TJS. A phylogenomic study of the non-ditrysian lineages (part of the LepTree project), of which he was a senior co-author, was submitted just weeks after his death (and was dedicated to him). But his most significant contribution in the last stage of his career was the discovery and description of an extraordinary homoneuran family from Kangaroo Island, Australia (the so-called Kangaroo Island Moth, or KIM) and its significance for modifying our understanding of early Lepidoptera evolution. The paper was, to his great pleasure, accepted for publication before his death.

Besides being a leading research scientist, Niels was a highly engaging and inspiring teacher and supervisor. Early in his career, he wrote, in Danish, a detailed yet concise compendium of systematic entomology (“Systematisk Entomologi” 1974), which for years was the standard textbook on the subject at the University of Copenhagen. For over two decades he was a driving force behind an advanced course in systematic entomology and insect morphology, which was taught biennially at the University. Niels’ lectures displayed not only the depth of his learning, but were rich in subtle humour, a quality of which he was a master. TJS recalls a particular entomology lecture (in 1996) in which Niels was explaining the morphology of the thorax, including flight mechanisms. To ensure that the students understood the complex ways in which insects move their wings to minimize drag, he demonstrated by lying face down on a table, still clad in jacket and tie and with his feet sticking out, waiving his arms in the air! While this performance was not characteristic of most university professors, it worked — TJS resolved there and then to do his graduate studies under Niels’ supervision.

Throughout his career, Niels supervised several Masters and PhD students and postdoctoral fellows. He took a deep interest in their well-being, both professional and personal, and derived immense pleasure from their subsequent successes while keeping in close contact with them after they graduated. (After graduating and moving away from Copenhagen, TJS spent numerous hours on the phone with Niels discussing his own work, Niels’ work, the world in general and entomology in particular.) It was therefore also with great sadness and regret that he found himself writing obituaries for two of his most talented PhD students, the coleopterist Michael Hansen and the lepidopterist Ebbe S. Nielsen, both of whom died prematurely in the year 2000. Niels considered the more sociological aspects of entomology and lepidopterology to be integral parts of the (informal) training of a student. TJS recalls numerous meetings with Niels, intended to be brief, but often extending to a couple of hours, and invariably covering a wide range of aspects such as the history of science, the works (present and past) of other entomologists, anecdotes, amateur entomology, entomology and society. One of Niels’ great qualities as a supervisor was that hallmark of all top supervisors — he had an intuitive understanding of the level and extent of supervision needed to fit the individual student, endeavouring always to bring out the best in him or her.

Niels started collecting butterflies and moths as a schoolboy, and although he never built up a large collection, this activity influenced his choice to become a biologist and a specialist in Lepidoptera. At that time there was no tradition for lepidopterology at the ZMUC, and Niels was the first academically trained lepidopterist at the museum — despite being advised by the head of the entomology department at that time, S. L. Tuxen, to find a more scientific group! Later he often defended collecting Lepidoptera, arguing that is an important way to get young people interested in entomology.

Niels appreciated deeply (serious) amateur lepidopterists, being well aware that major parts of the Lepidoptera collections in larger museums had been collected by them. He valued their efforts, spoke positively and warmly about them and did much to help them, for example through his work for societies with large amateur memberships, by advising on scientific matters and in providing help to get collecting permits.

While Niels enjoyed experiencing Lepidoptera and other insects in nature, and he would often run a mercury vapour light at his summer cottage, he was not primarily a field worker. He felt, and indeed demonstrated, that he could serve the study of Lepidoptera best by focusing his exceptional skills on the study of key taxa at the museum bench.

Besides his research and teaching, Niels shouldered a substantial administrative burden at the ZMUC, at which he spent his entire career. This included several stints as Chair or Deputy Chair of the Entomology Department, two periods as Deputy Director of the museum, and three years as Director. He was also Head of Zoology at the newly designated Natural History Museum of Denmark from 2004–2006. Although his heart remained in his research, he carried out these time-consuming administrative responsibilities conscientiously and with a great sense of love and concern for the museum.

