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Menno Schilthuizen

Prof. Dr. M. (Menno) Schilthuizen, senior researcher- Endless Forms


Email: menno.schilthuizen@naturalis.nl
Phone: +31 (0)71-75 17 285
Room number: 3rd fl., Vondellaan 55; mobile: 06-22030313
“I am fascinated by patterns in biodiversity.”

My personal drive is the endless diversity of life, which began when I was still a little boy, collecting bugs and botanising in our backyard. This childhood fascination morphed into a professional dedication to understanding the many facets involved in the evolution of species and the way communities of closely related species interact ecologically. The organisms I work with are mainly beetles and land snails, but I am not wedded to these taxa, and occasionally I choose additional kinds of organisms to suit certain research questions. Through my popular writing and public appearances I try to pre-digest complex evolutionary and ecological concepts for a general audience, and to share my personal fascinations with more than just my direct colleagues.


At Naturalis Biodiversity Center, I hold a position as research scientist and head of the Endless Forms group, which focuses on character evolution and speciation. Furthermore, I hold a professorship at Leiden University, and a research associateship at the Institute for Tropical Biology and Conservation at Universiti Malaysia Sabah. At the latter institute, I worked as an associate professor from 2000 until 2006. In addition to my scientific work, I have been active as a popular science writer and public speaker since 1994.


Research interest

My main research interest is to understand the causes for evolutionary radiations in groups of related species. Working with terrestrial gastropods and Coleoptera, I use taxonomy, phylogenetics, microanatomy, morphogenetic modeling, and field ecology to build a comprehensive analysis of the evolutionary processes involved in the radiation of the group under study. To facilitate research and share our results, I encourage web-based taxonomy initiatives for the groups we work on.

For Southeast-Asian gastropods, three Encyclopedia of Life pages have been set up by my former PhD student Liew Thor-Seng.

Borneo terrestrial mollusks
Opisthostoma (fam. Diplommatinidae)
Peninsular Malaysian terrestrial mollusks

For Cholevidae beetles, a scratchpad has been set up and maintained by my former MSc student Rik Lievers.

Cholevidae (also known as Cholevinae, Catopidae, and Catopinae) Scratchpad


land snails, community ecology, tropical biology, evolutionary radiations, terrestrial gastropods, morphology and anatomy, coleoptera

Current research topics

Evolution of endemism 

1. High-elevation endemics in Borneo

In 2012, Naturalis, together with Sabah Parks, organised a joint expedition to unravel the evolution of endemic species on Mount Kinabalu, Borneo. The expedition attracted a lot of attention in the media, for example on NBC and De Volkrant. Also, blogs were maintained, in Dutch on the Naturalis website, and in English on the Scientific American website. And animator Aart Taminiau produced a cute animation. This study eventually culminated in a paper in Nature.

More information:
Video "Jungle Genetics" produced by Nature
Research trailer "Tropical Mpuntains Are Hotspots of Biodiversity"


2. Evolution of a brass band 

In Southeast Asia, limestone (a calcareous, alkaline habitat) occurs as thousands of small, isolated, and scattered outcrops, surrounded by non-calcareous, mostly acidic substrates. Many groups of terrestrial gastropods occur only on these karsts, where they exhibit enormous morphological diversity including wonderful shapes.

Many of these species are endemic to single outcrops, and are sufficiently endangered to have been placed on the IUCN Red List. We are currently exploiting this fauna for studies to understand the diversification of shell shape as well as for community-ecological studies.

More information:
2006 Evolution paper
2011 Contributions to Zoology paper
Liew Thor-Seng's homepage
VPRO Noorderlicht: Video on extinction of limestone microsnails (in Dutch)

Rapid Evolution in the Anthropocene

1. Snail shell colour evolution

Using the classic model species Cepaea (the banded snail), this project aims to assess the evolutionary effects of human-induced climate and habitat change. Kick-started by the successful Darwin Year (2009) citizen science project Evolution Megalab, we are running studies in which we resample sites that have been sampled for Cepaea in the past. We also use transcriptomics to obtain information on the shell pigmentation genes involved. In combination with GIS and climate data, we assess and interpret the adaptive changes in morph frequencies.

More information:
Instruction video on YouTube (in Dutch)
The PLoS ONE paper


2. Evolution after invasion

A comparable study within this project is concerned with the evolutionary adaptation of native herbivorous insects to introduced plants (Evolution after invasion). Started in early 2008, we identify cases of normally oligophagous phytophagous insects that feed on woody plants without related native species. This usually includes trees and shrubs from other continents and/or genera with no European representatives. Once identified, the populations on the new host plant is screened for morphometric or genetic differentiation from the population on the native host plant, to look for incipient sympatric speciation.


Left hand, right hand

Chirality is the phenomenon that, like hands, asymmetric objects can come in two mirror-image forms. Many organs and entire organisms are asymmetric. However, in most species, only one of the two mirror-images actually exists. In snails, for example, almost all species are entirely clockwise or anti-clockwise coiled.

Rarely do both mirror-image forms occur mixed in the same population (a situation known as 'antisymmetry'). I am working on understanding the causes for this rarity of antisymmetry and the causes for antisymmetry when it does occur, in various groups of organisms, notably Amphidromus treesnails and insect genitalia.

More information: 
Interview on Noorderlicht Nieuws (in Dutch)
Journal of Evolutionary Biology paper
Masaki Hoso's homepage
My inaugural lecture 
Research trailer asymmetry


Satoshi Chiba, Tohoku University, Japan

Martin Haase, University of Greifswald, Germany

Joris Koene, VU University, Amsterdam, the Netherlands

Virginie Orgogozo and Michaël Lang, Institut Jacques Monod, Paris, France

Liew Thor Seng and Bakhtiar Effendi Yahya, Universiti Malaysia Sabah, Malaysia

Rampal Etienne, University of Groningen, the Netherlands

Michel Perreau and Louis Deharveng, Muséum National d'Histoire Naturelle, Paris, France

Supervision PhDs


Salmah Yaakop (Rijksuniversiteit Groningen, 2011); now lecturer at Universiti Kebangsaan Malaysia

Kim Meijer (Rijksuniversiteit Groningen, 2013)

Thor-Seng Liew (Universiteit Leiden, 2014); now lecturer at Universiti Malaysia Sabah

Kasper Hendriks (Rijksuniversiteit Groningen, projected for 2019); here his page at RUG.

