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Camiel Doorenweerd

C. (Camiel) Doorenweerd, PhD - Biodiversity Discovery


Email: camiel.doorenweerd@naturalis.nl
Phone: +31 (0) 71 7519 344
Room number: Vondellaan 55
“DNA functions as evolution's logbook.”

I study the evolutionary drivers that shaped the diversity of four species rich genera of leaf-mining moths. In the above picture I am searching for undescribed species in a rare Taiwanese beech forest (Fagus hyatae) that is currently only found at mid altitude mountain ridges in a few localities in Taiwan but has had a wider distribution in a cooler climate in the past. (Photo by Shipher Wu)


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Leafmining larva of Ectoedemia hannoverella.


Research interest

My work focuses on the underlying evolutionary processes that explain how and why life exists and diversifies, in order to contribute to our understanding of life itself.


phylogeography, evolution, dna barcoding, phylogenetics, population genetics, co-evolution, molecular biology, taxonomy

Current research topics

Ectoedemia heringi.

Tiny plant eating moths: leaf miners

The moths I use to study evolutionary processes are present everywhere but noticed by few. The word "moth" is commonly associated with the buggers that eat the clothes in your closet or your expensive rug. However, the vast majority of the estimated 300,000 species of moths worldwide would never touch your clothes; they feed on plants. There are moths with a wingspan of over 30cm, but the moths that have caught my interest have wingspans of  0.3 to 1.5cm. The caterpillars of these moths are so tiny they can actually live inside the leaves of plants and are referred to as leaf miners. This ecological group of species is comprised of different moth genera and families that have evolved independently. Unique to this group is their ultimately intimate relationship with their hosts plants, requiring them to be highly host-specific. A single species will feed on a single host plant species only, or at most some that are closely related. Moreover, closely related mining species will often be found on closely related plant species.

Scanning the fossil record

Leaf-mining moths include some of the oldest lineages of moths with fossils of leaf-mine traces going back 102 million years ago (Cretaceous), meaning that they were already present and abundant when the dinosaurs were around. By studying leaf-mine traces on fossilized leaves as well as Amber fossils with micro CT scanning we attempt to calibrate the molecular phylogenies.

DNA barcoding and interactive keys

DNA barcoding is a taxonomic tool invaluable to my research an applied on different groups, also outside the groups of my focus mentioned below. Because there are vast numbers of undescribed species, the wealth of data of COI barcodes aids for faster species delineation. By developing interactive keys for the groups we work on, we hope to gain more knowledge on the distribution of species, while at the same time allowing more people to identify species that would normally go to one of the few worlds' specialists of those groups.

Comparative phylogenetics

By studying their DNA, a reconstruction will be made of how four different lineages of moths were able to colonize different host plants in an era where the continents were forming, ice-ages came and went, forests developed and disappeared, and the flowering plants flourished. The results from this study will help to understand how moths were able to become so diverse.

The four genera that will be compared:

  • Stigmella (Nepticulidae), 350 species known
  • Ectoedemia (Nepticulidae), 212 species known
  • Phyllonorycter (Gracillariidae), 600 species known
  • Cameraria (Gracillariidae), 102 species known

A larva of Stigmella hybnerella, mining in a leaf of the common hawthorn, Crataegus monogyna.


There is close collaboration with many Dutch and international collegues, primarily lepidopterologists. International connections include collegues in Finland, France, Great-Britain, Australia, USA, Taiwan and Japan.



Not applicable.

Available student projects



Journals SCI, peer-reviewed

Nieukerken E.J. van, Lees D.C., Doorenweerd C., Koster J.C., Bryner R., Schreurs A., Timmermans M.J.T.N., Sattler K. 2018. Two European Cornus feeding leafmining moths, Antispila petryi Martini, 1899, sp. rev. and A. treitschkiella (Fischer von Röslerstamm, 1843) (Lepidoptera, Heliozelidae): an unjustified synonymy and overlooked range expansion. Nota Lepidopterologica 41: 39-86.
Go to website (DOI)


Journals SCI, peer-reviewed

Doorenweerd C., Nieukerken E.J. van, Hoare R.J.B. 2017. Phylogeny, classification and divergence times of pygmy leaf-mining moths (Lepidoptera: Nepticulidae): the earliest lepidopteran radiation on Angiosperms?. Systematic Entomology 42: 267–287.
Go to website (DOI)

Milla L., Nieukerken E.J. van, Vijverberg R., Doorenweerd C., Wilcox S.A., Halsey M., Young D.A., Jones T., Kallies A., Hilton D.J. 2017. A preliminary molecular phylogeny of shield-bearer moths (Lepidoptera: Adeloidea: Heliozelidae) highlights rich undescribed diversity. Molecular Phylogenetics and Evolution 120: 129-143.
Go to website (URI)


Journals SCI, peer-reviewed

Mutanen M., Kivelä S.M., Vos R.A., Doorenweerd C., Ratnasingham S., Hausmann A., Huemer P., Dincă V., Nieukerken E.J. van, Lopez-Vaamonde C., Vila R., Aarvik L., Decaëns T., Efetov K.A., Hebert P.D.N., Johnsen A., Karsholt O., Pentinsaari M., Rougerie R., Segerer A., Tarmann G., Zahiri R., Godfray C.J. 2016. Species-Level Para- and Polyphyly in DNA Barcode Gene Trees: Strong Operational Bias in European Lepidoptera. Systematic Biology 65: 1024-1040.
Go to website (DOI)

