In Memoriam: Niels Peder Kristensen (1943–2014)

2015·11 MB·Norwegian

by  Thomas Simonsen

NotaLepi.38(1)2015:89-102 DOI 10.3897/nl.38.4816
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In Memoriam: Niels Peder Kristensen (1943-2014)
Thomas J. Simonsen13 Ole Karsholt2 Malcolm J. Scoble1
, ,
1 NaturalHistoryMuseum, UnitedKingdom,London, UK
2 TheNaturalHistoryMuseumofDenmark, Copenhagen,Denmark
3 Currentaddress:NaturalHistoryMuseumAarhus,Aarhus,Denmark
http://zoobank.org/F762FCAl-0D5A-4CDF-9EB3-93A7800752FD
Received3March2015;accepted 18March2015;published: 12May2015
SubjectEditor:JadrankaRota.
Niels Peder Kristensen, Honorary Member and
former president of SEL, passed away on Satur-
day December 6th 2014 in Copenhagen. While his
death was not unexpected, its timing came earli-
er than we had thought or hoped. His loss is felt
widely and intensely.
Bom on March 2nd 1943, Niels was the second
child of Thorkil and Ellen Christine Kristensen
(nee Nielsen). His father was an academic, politi-
cian andthinkerwho servedas MinsterofFinance
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 entomolo-
gy and lepidopterology in particular at an ear-
Figure 1.Niels PederKristensen, March 2nd 1943 ly age, and once told TJS about the first time,
-December6th2014(photo: BirgitNielsen). when eight years old, he visited the Entomology
Department at the Zoological Museum in Copen-
hagen (ZMUC) ‘clutching his father’s hand’. After completing high school at Birkerod Statsskole
in 1961, Niels enrolled as a biology student at the University ofCopenhagen, and quickly became
a regular visitorto the Entomology Department ofthe 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,whichwasonthe faunisticsofDanishcicadas. Fromtheverystartofhis scientificcareer, one
ofNiels’ abiding interests was the evolution (particularly evolutionary morphology) ofprimitive
Lepidoptera. Indeed, the work for his Mag. Scient, degree was on the comparative morphology
ofthe primitive glossatan family, Eriocraniidae. During this study, Niels spent the academic year
1966-67 atthe University ofBristol, workingwiththe eminentand extremelyknowledgeable Brit-
90 SiMONSENetal: InMemoriam:NielsPederKristensen(1943-2014)
ish entomologist Howard E. Hinton, who was at that time pioneering the use ofscanning electron
microscopy in entomology. It was while working with Hinton that Niels came to appreciate the
value ofSEM in comparativemorphology. Through it, hemadethe surprising discovery(published
in 1970) thatthe mostprimitive Lepidopterahave flat, solidwing scales (i.e. lacking an internal lu-
men), acondition contrasting stronglywiththe hollowwing scales generallyfound inLepidoptera.
After obtaining his Mag. Scient, degree in 1968, Niels was offered a tenure-track position at
the ZMUC asAmanuensis (AssistantProfessor). He waspromotedtoAssociate Professor in 1972
and appointed as Full Professor ofEntomology in 1995. In 1970, Niels visited one ofEurope’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 ofcomparative morphology at
the University ofCopenhagen, unquestionably had a profound influence on Niels’ development
as a scientist. He oftenreferred to the effectthese three mentors had on his career. Niels was also
deeply interested in the analytical methods used in evolutionary research. Together with fellow
entomologistNils MöllerAndersenandthepalaeontologistNiels Bonde, hewas apioneerinDen-
mark, and more widely in Scandinavia, ofHennig’s phylogenetic systematics, and his cladistic
analyses ofthe higher-level relationships ofbutterflies in 1976 remainedthe standardworkonthe
subjectuntil the studyby de Jong etal. (1996) 20 years later.
From the very start ofhis career, Niels was deeply interested in the morphology and phylogeny
ofthe higher insects. In 1975 he published (Z. zool. Syst. Evolut.-forsch. 13, pp. 1^14) one ofhis
most influential papers: “The phylogeny ofhexapod ‘orders’. Acritical review ofrecent accounts”.
Thirtyyearslater,GrimaldiandEngel(2005,p. 144)referredtothisworkas“perhapsthesinglemost
important paper in systematic entomology”. This publication formed the basis ofhis five updated
reviewsofthesubject. ThelastofthesewaspublishedinEur. J. Ent. in 1999,whileperhapsthemost
notable ofthem is the 1991 text-book chapter “Phylogeny ofextant hexapods” in “The Insects of
Australia”, which should be mandatory reading for all students ofsystematic entomology. Overthe
yearsNiels authored orco-authoredanumberofpapers on higherHexapodrelationships especially
on the lower Hexapod orders, Trichoptera, the enigmatic New Zealand mecopteran familyNanno-
—
choristidae,Neuroptera,andofcourseMantophasmatodea thefirstnewinsectordertobedescribed
for90years, the descriptionofwhichhe co-authoredin2004.