Zacaery Mohd. Khalik (Universiteit Leiden, projected for 2019)

Kevin Beentjes (Universiteit Leiden, projected for 2019)



Student projects

All the projects I work on, have the possibility for students (Bachelor, Master, or HBO) to do smaller research projects. I also occasionally help students design a project that is tailor-made to their wishes, provided it fits within one of my research interests. Students are advised to contact me at least 6 months before they wish to start their project. If I have no available projects, I will try to find alternatives within the Character Evolution Group.

BSc courses

I teach the 1st-year sub-course "Basics of Systematics" (in the course description termed, "Fylogenie") in the Leiden University biology program. I am also initiator and one of the coordinators for the MOOC Evolution for Everyone, available at Coursera from September 2016. 

MSc courses

Together with Jeremy Miller, I am coordinator and lecturer of the course Orientation EBC, in the Leiden University MSc program Evolution, Biodiversity, and Conservation (EBC).

I also coordinate and teach the Field Course Tropical Biodiversity.

Other teaching

In addition to the above, I also perform the following teaching activities:

  • Students can read a book with me in the Leiden University 1st Yr BSc course "popular science book". For this year, the book is: A Guinea Pig's History of Biology (Jim Endersby)
  • I supervise one or two projects in the Evolution and Ecology mini-projects of the 1st Yr BSc (Leiden University); see here for a video a student group made a few years ago.
  • Two lectures on speciation in Genomic Architecture (MSc program EBC; Leiden University)
  • Until 2013, I taught the course Diversiteit Dieren (lectures and labs) at the University of Groningen

Supervision MSc Students

2014 Supervisor - P.A.J. Bakker - Evolution and diversification in euconulid-shaped snails
2014 Supervisor - Sipman, I.E., - Right-handed snails and Left-handed crabs (or vice versa)
2012 Supervisor - Tamara Hoogenboom - Evolution of asymmetric genitalia in Cholevini
2012 Supervisor - Rick van Beek - A search for asymmetry: a morphological analysis on the genitalia of Cholevini (Leiodidae: Cholevinae)
2012 Co-supervisor - Nikita van Zeijl - Heat islands and snails in big cities
2011 Supervisor - Els Baalbergen - Larvae of Drilini (Coleoptera: Elateridae) beetles preying on snails
2011 Supervisor - Renate Helwerda - Predation on Greek Albinaria (Pulmonata: Clausiliidae) as a possible driver of shell evolution
2011 Supervisor - Rense Schelfhorst - Natural selection on shell traits in Albinaria, driven by predators
2011 Co-supervisor - Annebelle Kok - Shell growth and development in two Opisthostoma species
2011 Supervisor - Sigrid Hendrikse - Differences between dextral and sinistral individuals of the tropical land snail Amphidromus inversus
2011 Supervisor - Youri Lammers - Rare alleles increase genome-wide in a hybrid zone
2010 Supervisor - Rik Lievers - DNA‐Barcoding and Web‐Based Taxonomy for Cholevinae (Coleoptera: Leiodidae)
2010 Co-supervisor - Irma Valera - Phenotypic plasticity, genetic assimilation and the evolution of annualism in Rivulidae
2010 Co-supervisor - Marian van Hulsel - BOSTRYX AND SCUTALUS IN PERU Comparing habitat for different morphologies
2010 Co-supervisor - Nienke Vastenhout - Systematics and biogeography of Avakubia (Gastropoda; Streptaxidae)
2010 Co-supervisor - Hidde Zemel - evolution of insect host-races specialized on Japan-derived invasive plants in Europe and vice versa in Japan
2010 Nicole Webster - Phylogenetic reconstruction and shell evolution of the Diplommatinidae (Gastropoda: Caenogastropoda)
2010 Paul Hoekstra - Phylogenetic relationships between isolated populations of the limestone-dwelling microsnail Gyliotrachela hungerfordiana (Gastropoda: Vertiginidae)
2009 Supervisor - Nicole Webster - Phylogenetic Reconstruction and Shell Evolution of the Subfamily Diplommatininae (Gastropoda: Caenogastropoda: Diplommatinidae)

Supervision BSc Students

2012 Supervisor - Thijmen Breeschoten - The evolution pattern of asymmetric genitalia in Cyclocephalini (Coleoptera: Scarabaeidae: Dynastinae) beetles
2012 Co-supervisor - Kasper Hendriks - A reconstruction of the phylogeny of Kelp Gull (Larus dominicanus)
2012 Co-supervisor - Ralph Werring - Herbivore insecten op de Amerikaanse vogelkers (Prunus serotina)
2012 Co-supervisor - Robin van der Ven - Herbivore insecten op de Amerikaanse vogelkers (Prunus serotina)
2012 Co-supervisor - Nils Sosef - Herbivore insecten op de Amerikaanse vogelkers (Prunus serotina)
2011 Supervisor - Devi van der Horst - Ontwikkeling van kweektechnieken voor Cholevidae
2011 Supervisor - Nils Beveridge - Host-shift and enemy-free space of herbivorous insects on exotic plants
2011 Co-supervisor - Frida Feijen - An Introduction to Phylogenetic Studies on Diopsidae (Diptera)
2010 Supervisor - Devi van der Horst - DNA-barcoding van Nederlandse Cholevidae ten behoeve van forensische entomologie
2010 Supervisor - Jessy Dommershuijzen - DNA-barcoding van Nederlandse Cholevidae ten behoeve van forensische entomologie
2010 Supervisor - Dwayne van der Klugt - Adaptive Host ­Shift in Fruit Flies (Rhagoletis cerasi & R. alternata, Diptera: Tephritidae) on exotic plants
2009 Supervisor - Remco Schipper - De successie van Cholevidae op kadavers
2009 Supervisor - Joyce van der Steen - De successie van Cholevidae op kadavers
2009 Supervisor - Melanie Meijer zu Schlochtern - De evolutie van asymmetrie en chiraliteit bij de mannelijke genitaliën van Sciodrepoides (Coleoptera: Cholevidae)
Available student projects