Throughout his career, Niels was deeply involved in entomological and lepidopterological societies. He was President of the Danish Entomological Society from 1989 to 1999, Council member of the International Congress of Entomology from 1988 to 2004 (Deputy Chair 2000–2004), and of course President of the SEL from 1998 to 2007, and Chair of the SEL congress in Korsør, Denmark in 2002. One of Niels’ long-standing ambitions was to hold a joint European-North American Lepidoptera Congress. Although this did not take place during his own tenure as President, he was very pleased to see the first joint meeting of the Lepidopterists’ Society and SEL in Denver, Colorado in 2012 (even if he could not attend the meeting himself).

During his career, Niels received many honours and awards, a testimony to his achievements and pre-eminence in his field. He was a member of the Danish Academy of Natural Sciences, a member of the Royal Danish Academy of Sciences and Letters, a corresponding member of the Finnish Entomological Society, an honorary ‘Foreign member’ of the Linnean Society of London (1998), an honorary member of Sociedad Hispano-Luso-Americana de Lepidopterologia (SHILAP), an honorary member of the Danish Entomological Society, an honorary research fellow at the Natural History Museum, London, an honorary member of Gesellschaft für Biologische Systematik, an honorary member of Societas Europaea Lepidopterologica, an honorary fellow of the Royal Entomological Society, and an honorary member of the Russian Entomological Society. In 1988 he was awarded the ‘Karl Jordan Medal’ (Lepidopterists’ Society) for “outstanding original research in lepidopterology”, in 1999 he received the Joachim Jungius-Medaille (J.J.Gesellschaft der Wissenschaften, Hamburg) for “herausragender Leistungen in Wissenschaft und Forschung”, and in 2014 shortly before his death he gained the 2014 Linnean Medal (Zoology).

Niels cared deeply about the future of European entomology and lepidopterology and was dismayed by the progressive decline of staff numbers and funds at several major research institutions (including his own in Denmark). He believed firmly in the need for basic research, and that in publicly funded institutions such as museums it should be possible for researchers to focus on academically interesting questions that do not necessarily have immediately obvious economic, social or medical benefits.

Unsurprisingly, Niels’ standing, awareness and understanding of wider socio-political issues in science led to him being asked to act as Director of the ZMUC. While he had an interest in university politics and administration, his real love was for his research, but he accepted the position (doing two terms of service) partly from a sense of duty, and partly because he felt that the position should be held by an acknowledged researcher rather than a mandarin. Having high ethical standards, Niels rarely sought the easiest solution to any problem, but rather the one that he thought to be right. He was a conscientious leader, made great efforts to keep abreast of relevant matters and always made time for his colleagues, whatever their level in the organization. He undoubtedly suffered during his extended directorship, both as a result of these personal qualities and through the loss of most of his time for research. Alas, he had to endure more frustration due to seemingly endless cuts to the museum’s funds. But when the Ministry of Education and Research ordered cuts of several positions at the museum, he felt he could no longer accept the responsibility for running the institution and stepped down in protest.

Niels was critical of the plans for a new natural history museum in Copenhagen, which would have resulted in newer but reduced facilities. He was particularly disturbed about the idea of demolishing the ZMUC building, which had been purpose built and which he considered to be still fit for purpose. During his later years he spoke and wrote against the idea, and was disappointed that the management of the Natural History Museum of Denmark, and many of his colleagues, did not agree with him. He also expressed concern about the appointment at the museum of scientists with little experience of collections-based research.

At the time of his death, Niels had settled into a productive retirement: relieved of administrative responsibilities, it was a phase of his life that he was enjoying thoroughly. So it is heart-breaking that he missed the prolonged and active retirement he would have found so fulfilling. Moreover, it leaves the scientific community bereft of the many works that would surely have been produced by him. The entomological world has become a much poorer place without Niels’ profound knowledge and insight, his generosity of spirit, his conscientiousness and his quiet humour. For all these qualities he is and will continue to be missed deeply. He was also a loving family man and our deepest condolences are extended to his wife Else and their daughters.

HAVAMAL (Our translation)

Livestock die

Kinsman dies

We all die just the same

Only one thing I know which never dies

The judgment of a dead man’s life.

HAVAMAL (Danish)

Fæ dør

Frænde dør

Dør selv på samme vis

Kun et ved jeg som aldrig dør

Dommen over død mands liv

We wish to thank Lars Vilhelmsen, Natural History Museum of Denmark for sharing biographical data he had collected. We thank Birgit Nielsen, Frederiksværk, Denmark for permission to use the portrait photo of Niels P. Kristensen.