Itwas, however, the LepidopterathatremainedNiels’main interest, andthe majorityofhis publi-
cations are on that order. They range in scope from nomenclatural and faunistic notes to higher-lev-
el phylogenetics and to the exceptionally detailed, comparative morphological studies ofprimitive
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
electronmicroscopyandbecamemethodologicalcornerstonesinhisworkthroughouthisworkinglife.
Muchofhisproductivity,particularlyinthefirsthalfofhiscareer, ledtohighlydetailedstudiesoflit-
tle-understoodstructuresandorgansystemsofprimitiveLepidoptera,includingoverallheadandneck
anatomy, mouthpart morphology, anatomy ofthe alimentary canal, structure ofthe trachaea system,
comparative morphology and anatomy ofmale and female genitalia, and wing scales and vestiture.
Niels’workonprimitive Lepidopteramorphologyandanatomywas always embedded inthe context
ofhigherLepidopteraevolution, andhis ultimategoalwasto establishtheearlyevolutionarypatterns
within the order, thereby creating a soundbasis forfurther studies higherup the lepidopteran tree. In
1978and 1979healsodescribedtwonewfamiliesofprimitiveLepidoptera,thebasalhepialoidfamily
NotaLepi.38(1):89-102 91
Figure2.NielsstudyinghisbelovedhomoneuranLepidoptera(photo: TJS).
Neotheoridae andthe non-ditrysian familyHeterobathmiidae (the latterin collaborationwiththe late
Ebbe S.Nielsen). HisworkonprimitiveLepidopteraphylogenyandcomparativemorphologyculmi-
natedinhisDrScient, dissertation“StudiesonthemorphologyandsystematicsofprimitiveLepidop-
tera”published in Steenstrupia in 1984. Until the modifications introducedbytheveryrecent advent
ofphylogenomic studiesandespeciallythe surprisingdiscoveryofanewprimitivemoth familyfrom
Australia,thisremainedthe standardworkontheevolutionofthehomoneurousLepidoptera.
In the early 1990s Niels was appointed the editor-in-chiefofthe two Lepidoptera volumes of
the Handbook ofZoology. This immense undertaking was to dominate his professional life forthe
followingdecade. Thetwovolumes,whichwerepublishedin 1998 and2003, definedthe latterpart
ofhiscareerasmuchashisworkonhigherHexapodphylogenyandcomparativeLepidopteramor-
phology had shaped his early and mid career, although he continued his work on these topics until
illness forcedhimto stopjustweeksbefore his death. Nielshadanticipatedwritingorco-authoring
a substantive part ofthe first volume. He did not, however, expect to have made a similar input to
the secondvolume, whichwas on morphologyandphysiology. Having to do so resulted in amuch
greaterefforton his partthan hehad intended: moreover, itrequired himtowrite about subjects on
which he did not consider himselfan expert. The result, nevertheless, stands as a landmark publi-
cation and a tribute to Niels’ capacity and breadth ofknowledge. The Handbook would have been
more than enough ofa mega-project formost ofus, so it is remarkable thatNiels also spent much
timeandeffortduringhislastyearseditingabookontheinsectsofGreenlandinsteadofcompleting
some ofhis own research projects. While he certainlybelieved in thevalue ofthe Greenlandwork,
his resolve was propelled by that innate sense ofresponsibility and conscientiousness that were so
evident in hispersonal makeup.
92 SiMONSENetal:InMemoriam:NielsPederKristensen(1943-2014)
Afterthe m—ammoth task ofcompleting the Handbook, Niels returned to his work on primitive
Lepidoptera at least as much as his administrative duties permitted. At the time ofhis death,
he was involved in long-term studies ofseveral groups: Micropterigidae with G. W. Gibbs andD.