Journals SCI, peer-reviewed

Njunjić I., Perrard A., Hendriks K., Schilthuizen M., Perreau M., Merckx V., Baylac M., Derharveng L. 2018. Comprehensive evolutionary analysis of the Anthroherpon radiation (Coleoptera, Leiodidae, Leptodirini). PLOS ONE 13: e0198367.
Go to website (DOI)

Beentjes K.K., Speksnijder A.G.C.L., Schilthuizen M., Schaub B.E.M., Hoorn B.B. van der 2018. The influence of macroinvertebrate abundance on the assessment of freshwater quality in The Netherlands. Metabarcoding and Metagenomics 2: e26744.
Go to website (DOI)

Schilthuizen M., Clavera A.P., Khoo M.S., Bondar C.A., Elder C.H.S., Bouma A.M., Eddington T.D., Reisinger C., Cosentino E., Rossato M., Delledonne M. 2018. Bringing the lab to the field: a new lowland Microparmarion semi-slug (Gastropoda: Ariophantidae) described and DNA-barcoded in the forest. Journal of Molluscan Studies: eyy052.
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Nekola J.C., Schilthuizen M., Szabo K., Horsáková V., Horsák M. 2018. First evidence for long‐term stasis in wet‐tropics land snail community composition. Ecography: pending.
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Velzen R. van, Holmer R., Bu F., Rutten L., Zeijl A. van, Liu W., Santuari L., Cao Q., Sharma T., Shen D., Roswanjaya Y., Wardhani T.A.K., Kalhor M.S., Jansen J., Hoogen J. van den, Güngör B., Hartog M., Hontelez J., Verver J., Yang W.-C., Schijlen E., Repin R., Schilthuizen M., Schranz M.E., Heidstra R., Miyata K., Fedorova E., Kohlen W., Bisseling T., Smit S., Geurts R. 2018. Comparative genomics of the nonlegume Parasponia reveals insights into evolution of nitrogen-fixing rhizobium symbioses. Proceedings of the National Academy of Sciences of the United States of America 115: E4700–E4709.
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Schilthuizen M., Perreau M., Njunjić I. 2018. A review of the Cholevinae from the island of Borneo (Coleoptera, Leiodidae). ZooKeys 777: 57-108.
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Khalik M.Z., Hendriks K., Vermeulen J.J., Schilthuizen M. 2018. A molecular and conchological dissection of the “scaly” Georissa of Malaysian Borneo (Gastropoda, Neritimorpha, Hydrocenidae). ZooKeys 773: 1-55.
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Journals non-SCI, peer-reviewed

Miller J.A., Freund C., Rambonnet L., Koets L., Barth N., Linden C. van der, Geml J., Schilthuizen M., Burger R., Goossens B. 2018. Dispatch from the field II: the mystery of the red and blue Opadometa male (Araneae, Tetragnathidae, Opadometa sarawakensis). Biodiversity Data Journal 6: e24777.
Go to website (URI)

Authored books

Schilthuizen M. 2018. Darwin in de Stad. Atlas Contact,

Schilthuizen M. 2018. Darwin Comes to Town; how the urban jungle drives evolution. 304 p. Picador USA,


Journals SCI, peer-reviewed

Miller J.A., Riemsdijk I. van, Khalik M.Z., Scager D.J., Schilthuizen M. 2017. Comment on Falade et al. (2016) DNA-barcoding of Clarias gariepinus, Coptedon zilii and Sarotherodon melanotheron from Southwestern Nigeria. F1000Research -
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Geml J., Morgado L.N., Semenova-Nelsen T.A., Schilthuizen M. 2017. Changes in richness and community composition of ectomycorrhizal fungi among altitudinal vegetation types on Mount Kinabalu in Borneo. New Phytologist 215: 454-468.
Go to website (URI)

Njunjić I., Schilthuizen M., Pavićević D., Perreau M. 2017. Further clarifications to the systematics of the cave beetle genera Remyella and Rozajella (Coleoptera: Leiodidae: Cholevinae: Leptodirini). Arthropod Systematics and Phylogeny 75: 141-158.

Haren M.M. van, Liljestrand Rönn J., Schilthuizen M., Arnqvist G. 2017. Postmating sexual selection and the enigmatic jawed genitalia of Callosobruchus subinnotatus. Biology Open: pending.
Go to website (DOI)

Hudson L.N. et al. (>100 authors) 2017. The database of the PREDICTS (Projecting Responses of Ecological Diversity In Changing Terrestrial Systems) project. Ecology and Evolution 7: 145-188.
Go to website (DOI)

Schilthuizen M., Sipman I., Zwaan H. 2017. Sexual dimorphism in shell coloration of Plectostoma (Caenogastropoda: Diplommatinidae) is caused by polyenes. Journal of Molluscan Studies: pending.
Go to website (DOI)

Ożgo M., Liew T.S., Webster N.B., Schilthuizen M. 2017. Inferring microevolution from museum collections and resampling: lessons learned from Cepaea. PeerJ 5: e3938.
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Kerkvliet J., Boer T. de, Schilthuizen M., Kraaijeveld K. 2017. Candidate genes for shell colour polymorphism in Cepaea nemoralis. PeerJ: e3715.
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Schilthuizen M., Langelaan R., Hemmings N., Oostenbrugge W. van, Visser S. 2017. An unexpected twist: Sperm cells coil to the right in land snails and to the left in song birds. Contributions to Zoology 86: 297-302.
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Schilthuizen M., Sipman I., Zwaan J.C. 2017. Sexual dimorphism in shell coloration of Pletostoma (Caenogastropoda: Diplommatinidae) is caused by polyenes. Journal of Molluscan Studies: 1-3.
Go to website (URI)

Journals non-SCI, peer-reviewed

Schilthuizen M., Seip L.A., Otani S., Suhaimi J., Njunjić I. 2017. Three new minute leaf litter beetles discovered by citizen scientists in Maliau Basin, Malaysian Borneo (Coleoptera: Leiodidae, Chrysomelidae). Biodiversity Data Journal: e21947.
Go to website (DOI)