Breeschoten T., Doorenweerd C., Tarasov S., Vogler A.P. 2016. Phylogenetics and biogeography of the dung beetle genus Onthophagus inferred from mitochondrial genomes. Molecular Phylogenetics and Evolution 105: 86-95.
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Nieukerken E.J. van, Doorenweerd C., Nishida K., Snyers C. 2016. New taxa, including three new genera show uniqueness of Neotropical Nepticulidae (Lepidoptera). ZooKeys 628: 1-63.
Go to website (URI)

Nieukerken E.J. van, Doorenweerd C., Hoare R.J.B., Davis D.R. 2016. Revised classification and catalogue of global Nepticulidae and Opostegidae (Lepidoptera: Nepticuloidea). ZooKeys 628: 65-246.
Go to website (URI)

Journals non-SCI, peer-reviewed

Jordan M.P., Langmaid J.R., Doorenweerd C. 2016. Morphological difference between upperside and underside leaf-mining larvae of Phyllocnistis unipunctella (Stephens, 1834) (Lep.:Gracillariidae) and its changing phenology. Entomologist's Record and Journal of Variation 128: 121-127.

Chapters in books

Doorenweerd C., Nieukerken E.J. van, Lopez-Vaamonde C., Menken S.B.J. 2016. Increased diversification rates across lineages of leaf-mining moths after entering the adaptive zone of the temperate region. 183-206:


Journals SCI, peer-reviewed

Doorenweerd C., Nieukerken E.J. van, Menken S.B. 2015. A global phylogeny of leafmining Ectoedemia moths (Lepidoptera: Nepticulidae): exploring host plant family shifts and allopatry as drivers of speciation. PLOS ONE 10: e0119586.
Go to website (DOI)

Doorenweerd C., Nieukerken E.J. van, Sohn J.C, Labandeira C.C. 2015. A revised checklist of Nepticulidae fossils (Lepidoptera) indicates an Early Cretaceous origin. Zootaxa 3963: 295-334.
Go to website (DOI)

Journals non-SCI, peer-reviewed

Roosmalen, J.A.M. van & Doorenweerd C. 2015. Coleophora gryphipennella (Hübner, 1796) (Lepidoptera, Coleophoridae) on Fragaria vesca L. (Rosaceae), a novel host, in the coastal dunes of The Netherlands. Nota Lepidopterologica 38: 147-155.
Go to website (URI)


Journals SCI, peer-reviewed

Trimbos K.B., Doorenweerd C., Kraaijeveld K., Musters C.J.M., Groen N.M., Knijff P. de, Piersma T., de Snoo G.R. 2014. Patterns in nuclear and mitochondrial DNA reveal historical and recent isolation in the Black-tailed Godwit (Limosa limosa). PLOS ONE 9: e83949.
Go to website (DOI)

Doorenweerd C., Haren M.M. van, Schermer M., Pieterse S., Nieukerken E.J. van 2014. A Linnaeus NG (TM) interactive key to the Lithocolletinae of North-West Europe aimed at accelerating the accumulation of reliable biodiversity data (Lepidoptera, Gracillariidae). ZooKeys 422: 87-101.
Go to website (DOI)

Journals non-SCI, peer-reviewed

Doorenweerd C., As B., Scheffers J. 2014. Explosieve verspreiding van de lindevouwmot: nu ook in Nederland?. The explosive expansion of the lime leaf miner in Europe: invading the Netherlands?. Entomologische Berichten 74: 111--114.
Go to website (URI)


Journals non-SCI, peer-reviewed

Groenen F., Huisman K.H., Doorenweerd C. 2013. Phalonidia manniana, een complex van twee soorten: Ph. manniana en Ph. udana (Lepidoptera: Tortricidae). Entomologische Berichten 73: 191--196.


Journals SCI, peer-reviewed

Nieukerken E.J. van, Wagner D.L., Baldessari M., Mazzon L., Angeli G., Girolami V., Duso C., Doorenweerd C. 2012. Antispila oinophylla new species (Lepidoptera, Heliozelidae), a new North American grapevine leafminer invading Italian vineyards: taxonomy, DNA barcodes and life cycle. ZooKeys 170: 29-77.
Go to website (DOI)

Nieukerken E.J. van, Doorenweerd C., Stokvis F.R., Groenenberg D.S.J. 2012. DNA barcoding of the leaf-mining moth subgenus Ectoedemia s. str. (Lepidoptera: Nepticulidae) with COI and EF1-alpha: two are better than one in recognising cryptic species. Contributions to Zoology 81: 1-24.

Journals non-SCI, peer-reviewed

Nieukerken E.J. van, Doorenweerd C., Ellis W., Huisman K., Koster J., Mey W., Muus T., Schreurs A. 2012. Bucculatrix ainsliella Murtfeldt, a new North American invader already widespread on northern red oaks (Quercus rubra) in Western Europe (Bucculatricidae). Nota Lepidopterologica 35: 135--159.

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