L. Lees; Mnesarchaeidae with G. W. Gibbs; Hepialoidea with TJS. Aphylogenomic study ofthe
non-ditrysian lineages (part ofthe LepTree project), ofwhich he was a senior co-author, was sub-
mittedjustweeks afterhis death (andwas dedicatedto him). Buthis most significant contribution
in the last stage ofhis career was the discovery and description ofan extraordinary homoneuran
family from Kangaroo Island, Australia (the so-called Kangaroo Island Moth, or KIM) and its
significanceformodifyingourunderstandingofearlyLepidopteraevolution. Thepaperwas,tohis
greatpleasure, accepted forpublicationbefore 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 foryears was the standard text-
bookonthe subjectattheUniversity ofCopenhagen. Forovertwo decades hewas adriving 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 ofhis learning, but were
rich in subtle humour, a quality ofwhich he was a master. TJS recalls a particular entomology
lecture (in 1996) in which Niels was explaining the morphology ofthe thorax, including flight
mechanisms. To ensurethatthe studentsunderstoodthe complexways inwhichinsectsmovetheir
wings to minimize drag, he demonstratedby lying face downon atable, still clad injacket andtie
andwithhis feet sticking out, waivinghis arms i—nthe air! Whilethis performancewas notcharac-
teristic ofmostuniversityprofessors, itworked TJS resolvedthere andthen to do his graduate
studies underNiels’ supervision.
Throughout his career, Niels supervised several Masters and PhD students and postdoctoral fel-
lows. He took a deep interest in their well-being, both professional and personal, and derived im-
mensepleasure from theirsubsequent successes while keeping in close contactwiththem afterthey
graduated. (After graduating and moving away from Copenhagen, TJS spent numerous hours on
thephonewithNiels discussinghis ownwork,Niels’work, theworld in general andentomology in
particular.)Itwasthereforealsowithgreatsadnessandregretthathefoundhimselfwritingobituaries
fortwoofhismosttalentedPhDstudents,thecoleopteristMichaelHansenandthelepidopteristEbbe
S.Nielsen, bothofwhomdiedprematurelyintheyear2000. Niels consideredthemore sociological
aspectsofentomologyandlepidopterologytobeintegralpartsofthe(informal)trainingofastudent.
TJS recalls numerous meetings with Niels, intended to be brief, but often extending to a couple of
hours,andinvariablycoveringawiderangeofaspectssuchasthehistoryofscience,theworks(pres-
ent and past) ofother entomologists, anecdotes, amateur entomology, entomol—ogy and society. One
ofNiels’greatqualitiesas asupervisorwasthathallmarkofalltopsupervisors hehadanintuitive
understandingofthelevelandextentofsupervisionneededtofittheindividualstudent,endeavouring
alwaystobring outthebestinhim orher.
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 Lep-
idoptera. At that time there was no tradition for lepido—pterology at the ZMUC, and Niels was the
first academically trained lepidopterist atthe museum despite being advised bythe head ofthe
entomology department at that time, S. L. Tuxen, to find a more scientific group! Later he often
NotaLepi.38(1):89-102 93
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 majorparts of
theLepidopteracollections inlargermuseums hadbeencollectedbythem. Hevaluedtheirefforts,
spokepositivelyandwarmlyaboutthemand didmuchtohelpthem, forexamplethroughhiswork
for societies with large amateur memberships, by advising on scientific matters and in providing
help to get collectingpermits.
While Niels enjoyed experiencing Lepidoptera and other insects in nature, and he would often
run amercuryvapourlightathis summercottage, hewasnotprimarilyafieldworker. He felt, and
indeeddemonstrated,thathe couldservethe studyofLepidopterabestbyfocusinghisexceptional
skills onthe study ofkeytaxa atthe museumbench.
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
ofthe EntomologyDepartment,twoperiods as DeputyDirectorofthemuseum, andthreeyears as
Director. He was also Head ofZoology atthe newly designatedNatural HistoryMuseum ofDen-
markfrom 2004-2006.Althoughhisheartremained inhisresearch, he carriedoutthesetime-con-
suming administrative responsibilities conscientiously and with a great sense oflove and concern
forthe museum.
Throughouthis career,Nielswas deeplyinvolvedin entomological andlepidopterological soci-
eties. Hewas Presidentofthe DanishEntomological Society from 1989to 1999, Council member
oftheInternationalCongressofEntomologyfrom 1988to2004(DeputyChair2000-2004),andof
coursePresidentofthe SELfrom 1998to2007,andChairofthe SELcongressinKorsor,Denmark
in2002. OneofNiels’long-standingambitionswastoholdajointEuropean-NorthAmericanLepi-
dopteraCongress.Althoughthisdidnottakeplace duringhis owntenureasPresident,hewasvery
pleasedto seethefirstjointmeetingoftheLepidopterists’ Societyand SELinDenver, Colorado in
2012 (even ifhe could not attendthe meetinghimself).