Journals SCI, peer-reviewed

T.-S. Liew, Schilthuizen M. 2016. A method for quantifying, visualising, and analysing gastropod shell form. PLOS ONE 11: e0157069.
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Meijer K., Smit C., Schilthuizen M., Beukeboom L.W. 2016. Fitness benefits of the fruit fly Rhagoletis alternata on a non-native rose host. Oecologia 181: 185-192.
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Schilthuizen M., Jong P. de, Beek R. van, Hoogenboom T., Schlochtern M.M.Z. 2016. The evolution of asymmetric genitalia in Coleoptera. Philosophical Transactions of the Royal Society B: Biological Sciences 371: 20150400.
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Meijer K., Schilthuizen M., Beukeboom L., Smit C. 2016. A review and meta-analysis of the enemy release hypothesis in plant–herbivorous insect systems. PeerJ 4: e2778.
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Rhebergen F.T., Courtier-Orgogozo V., Dumont J., Schilthuizen M., Lang M. 2016. Drosophila pachea asymmetric lobes are part of a grasping device and stabilize one-sided mating. BMC Evolutionary Biology 16: 176.
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Schilthuizen M., Santos Pimenta L.P., Lammers Y., Steenbergen P.J., Flohil M., Beveridge N.G.P., Duijn, P.T. van, Meulblok M.M., Sosef N., Ven R. van de, Werring R., Beentjes K.K., Meijer K., Vos R.A., Vrieling K., Gravendeel B., Choi Y., Verpoorte R., Smit C., Beukeboom L.B. 2016. Incorporation of an invasive plant into a native insect herbivore food web. PeerJ 4: e1954.
Go to website (DOI)

Meijer K., Smit C., Schilthuizen M., Beukeboom L.W. 2016. Fitness benefits of the fruit fly Rhagoletis alternata on a non-native rose host. Oecologia 181: 185-192.
Go to website (DOI)

Njunjić I., Perreau M., Hendriks K., Schilthuizen M., Deharveng L. 2016. The cave beetle genus Anthroherpon is polyphyletic; molecular phylogenetics and description of Graciliella n. gen. (Leiodidae, Leptodirini). Contributions to Zoology 85: 337-359.
Go to website (URI)

Hudson L.N., Newbold T., Contu S., et al. 2016. The database of the PREDICTS (Projecting Responses of Ecological Diversity in Changing Terrestrial Systems) project. Ecology and Evolution 7: 145–188.
Go to website (DOI)

Journals non-SCI, peer-reviewed

Gravendeel B., Schilthuizen M. 2016. Inheemse insecten hebben aanval op Bospest ingezet. De Levende Natuur 117: 208-209.

Kolfschoten L. van, Selveno S., Danica J. Stark D.J., Schilthuizen M. 2016. Nest density of the Black-and-Red Broadbill (Cymbirhynchus macrorhyncos) along the Kinabatangan River, in Relation to Riverine Habitat Reduction. Journal of Tropical Biology and Conservation 13: 157–168.

Baalbergen E., Schelfhorst R., Schilthuizen M. 2016. Drilus larvae in the Netherlands. Entomologische Berichten 76: 165-173.

Authored books

Schilthuizen M. 2016. Organi sessuali, evoluzione e biodiversità. 324 p. Bollati Boringhieri, Torino. ISBN 9788833974484.

Schilthuizen M. 2016. Waarom Zijn Er Zoveel Soorten?. 304 p. Atlas Contact, Amsterdam. ISBN 9789045031415.

Schilthuizen M. 2016. Comme les Bêtes. Ce que les animaux nous apprennent de notre sexualité. 320 p. Flammarion, Paris. ISBN 208134839X.

Chapters in books

Schilthuizen M. 2016. Robin Baker and Mark Bellis. 1-3:
Go to website (DOI)


Journals SCI, peer-reviewed

Kundrata R., Baalbergen E., Bocak L., Schilthuizen M. 2015. The origin and diversity of Drilus Olivier, 1790 (Elateridae: Agrypninae: Drilini) in Crete based on mitochondrial phylogeny. Systematics and Biodiversity 13: 52-75.
Go to website (DOI)

Schilthuizen M., Vairappan C.S., Slade E.M., Mann D.J., Miller J.A. 2015. Specimens as primary data: museums and ‘open science’. Trends in Ecology & Evolution 30: 237-238.
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Meijer K., Zemel H., Chiba S., Smit C., Beukeboom L.W., Schilthuizen M. 2015. Phytophagous insects on native and non-native host plants: combining the community approach and the biogeographical approach. PLOS ONE 10: e0125607.
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Merckx V.S.F.T., Hendriks K.P., Beentjes K.K., Mennes C.B., Becking L.E., Peijnenburg K.T.C.A., Afendy A., Arumugam N., de Boer H., Biun A., Buang M.M., Chen P.-P., Chung A.Y.C., Dow R., Feijen F.A.A., Feijen H., Feijen-van Soest C, Geml J., Geurts R., Gravendeel B., Hovenkamp P., Imbun P., Ipor I., Janssens S.B., Jocqué M. Kappes H., Khoo E., Koomen P., Lens F., Majapun R.J., Morgado L.N., Neupane S., Nieser N., Pereira J.T., Rahman H., Sabran S., Sawang A., Schwallier R.M., Shim P.-S., Smit H., Sol N., Spait M., Stech M., Stokvis F., Sugau J.B., Suleiman M., Sumail S., Thomas D.C., van Tol J., Tuh F.Y.Y., Yahya B.E., Nais J., Repin R., Lakim M. & Schilthuizen M. 2015. Evolution of endemism on a young tropical mountain. Nature 524: 347-350.
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Vermeulen J.J., T.-S. Liew, Schilhuizen M. 2015. Additions to the knowledge of the land snails of Sabah (Malaysia, Borneo), including 48 new species. ZooKeys 531: 1-139.
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Journals SCI, peer-reviewed

Baalbergen E., Helwerda R., Schelfhorst R., Cajas R.F. Castillo, Moorsel C.H.M. van, Kundrata R., Welter-Schultes F.W., Giokas S., Schilthuizen M. 2014. Predator-prey interactions between shell-boring beetle larvae and rock-dwelling land snails. PLOS ONE 9: e100366.
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Schilthuizen M., Kellermann V. 2014. Contemporary climate change and terrestrial invertebrates: evolutionary versus plastic changes. Evolutionary Applications 7: 56-67.
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Santos Pimenta L.P., Schilthuizen M., Verpoorte R., Choi Y.H. 2014. Quantitative analysis of amygdalin and prunasin in Prunus serotina Ehrh. using H-1-NMR spectroscopy. Phytochemical Analysis 25: 122-126.
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Haelewaters D., Schilthuizen M., Pfister D.H. 2014. On Diphymyces (Laboulbeniales, Ascomycota) in Malaysian Borneo. Plant Ecology and Evolution 147: 93-100.
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Kappes H., Schilthuizen M. 2014. Habitat effects on slug assemblages and introduced species. Journal of Molluscan Studies 80: 47-54.
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Liew T.-S., Vermeulen J.J., Marzuki M.E. bin, Schilthuizen M. 2014. A cybertaxonomic revision of the micro-landsnail genus Plectostoma Adam (Mollusca, Caenogastropoda, Diplommatinidae), from Peninsular Malaysia, Sumatra and Indochina. ZooKeys 393: 1-107.
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Capinha C., Vermeulen J.J., Lakim M. bin, Schilthuizen M., Kappes H. 2014. Susceptibility of tropical mountain forests to biological invasions from the temperate and subtropical zone, exemplified by Zonitoides snails (Gastropoda: Gastrodontidae). Raffles Bulletin of Zoology 62: 600-609.
Go to website (URI)

Helwerda R.A., Schilthuizen M. 2014. Predation on Greek Albinaria (Pulmonata: Clausiliidae) by Poiretia (Pulmonata: Oleacinidae) and by an unknown organism making circular holes: possible drivers of shell evolution. Journal of Molluscan Studies 80: 272-279.
Go to website (DOI)

Liew T.S., Kok A.C.M., Schilthuizen M., Urdy S. 2014. On growth and form of irregular coiled-shell of a terrestrial snail: Plectostoma concinnum (Fulton, 1901) (Mollusca: Caenogastropoda: Diplommatinidae). PeerJ 2: e383.
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Journals non-SCI, peer-reviewed

Miller J.A., Schilthuizen M., Lilliendahl Burmester J., Graaf L. van der, Merckx V., Jocqué M., Kessler P.J.A., Fayle T.M., Breeschoten T., Broeren R., Bouman R., Chua W.-J., Feijen F., Fermont T., Groen K., Groen M., Kil N.J.C., Laat H.A. de, Moerland M.S., Moncoquet C., Panjang E., Philip A.J., Roca-Eriksen R., Rooduijn B., Santen M. van, Swakman V., Evans M.N., Evans L.J., Love K., Joscelyne S.H., Tober A.V., Wilson H.F., Ambu L.N., Goossens B. 2014. Dispatch from the field: ecology of ground-web-building spiders with description of a new species (Araneae, Symphytognathidae). Biodiversity Data Journal 2: 1-13.
Go to website (URI)

Liew T.S., Schilthuizen M. 2014. Association between shell morphology of micro-land snails (genus Plectostoma) and their predator’s predatory behaviour. PeerJ 2: 1e329.

Authored books

Schilthuizen M. 2014. Darwins Peep Show. Atlas Contact,

Schilthuizen M. 2014. Nature's Nether Regions: What the Sex Lives of Bugs, Birds, and Beasts Tell Us About Evolution, Biodiversity, and Ourselves. 256 p. Viking Adult, ISBN 9780670785919 |.

Schilthuizen M. 2014. Anche le Coccinelle nel loro piccolo..... Organi sessuali ed evoluzione. Bollati Boringhieri, ISBN 978-8833924359.


Journals SCI, peer-reviewed

Schilthuizen M. 2013. Rapid, habitat-related evolution of land snail colour morphs on reclaimed land. Heredity 110: 247-252.
Go to website (DOI)

Schilthuizen M. 2013. Something gone awry: unsolved mysteries in the evolution of asymmetric animal genitalia. Animal Biology 63: 1-20.
Go to website (DOI)

Schilthuizen M., Lammers Y. 2013. Hybrid zones, barrier loci and the 'rare allele phenomenon'. Journal of Evolutionary Biology 26: 288-290.
Go to website (DOI)

Breeschoten T., Clark D.R., Schilthuizen M. 2013. Evolutionary patterns of asymmetric genitalia in the beetle tribe Cyclocephalini (Coleoptera: Scarabaeidae: Dynastinae). Contributions to Zoology 82: 95-106.

Semenov A.V., Elsas J.D. van, Glandorf D.C.M., Schilthuizen M., Boer W.F. de 2013. The use of statistical tools in field testing of putative effects of genetically modified plants on nontarget organisms. Ecology and Evolution 3: 2739-2750.
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Schilthuizen M., Liew T.-S., Liew T.-H., Berlin P., King J.P., Lakim M. 2013. Species diversity patterns in insular land snail communities of Borneo. Journal of the Geological Society 170: 539-545.
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Koene J.M., Liew T.S., Montagne-Wajer K., Schilthuizen M. 2013. A syringe-like love dart injects male accessory gland products in a tropical hermaphrodite. PLOS ONE 8: e69968.
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Journals SCI, peer-reviewed