Duringhis career,Nielsreceivedmanyhonours andawards, atestimonytohis achievements and
pre-eminence inhisfield. Hewas amemberofthe DanishAcademyofNatural Sciences, amember
ofthe Royal DanishAcademy ofSciences and Letters, a corresponding memberofthe Finnish En-
tomological Society, an honorary ‘Foreign member’ ofthe Linnean Society ofLondon (1998), an
honorarymemberofSociedadHispano-Luso-Americanade Lepidopterologia(SHILAP), an honor-
arymemberoftheDanishEntomological Society,anhonoraryresearchfellowattheNaturalHistory
Museum, London, an honorary member ofGesellschaft für Biologische Systematik, an honorary
member ofSocietas Europaea Lepidopterologica, an honorary fellow ofthe Royal Entomological
Society,andanhonorarymemberoftheRussianEntomological Society. In 1988hewasawardedthe
‘KarlJordanMedal’(Lepidopterists’ Society) for“outstandingoriginalresearchinlepidopterology”,
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
deathhe gainedthe 2014LinneanMedal (Zoology).
Niels cared deeply about the future ofEuropean entomology and lepidopterology and was dis-
mayedbytheprogressive decline ofstaffnumbers and funds at several majorresearch 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
94 SiMONSENetal.\InMemoriam:NielsPederKristensen(1943-2014)
academically interesting questions that do not necessarily have immediately obvious economic,
social ormedical benefits.
Unsurprisingly, Niels’ standing, awareness and understanding ofwider socio-political issues in
science led to him being asked to act as Director ofthe ZMUC. While he had an interest in uni-
versitypolitics and administration, his real love was forhis research, but he acceptedthe position
(doingtwoterms ofservice)partlyfromasense ofduty, andpartlybecausehefeltthattheposition
shouldbe held by an acknowledged researcherratherthan amandarin. Having high ethical stand-
ards, Niels rarely sought the easiest solution to anyproblem, but ratherthe one that he thought to
be right. He was a conscientious leader, made great efforts to keep abreast ofrelevantmatters and
always made time for his colleagues, whatever their level in the organization. He undoubtedly
suffered during his extended directorship, both as a result ofthese personal qualities and through
the loss ofmost ofhis time forresearch.Alas, he had to endure more frustration due to seemingly
endlesscutstothemuseum’sfunds. ButwhentheMinistryofEducationandResearchorderedcuts
ofseveralpositions atthe museum, he felthe couldno longeraccepttheresponsibilityforrunning
the institution and stepped down inprotest.
Niels was critical ofthe 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 ofde-
molishingtheZMUCbuilding,whichhadbeenpurposebuiltandwhichheconsideredtobe stillfit
forpurpose. During his lateryears he spoke andwrote againstthe idea, andwas disappointedthat
the management oftheNatural HistoryMuseum ofDenmark, andmany ofhis colleagues, didnot
agreewithhim. Healso expressedconcernaboutthe appointmentatthe museumofscientistswith
little experience ofcollections-based research.
Atthetime ofhis death,Nielshad settled into aproductiveretirement: relieved ofadministrative
responsibilities, itwasaphaseofhislifethathewas enjoyingthoroughly. So itisheart-breakingthat
hemissedtheprolongedandactiveretirementhewouldhave foundso fulfilling. Moreover, itleaves
thescientificcommunitybereftofthemanyworksthatwouldsurelyhavebeenproducedbyhim.The
entomological world has become a much poorer place without Niels’ profound knowledge and in-
sight,hisgenerosityofspirit, his conscientiousnessandhis quiethumour. Forallthesequalitieshe is
andwillcontinuetobemisseddeeply. Hewasalsoalovingfamilymanandourdeepestcondolences
are extendedtohiswifeElse andtheirdaughters.
HAVAMAL
(Ourtranslation)
Livestockdie
Kinsman dies
We all diejustthe same
Only one thing I knowwhichneverdies
Thejudgment ofadead man’s life.
HAVAMAL
(Danish)
Fæ dor
Frænde dor
Dor selvpâ samme vis
Kun etvedjeg som aldrig dor
Dommen overdod mands liv
.
NotaLepi.38(1):89-102 95
Acknowledgements
WewishtothankLarsVilhelmsen,NaturalHistoryMuseumofDenmarkforsharingbiographicaldatahehadcollected.We
thankBirgitNielsen,Frederiksværk,DenmarkforpermissiontousetheportraitphotoofNielsP.Kristensen.
References
deJongR,Vane-WrightRI,AckeryPR(1996)Thehigherclassificationofbutterflies(Lepidoptera):problems
andprospects. Entomologicascandinavica27: 65-101. doi: 10.1163/187631296X00205
GrimaldiD,EngelMS(2005)EvolutionoftheInsects.CambridgeUniversityPress,Cambridge,UK,755pp.
Niels PederKristensen, publications:
M
156.KarsholtO,KristensenNP, SimonsenTJ,Ahola (inpress)Lepidoptera(mothsandbutterflies). 51 pp.