Ozgo M., Schilthuizen M. 2012. Evolutionary change in Cepaea nemoralis shell colour over 43 years. Global Change Biology 18: 74-81.
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Butlin R., Debelle A., Kerth C., Snook R.R., Beukeboom L.W., Cajas R.F.C., Diao W., Maan M.E., Paolucci S., Weissing F.J., Zande L. van de, Hoikkala A., Geuverink E., Jennings J., Kankare M., Knott K.E., Tyukmaeva V.I., Zoumadakis C., Ritchie M.G., Barker D., Immonen E., Kirkpatrick M., Noor M., Macias G.C., Schmitt T., Schilthuizen M. 2012. What do we need to know about speciation?. Trends in Ecology & Evolution 27: 27-39.
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Schilthuizen M., Rutten E.M.J., Haase M. 2012. Small-scale genetic structuring in a tropical cave snail and admixture with its above-ground sister species. Biological Journal of the Linnean Society 105: 727-740.
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Schilthuizen M. 2012. Contemporary evolution Scrutinising snail shells. Heredity 108: 364-365.
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Groenenberg D.S.J., Pirovano W., Gittenberger E., Schilthuizen M. 2012. The complete mitogenome of Cylindrus obtusus (Helicidae, Ariantinae) using Illumina next generation sequencing. BMC Genomics 13: 114.
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Webster N.B., Dooren T.J.M. van, Schilthuizen M. 2012. Phylogenetic reconstruction and shell evolution of the Diplommatinidae (Gastropoda: Caenogastropoda). Molecular Phylogenetics and Evolution 63: 625-638.
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Schilthuizen M., Gravendeel B. 2012. Left-right asymmetry in plants and animals: a gold mine for research. Contributions to Zoology 81: 75-78.

Schilthuizen M., Haase M., Koops K., Looijestijn S.M., Hendrikse S. 2012. The ecology of shell shape difference in chirally dimorphic snails. Contributions to Zoology 81: 95-101.

Journals non-SCI, peer-reviewed

Meijer K., Smit C., Beukeboom L.W., Schilthuizen M. 2012. Native insects on non-native plants in The Netherlands: curiosities or common practice?. Entomologische Berichten 72: 288-293.


Journals SCI, peer-reviewed

Schilthuizen M. 2011. Lopsided love. New Scientist 210: 42-45.

Silvertown J., Cook L., Cameron R., Dodd M., McConway K., Worthington J., Skelton P., Anton C., Bossdorf O., Baur B., Schilthuizen M., Fontaine B., Sattmann H., Bertorelle G., Correia M., Oliveira C., Pokryszko B., Ozgo M., Stalazas A., Gill E., Rammul U., Solymos P., Feher Z., Juan X. 2011. Citizen Science Reveals Unexpected Continental-Scale Evolutionary Change in a Model Organism. PLOS ONE 6: e18927.
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Schilthuizen M., Heuven B.-J. van 2011. Dextral and sinistral Amphidromus inversus (Gastropoda: Pulmonata: Camaenidae) produce dextral sperm. Zoomorphology 130: 283-287.
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Hoekstra P., Schilthuizen M. 2011. Phylogenetic relationships between isolated populations of the limestone-dwelling microsnail Gyliotrachela hungerfordiana (Gastropoda: Vertiginidae). Journal of Zoological Systematics and Evolutionary Research 49: 266-272.
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Schilthuizen M. 2011. Community ecology of tropical forest snails: 30 years after Solem. Contributions to Zoology 80: 1-15.

Schilthuizen M., Scholte C., Wijk R.E.J. van, Dommershuijzen J., Horst D. van der, Schlochtern M.M.Z., Lievers R., Groenenberg D.S.J. 2011. Using DNA-barcoding to make the necrobiont beetle family Cholevidae accessible for forensic entomology. Forensic Science International 210: 91-95.
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Schilthuizen M., Giesbers M.C.W.G., Beukeboom L.W. 2011. Haldane's rule in the 21st century. Heredity 107: 95-102.
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Hoekstra P., Schilthuizen M. 2011. Phylogenetic relationships between isolated populations of the limestone-dwelling microsnail Gyliotrachela hungerfordiana (Gastropoda: Vertiginidae). Journal of Zoological Systematics and Evolutionary Research 49: 266-272.
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Journals SCI, peer-reviewed

Schilthuizen M., Haase M. 2010. Disentangling true shape differences and experimenter bias: are dextral and sinistral snail shells exact mirror images?. Journal of Zoology 282: 191-200.
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Liew T.S., Schilthuizen M., Lakim M. 2010. The determinants of land snail diversity along a tropical elevational gradient: insularity, geometry and niches. Journal of Biogeography 37: 1071-1078.
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Schilthuizen M., Raes N., Linder P., Triantis K., Fernandez-Palacios J.M., Whittaker R.J. 2010. Special Issue: Evolutionary Islands - 150 Years After Darwin. Journal of Biogeography 37: 983-983.
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Journals non-SCI, peer-reviewed

Breure A.S.H., Groenenberg D.S.J., Schilthuizen M. 2010. New insights in the phylogenetic relations within the Orthalicoidea (Gastropoda, Stylommatophora) based on 28S sequence data. Basteria 74: 25-31.

Authored books

Schilthuizen M. 2010. Waarom zijn er zoveel soorten?. KNNV Uitgeverij,


Journals SCI, peer-reviewed

Liew T.S., Schilthuizen M., Vermeulen J.J. 2009. Systematic revision of the genus Everettia Godwin-Austen, 1891 (Mollusca: Gastropoda: Dyakiidae) in Sabah, northern Borneo. Zoological Journal of the Linnean Society 157: 515-550.
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Sheldon F.H., Lim H.C., Nais J., Lakim M., Tuuga A., Malim P., Majuakim J., Lo A., Schilthuizen M., Hosner P.A., Moyle R.G. 2009. Observations on the ecology, distribution and biogeography of forest birds in Sabah, Malaysia. Raffles Bulletin of Zoology 57: 577-586.

Schilthuizen M., Looijestijn S. 2009. The sexology of the chirally dimorphic snail species Amphidromus inversus (Gastropoda: Camaenidae). Malacologia 51: 379-387.

Journals non-SCI, peer-reviewed

Schilthuizen M., Liew T.S. 2009. The slugs and semislugs of Sabah, Malaysian Borneo (Gastropoda, Pulmonata: Veronicellidae, Rathouisiidae, Ariophantidae, Limacidae, Philomycidae). Basteria 72: 287-306.


Journals SCI, peer-reviewed

Liew T.S., Clements R., Schilthuizen M. 2008. Sampling micromolluscs in tropical forests: one size does not fit all. Zoosymposia 1: 271-280.

Gomes S.R., Picanco J.B., Schilthuizen M., Thome J.W. 2008. Valiguna flava (Heynemann, 1885) from Indonesia and Malaysia: Redescription and comparison with Valiguna siamensis (Martens, 1867) (Gastropoda : Soleolifera : Veronicellidae). Veliger 50: 163-170.

Schilthuizen M. 2008. A parasitic moment - Menno Schilthuizen of the University of Malaysia Sabah replies. Natural History 117: 58-58.

Clements R., Ng P.K.L., Lu X.X., Ambu S., Schilthuizen M., Bradshaw C.J.A. 2008. Using biogeographical patterns of endemic land snails to improve conservation planning for limestone karsts. Biological Conservation 141: 2751-2764.
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Clements Re., Liew T.-S., Vermeulen J.Jan, Schilthuizen M. 2008. Further twists in gastropod shell evolution. Biology Letters 4: 179-182.
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Authored books

Schilthuizen M. 2008. The Loom of Life: Unravelling Ecosystems. Springer Verlag,


Journals SCI, peer-reviewed

Blakemore R.J., Csuzdi C., Ito M.T., Kaneko N., Kawaguchi T., Schilthuizen M. 2007. Taxonomic status and ecology of Oriental Pheretima darnleiensis (Fletcher, 1886) and other earthworms (Oligochaeta : Megascolecidae) from Mt Kinabalu, Borneo. Zootaxa 1613: 23-44.

Haase M., Schilthuizen M. 2007. A new Georissa (Gastropoda : Neritopsina : Hydrocenidae) from a limestone cave in Malaysian Borneo. Journal of Molluscan Studies 73: 215-221.
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Schilthuizen M., Craze P.G., Cabanban A.S., Davison A., Stone J., Gittenberger E., Scott B.J. 2007. Sexual selection maintains whole-body chiral dimorphism in snails. Journal of Evolutionary Biology 20: 1941-1949.
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Benedick S., White T.A., Searle J.B., Hamer K.C., Mustaffa N., Vun Khen C., Mohamed M., Schilthuizen M., Hill J.K. 2007. Impacts of habitat fragmentation on genetic diversity in a tropical forest butterfly on Borneo. Journal of Tropical Ecology 23: 623-634.
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Journals SCI, peer-reviewed

Schilthuizen M., Til A. van, Salverda M., Liew T.-S., James S., Elahan B. bin, Vermeulen J.J. 2006. Microgeographic evolution of snail shell shape and predator behavior. Evolution 60: 1851-1858.

Craze P.G., Elahan B., Schilthuizen M. 2006. Opposite shell-coiling morphs of the tropical land snail Amphidromus martensi show no spatial-scale effects. Ecography 29: 941-941.

Benedick S., Hill J.K., Mustaffa N., Chey V.K., Maryati M., Searle J.B., Schilthuizen M., Hamer K.C. 2006. Impacts of rain forest fragmentation on butterflies in northern Borneo: species richness, turnover and the value of small fragments. Journal of Applied Ecology 43: 967-977.
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Clements R., Sodhi N.S., Schilthuizen M., Ng P.K.L. 2006. Limestone karsts of southeast Asia: Imperiled arks of biodiversity. BioScience 56: 733-742.

Weerd D.R. uit de, Groenenberg D.S.J., Schilthuizen M., Gittenberger E. 2006. Reproductive character displacement by inversion of coiling in clausiliid snails (Gastropoda, Pulmonata). Biological Journal of the Linnean Society 88: 155-164.


Journals SCI, peer-reviewed

Charlat S., Hornett E.A., Dyson E.A., Ho P.P.Y., Loc N.T., Schilthuizen M., Davies N., Roderick G.K., Hurst G.D.D. 2005. Prevalence and penetrance variation of male-killing Wolbachia across Indo-Pacific populations of the butterfly Hypolimnas bolina. Molecular Ecology 14: 3525-3530.
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Schilthuizen M., Liew T.S., Elahan B. bin, Lackman-Ancrenaz I. 2005. Effects of karst forest degradation on pulmonate and prosobranch land snail communities in Sabah, Malaysian Borneo. Conservation Biology 19: 949-954.

Schilthuizen M. 2005. The darting game in snails and slugs. Trends in Ecology & Evolution 20: 581-584.
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Schilthuizen M., Davison A. 2005. The convoluted evolution of snail chirality. Naturwissenschaften 92: 504-515.
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Moyle R.G., Schilthuizen M., Rahman M.A., Sheldon F.H. 2005. Molecular phylogenetic analysis of the white-crowned forktail Enicurus leschenaulti in Borneo. Journal of Avian Biology 36: 96-101.

Schilthuizen M., Cabanban A.S., Haase M. 2005. Possible speciation with gene flow in tropical cave snails. Journal of Zoological Systematics and Evolutionary Research 43: 133-138.
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Schilthuizen M., Scott B.J., Cabanban A.S., Craze P.G. 2005. Population structure and coil dimorphism in a tropical land snail. Heredity 95: 216-220.
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Journals SCI, peer-reviewed

Schilthuizen M., Gutteling E., Moorsel C.H.M. van, Welter-Schultes F.W., Haase M., Gittenberger E. 2004. Phylogeography of the land snail Albinaria hippolyti (Pulmonata : Clausiliidae) from Crete, inferred from ITS-1 sequences. Biological Journal of the Linnean Society 83: 317-326.

Schilthuizen, M., Hoekstra, R.F., Gittenberger, E. 2004. Hybridization, rare alleles and adaptive radiation. Trends in Ecology & Evolution 19: 404-405.


Journals SCI, peer-reviewed

Schilthuizen M. 2003. Sexual selection on land snail shell ornamentation: a hypothesis that may explain shell diversity. BMC Evolutionary Biology 3: 13.

Schilthuizen M., Chai H.N., Kimsin T.E., Vermeulen J.J. 2003. Abundance and diversity of land-snails (Mollusca : Gastropoda) on limestone hills in Borneo. Raffles Bulletin of Zoology 51: 35-42.


Journals SCI, peer-reviewed

Schilthuizen M. 2002. Mollusca: an evolutionary Cornucopia. Trends in Ecology & Evolution 17: 8-9.