In Böcher J, Kristensen NP, Pape T, Vilhelmsen L (Eds). The Greenlandentomofauna. An identification
manualofinsects, spidersandtheirallies. E. J. Brill, Leiden.
155. Kristensen NP, Hilton DJ, KalliesA, Milia L, Rota J, Wahlberg N, Wilcox SA, Glatz RV, Young DA,
Cocking G, Edwards ED, Gibbs GW, Halsey M (2015) A new extant family ofprimitive moths from
KangarooIsland,Australia,anditssignificanceforunderstandingearlyLepidopteraevolution. Systematic
Entomology40: 5-16. doi: 10.1111/syen.12115
154.KristensenNP,RotaJ,FischerS (2014)Notableplesiomorphiesandnotablespecializations:headstruc-
ture ofthe primitive “tongue moth” Acanthopteroctetes unifascia (Lepidoptera: Acanthopteroctetidae).
Journal ofMorphology275: 153-172. doi: 10.1002/jmor.20205
153. KristensenNP (2013) Intraspecific variability in gross design ofmothbrains:Acaveat concerning ex-
pectedly ‘good’characters’ in systematic entomology(Insecta: Lepidoptera). ZoologischerAnzeiger253:
112-113. doi: 10.1016/j.jcz.2013.09.002
152. EngelMS,KristensenNP(2013)Ahistoryofentomologicalclassification.AnnualReviewofEntomol-
ogy58: 585-607. doi: 10.1146/annurev-ento-120811-153536
151. Hünefeld F, Kristensen NP (2012) The female postabdomen and genitalia ofthe basal moth family
&
Heterobathmiidae(Insecta: Lepidoptera): Structureandphylogenetic significance.Arthropod Structure
Development41: 395^107. doi: 10.1016/j.asd.2012.05.001
150. Beutel RG, KristensenNP(2012) Morphologyand insect systematics inthe era ofphylogenomics.Ar-
thropodStructure& Development41: 303-305. doi: 10.1016/j.asd.2012.05.003
149.HünefeldF.KristensenNP(2012)Twonewheterobathmiidmothspecieswithdistinctivefemalegenital
configurations(Lepidoptera: Heterobathmiidae). Zootaxa3281: 61-68.
148. Kristensen NP (2012) Molecular phylogenies, morphological homologies and the evolution ofmoth
‘ears’. SystematicEntomology37: 237-239. doi: 10.111l/j.1365-3113.2012.00619.x
147. Gibbs GW, KristensenNP (2011) Agrionympha, the long-known SouthAfricanjaw moths: a revision
withdescriptions ofnewspecies(Lepidoptera,Micropterigidae). Zootaxa2764: 1-21
146. Djemæs M, Kristensen NP (2011) Derived morphology in a basal moth: The uniquely specialized
sternum V glands ofAgathiphaga. Arthropod Structure & Development 40: 559-569. doi: 10.1016/j.
asd.2011.06.001
145. RotaJ, KristensenNP (2011) Notes ontaxonomic history, thoraco-abdominal articulation, and current
placementofMillieriidae(Insecta: Lepidoptera). Zootaxa3032: 65-68.
144.vanNieukerkenE,KailaL,KitchingIJ,KristensenNP,Lees,DC,MinetJ,MitterC,MutanenM,Regier
JC, SimonsenTJ,WahlbergN,YenS-H,ZahiriR, etal. (38 additional authors)(2011)OrderLepidoptera.
96 SiMONSENetal: InMemoriam:NielsPederKristensen(1943-2014)
In: Zhang Z-Q (Ed.)Animal biodiversity: An outline ofhigher-level classification and surveyoftaxono-
micrichness. Zootaxa3148: 212-221.
143.KristensenNP(2011)MichaelFibiger29.juni 1945- 16. februar2011.EntomologiskeMeddelelser79:
153-165.
142. Hünefeld F, Kristensen NP (2010) The female postabdomen and internal genitalia ofthe basal moth
genusAgathiphaga (Insecta: Lepidoptera: Agathiphagidae): Morphology and phylogenetic implications.
ZoologicalJournaloftheLinneanSociety 159: 905-920. doi: 10.1111/j.1096-3642.2009.00590.x
141. KristensenNP, Gaedike R(2010) Extraordinarymoths andan extraordinarymothresearcher:An essay
review ofG. S. Robinson’s Biology, distribution and diversity oftineid moths. Nota Lepidopterologica
33: 3-8.
140. Lees DC, Rougerie R, Zeller-Lukaschort C, Kristensen NP (2010) DNA mini-barcodes in taxonomic
assignment: amorphologicallyuniquenewhomoneurousmothcladefromtheIndianHimalayasdescribed
in Micropterix (Lepidoptera, Micropterigidae). Zoologica Scripta 39: 642-661. doi: 10.1111/j.1463-
6409.2010.00447.x
139. Kristensen N P, Nielsen PS (2010) Et Hoffmeyer-manuscript om vore ’stribede’ kollesværmere. Lepi-
doptera9: 289-307.
138. Beutel RG, Kristensen NP, Pohl H (2009) Resolving insect phylogeny: The significance ofcephalic
structures oftheNannomecopterainunderstanding endopterygoterelationships.Arthropod Structure and
Devopment38: 427^160. doi: 10.1016/j.asd.2009.05.002
137.KaaberS,KristensenNP, SimonsenTJ(2009)Sexualdimorphismandgeographicalmalepolymorphism
in the ghost moth Hepialus humuli (Lepidoptera: Hepialidae): Scale ultrastructure and evolutionary as-
pects. EuropeanJournal ofEntomology 106: 303-313. doi: 10.14411/eje.2009.036
M
136. Béthoux O, KristensenNP, Engel (2008) Hennigian phylogenetic systematics and the ‘groundplan’
vs. ‘post-groundplan’ approaches: A Reply to Kukalovâ-Peck. Evolutionary Biology 35: 317-323. doi:
10.1007/sl1692-008-9035-6
135.KristensenNP(2008)Natsværmeren ‘humleæder’,densflyveaktivitet,vingeskæl-strukturogvariationI
Nordeuropa. DyriNaturogMuseum2008/2: 22-25.
134. KristensenNP(2008)EarlyLepidopteraevolution. GesellschaftfürBiologische SystematikNewsletter
20: 50-55.
133.KristensenNP,ScobleMJ,KarsholtO(2007)Lepidopteraphylogenyandsystematics:thestateofinven-
toryingmothandbutterflydiversity. Zootaxa 1668: 699-747.
132 KristensenNP(2007)NilsMöllerAndersen. DanskNaturhistoriskForeningÂrsskrift 16/17: 50-51.
.
131.KristensenNP(2007)LeifLyneborg-3.januar 1932- 10. September2006.DanskNaturhistoriskForen-
ingsÂrsskrift 16/17: 68-75.
130. Faucheux MJ, Kristensen NP, Yen S-H (2006) The antennae ofneopseustid moths: Morphology and
phylogenetic implications,withspecialreferencetothe sensilla(Insecta,Lepidoptera,Neopseustidae). Zo-
ologischerAnzeiger245: 131-142. doi: 10.1016/j.jcz.2006.05.004
129 KristensenNP(2006)Nils MöllerAndersen21. november 1940- 12. maj 2004. DetKongelige danske
.
videnskabemes Selskab. OversigtoverSelskabetsVirksomhed2004-05: 217-224.
128. Kristensen NP (2005) Jens Bodtker Rasmussen - Obituary. Tropical Zoology 18: 149-149. doi:
10.1080/03946975.2005 10531217
.
127.KrennHW, KristensenNP(2004)EvolutionofproboscismusculatureinLepidoptera. EuropeanJournal
ofEntomology 101: 565-575. doi: 10.1441l/eje.2004.080
126. KristensenNP (2004) Om indsamling ogudskillelse. NoglenaturhistoriskePerspektiven DanskeMuseer
17/5:20-21.
125. Kristensen NP (2004) Paul Johannes Holst-Christensen. Dansk Naturhistorisk Forening Ârsskrift 14:
86-89.
NotaLepi.38(1):89-102 97
124. KristensenNP(2003)ResolvingthebasalphylogenyofLepidoptera: morphologicalevidence. Entomo-
logischeAbhandlungen61:167-169.
123. KristensenNP (2003) Reproductive organs. Pp. 427-447 inN.P.Kristensen (Ed.) Lepidoptera: Moths and
butterflies2.HandbuchderZoologie/HandbookofZoologyIV/36:WalterdeGruyter,Berlin&NewYork.
122.Akai H, Hakim RS, KristensenNP(2003) Labial glands, silkandsaliva. Pp. 377-388 inN.P.Kristensen
(ed.) Lepidoptera: Moths andbutterflies 2. Handbuch derZoologie/HandbookofZoologyIV/36: Walterde
Gruyter,Berlin&NewYork.
121. Warrant E, KelberA, KristensenNP (2003) Eyes and vision. Pp. 325-359 inN.P.Kristensen (ed.) Lep-
idoptera: Moths andbutterflies 2. Handbuch derZoologie/HandbookofZoology IV/36: Walterde Gruyter,
Berlin&NewYork.