Schilthuizen M. 2002. Caution: Species crossing. Natural History 111: 62-71.

Schilthuizen M. 2002. The unselfish genome - The case for cooperating genes. Natural History 111: 68-69.

Schilthuizen M. 2002. What evolution is. Science 295: 50-50.

Schilthuizen M., Teravainen M.I.F., Faezamoltha N., Tawith K., Ibrahim H., Chea S.M., Chuan C.P., Daim L.J., Jubaidi A., Madjapuni M.J., Sabeki M., Mokhtar A. 2002. Microsnails at microscales in Borneo: Distributions of Prosobranchia versus Pulmonata. Journal of Molluscan Studies 68: 255-258.


Journals SCI, peer-reviewed

Schilthuizen M., Rutjes H.A. 2001. Land snail diversity in a square kilometre of tropical rainforest in Sabah, Malaysian Borneo. Journal of Molluscan Studies 67: 417-423.

Schilthuizen M., Hoekstra R.F., Gittenberger E. 2001. The 'rare allele phenomenon' in a ribosomal spacer. Molecular Ecology 10: 1341-1345.


Journals SCI, peer-reviewed

Schilthuizen M. 2000. Dualism and conflicts in understanding speciation. BioEssays 22: 1134-1141.

Schilthuizen M. 2000. Ecotone: speciation-prone. Trends in Ecology & Evolution 15: 130-131.

Schilthuizen M. 2000. Bimodal hybrid zones and the scale of a snail. Trends in Ecology & Evolution 15: 469-469.


Journals SCI, peer-reviewed

Schilthuizen M. 1999. Synchronised sex - How a jolt to the body clock can drive evolution. New Scientist 161: 15-15.

Schilthuizen M. 1999. Cloning Odysseus and the seed of speciation. Trends in Ecology & Evolution 14: 90-91.

Schilthuizen M., Hoekstra R.F., Gittenberger E. 1999. Selective increase of a rare haplotype in a land snail hybrid zone. Proceedings of the Royal Society B: Biological Sciences 266: 2181-2185.

Schilthuizen M., Vermeulen J.J., Davison G.W.H., Gittenberger E. 1999. Population structure in a snail species from isolated Malaysian limestone hills, inferred from ribosomal DNA sequences. Malacologia 41: 283-296.


Journals SCI, peer-reviewed

Schilthuizen M., Gittenberger E. 1998. Screening mollusks for Wolbachia infection. Journal of Invertebrate Pathology 71: 268-270.

Schilthuizen M., Honda J., Stouthamer R. 1998. Parthenogenesis-inducing Wolbachia in Trichogramma kaykai (Hymenoptera : Trichogrammatidae) originates from a single infection. Annals of the Entomological Society of America 91: 410-414.

Schilthuizen M., Nordlander G., Stouthamer R., Alphen J.J.M. van 1998. Morphological and molecular phylogenetics in the genus Leptopilina (Hymenoptera : Cynipoidea : Eucoilidae). Systematic Entomology 23: 253-264.

Schilthuizen M., Stouthamer R. 1998. Distribution of Wolbachia among the guild associated with the parthenogenetic gall wasp Diplolepis rosae. Heredity 81: 270-274.

Hurst G.D.D., Schilthuizen M. 1998. Selfish genetic elements and speciation. Heredity 80: 2-8.

Jochemsen P., Schilthuizen M., Stouthamer R. 1998. Transmission of Wolbachia between Trichogramma species and their lepidopteran hosts?. Proceedings of the section Experimental and Applied Entomology of the Netherlands Entomological Society (NEV) 9: 131-136.


Journals SCI, peer-reviewed

Schilthuizen M., Stouthamer R. 1997. Horizontal transmission of parthenogenesis-inducing microbes in Trichogramma wasps. Proceedings of the Royal Society B: Biological Sciences 264: 361-366.


Journals SCI, peer-reviewed

Kan F.J.P.M. van, Silva I.M.M.S., Schilthuizen M., Pinto J.D., Stouthamer R. 1996. Use of DNA-based methods for the identification of minute wasps of the genus Trichogramma. Proceedings of the section Experimental and Applied Entomology of the Netherlands Entomological Society (NEV) 7: 233-237.


Journals SCI, peer-reviewed

Schilthuizen M., Lombaerts M. 1995. Life on the edge - A hybrid zone in Albinaria hippolyti (Gastropoda, Clausiliidae) from Crete. Biological Journal of the Linnean Society 54: 111-138.

Schilthuizen M. 1995. A comparative study of hybrid zones in the polytypic land snail Albinaria hippolyti (Gastropoda Pulmonata: Clausiliidae). Netherlands Journal of Zoology 45: 261-290.

Schilthuizen M., Gittenberger E., Gultyaev A.P. 1995. Phylogenetic relationships inferred from the sequence and secondary structure of ITS1 rRNA in Albinaria and putative Isabellaria species (Gastropoda, Pulmonata, Clausiliidae). Molecular Phylogenetics and Evolution 4: 457-462.


Journals SCI, peer-reviewed

Schilthuizen M., Lombaerts M. 1994. Population structure and levels of gene flow in the Mediterranean and snail Albinaria corrugata (Pulmonata, Clausiliidae). Evolution 48: 577-586.

Huis A. van, Arendse P.W., Schilthuizen M., Wiegers P.P., Heering H., Hulshof M., Kaashoek N.K. 1994. Uscana lariophaga, egg parasitoid of bruchid beetle storage pests of cowpea in West-Africa - the effect of temperature and humidity. Entomologia Experimentalis et Applicata 70: 41-53.

Schilthuizen M., Gittenberger E. 1994. Parallel evolution of an sAat-hybrizyme in hybrid zones in Albinaria hippolyti (Boettger). Heredity 73: 244-248.

Schilthuizen M. 1994. Reproductive isolation in snails of the genus Albinaria (Gastropoda, Clausiliidae). Biological Journal of the Linnean Society 52: 317-324.


Journals SCI, peer-reviewed

Schilthuizen M. 1992. Allozyme variation in Cretan Albinaria (Gastropoda Pulmonata: Clausiliidae). Belgian Journal of Botany 125: 306-309.