120. Barbehenn RV, Kristensen NP (2003) Digestive and excretory system. Pp. 165-187 in N.P.Kristensen
(ed.) Lepidoptera: Moths andbutterflies 2. Handbuch derZoologie/HandbookofZoology'IV/36: Walterde
Gruyter,Berlin&NewYork.
119. Hasenfuss I, Kristensen NP (2003) Skeleton and muscles: immatures. Pp. 133-164 in N.P.Kristensen
(ed.) Lepidoptera: Moths andbutterflies 2. Handbuch derZoologie/HandbookofZoology IV/36: Walterde
Gruyter,Berlin&NewYork.
118. Kristensen NP (2003) Skeleton and muscles: adults. Pp. 39-131 in N.P.Kristensen (ed.) Lepidoptera:
Moths andbutterflies 2. Handbuch derZoologie/HandbookofZoology IV/36: Walterde Gruyter, Berlin &
NewYork.
117. Kristensen NP, Simonsen TJ (2003) ‘Hairs’ and scales. Pp. 9-22 in N.P.Kristensen (ed.) Lepidoptera:
Moths andbutterflies 2. Handbuch derZoologie/HandbookofZoology IV/36: Walter de Gruyter, Berlin &
NewYork.
116. Chauvin G, Kristensen NP (2003) Integument. Pp. 1-8 in N.P.Kristensen (ed.) Lepidoptera: Moths and
butterflies2. HandbuchderZoologie/HandbookofZoologyIV/36:WalterdeGruyter,Berlin&NewYork.
115.KristensenNP, SimonsenTJ(2003) Scalelength/winglengthcorrelationinLepidoptera.JournalofNat-
uralHistory37: 673-679. doi: 10.1080/00222930110096735
114. Karsholt O, Kristensen NP (2003) Plesiozela , gen. nov. from temperate SouthAmerica: apparent sis-
ter-group ofthepreviouslyknown Heliozelidae (Lepidoptera: Incurvarioidea: Heliozelidae). Invertebrate
Systematics 17: 39-46. doi: 10.1071/IS02047
113. KarsholtO, KristensenNP(2003) Kastaniemollet: etkontnyt skadedyriDanmark. DyriNaturogMu-
seum2003/1: 9-11.
112. Kristensen NP (2002) Mantophasmatodea: en nyopdaget orden afnulevende insekter. Dyr i Natur og
Museum2002/2: 24-27.
111.KlassK-D,ZomproO,KristensenNP,AdisJ(2002)Mantophasmatodea: anew insectorderwithextant
members intheAfrotropics. Science206: 1456-1459. doi: 10.1126/science.1069397
110.KristensenNP(2002)EbbeSchmidtNielsen. DanskNaturhistoriskForeningÂrsskrift 12: 58-60.
109.KristensenNP(1994—2002)Contributionsto ‘DanmarksNationalleksikonYDenStoreDanskeEncyklopce-
di’(vols 1-20, Gyldendal, Copenhagen), including ‘Dagsommerfugle’ [butterflies], ‘Insekter’ [insects], ‘Led-
dyr’ [arthropods], ‘Sommerfugle’ [Lepidoptera], ‘Urinsekter’ [apterygotes] andnumerous shorter(<50 lines)
articlesonzoological(mostlyentomological)subjects,includingbiographiesofzoologists.
108. Simonsen TJ, Kristensen NP (2001) Agathiphaga wing vestiture revisited: evidence for complex ear-
ly evolution oflepidopteran scales (Lepidoptera: Agathiphagidae). Insect Systematics and Evolution 32:
169-175. doi: 10.1163/187631201X00128
107. KlassK-D, KristensenNP(2001)Thegroundplanandaffinities ofhexapods: recentprogress andopen
problems.Annalesdela SociétéEntomologiquedeFrance(N. S.)37: 265-298.
106.KristensenNP(2001)EbbeSchmidtNielsen7June 1950-6March2001.NotaLepidopterologica24/3:
3-9.
98 SiMONSENetal:InMemoriam:NielsPederKristensen(1943-2014)
105.KristensenNP(2001)HenningAnthon. EntomologiskeMeddelelser69: 65-68.
104.KristensenNP(2001)MichaelHansen. DanskNaturhistoriskForeningÂrsskrift 11: 67-69.
103.KrennHW,KristensenNP(2000)EarlyevolutionoftheproboscisofLepidoptera(Insecta):externalmor-
phologyofthegaleainbasalglossatanmothlineages,withremarksontheoriginofthepilifers.Zoologisher
Anzeiger239: 179-196.
102. KristensenNP(1999) Phylogenyofendopterygote insects, themostsuccessful lineage ofliving organ-
isms. EuropeanJournalofEntomology96: 237-253.
101.DugdaleJS,KristensenNP,Robinson,GS,ScobleMJ(1998)Thesmallermicrolepidoptera-gradesuper-
families. Pp. 217-232 inN.P.Kristensen(ed.)Lepidoptera:Mothsandbutterflies 1.HandbuchderZoologie/
HandbookofZoologyIV/35(1999): 51-63.WalterdeGruyter,Berlin&NewYork.
100. Edwards ED, Gentili P, HorakM, KristensenNP,NielsenES (1998) The Cossoidea and Sesioidea. Pp.
181-197 inN.P.Kristensen(ed.)Lepidoptera:Mothsandbutterflies 1.HandbuchderZoologie/Handbookof
ZoologyIV/35: (1999)51-63.WalterdeGruyter,Berlin&NewYork.
99. Dugdale JS, Kristensen NP, Robinson GS, Scoble MJ (1998) The Yponomeutoidea. Pp. 119-130 in
N.P.Kristensen (ed.) Lepidoptera: Moths and butterflies 1. Handbuch der Zoologie/Handbook ofZoology
IV/35(1999): 51-63.WalterdeGruyter,Berlin&NewYork.
98. KristensenNP(1998) The homoneurous Glossata. Pp. 51-63 inN.P.Kristensen(Ed.)Lepidoptera: Moths
andbutterflies 1.HandbuchderZoologie/HandbookofZoologyIV/35(1999): 51-63.WalterdeGruyter,Ber-
lin&NewYork.
97.KristensenNP(1998)Thenon-glossatanmoths. Pp.41^49inN.P.Kristensen(Ed.)Lepidoptera:Mothsand
butterflies 1. Handbuch derZoologie/Handbook ofZoology IV/35: 51-63(1999). Walterde Gruyter, Berlin
&NewYork.
96. CarterD, KristensenNP(1998) Classification andkeysto highertaxa. Pp. 27—40 inN.P.Kristensen(ed.)
Lepidoptera: Moths and butterflies 1. Handbuch der Zoologie/Handbook ofZoology IV/35(1999): 51-63.
WalterdeGruyter,Berlin&NewYork.
95.KristensenNP, SkalskiA(1998)Palaeontologyandphylogeny. Pp. 7-25 inN.P.Kristensen(ed.)Lepidop-
tera: Moths andbutterflies 1. HandbuchderZoologie/HandbookofZoologyIV/35(1999): 51-63.Walterde
Gruyter,Berlin&NewYork.
94. Kristensen NP (1998) Historical Introduction. Pp. 1-5 in N.P.Kristensen (ed.) Lepidoptera: Moths and
butterflies 1. Handbuch derZoologie/HandbookofZoologyIV/35(1999): 51-63. Walterde Gruyter, Berlin
&NewYork.
93.KristensenNP,NielsenE S (1999).Heterobathmiavalvifern.sp.:Amothwithlargeapparent“ovipositor
valves”(Lepidoptera, Heterobathmiidae). Steenstrupia24(1998): 141-156.
92.KristensenNP(1997)EarlyevolutionoftheLepidoptera+Trichopteralineage:phylogenyandtheecologi-
calscenario.MémoiresduMuséumNationald'histoireNaturelle 173:253-271 [Japanesetranslation,with
.
minordifferences: pp. 182-200 inT.Yasuda(Ed.)(1988):BiologyofMicrolepidoptera]
91. KristensenNP(1997)Thegroundplan andbasal diversificationofthehexapods. Pp. 281-293 inFortey,
R.A. &Thomas,R. H. (eds),ArthropodRelationships, Chapman&Hall, London.
90. KristensenNP(1997)Japetus Steenstrup- 100ârefter. DyriNaturogMuseum 1997/2: 21-25.
89. KristensenNP(1997)MyreloveriDanmark. DyriNaturogMuseum 1997/1:24—27.
88.NielsenES, KristensenNP(1996)TheAustralianmothfamilyLophocoronidae andthebasalphylogeny
oftheLepidoptera-Glossata. InvertebrateTaxonomy 10: 1199-1302. doi: 10.1071/IT9961199
87.DavisDR,KarsholtO,KristensenNP,NielsenES(1995)ArevisionofthegenusOgygioses(Lepidoptera:
Palaeosetidae). InvertebrateTaxonomy9: 1231 1263. doi: 10.1071/IT9951231
86. KristensenNP(1995)Fortyyears’insectphylogeneticsystematics. Hennig’s“KritischeBemerkungen...”
and subsequentdevelopments. Zoologische Beiträge(N. F.)36: 83-124.
85. KristensenNP(1995) Sydbogemollene. DyriNaturogMuseum 1995/1: 30-31.