Prof. Dr. M. (Menno) Schilthuizen
Permanent research scientist

- schilthuizen@naturalis.nl
- Phone
- +31 (0)71 5687769
I work in the fields of evolutionary biology, systematics, and ecology, mostly studying tropical and mediterranean land snails, although I also work with parasitic wasps, leiodid scavenging beetles, birds, and the bacterial symbiont Wolbachia. I hold a position as research scientist at Naturalis, and as extraordinary professor for Insect Biodiversity at the University of Groningen. I also hold a research associateship at the Institute for Tropical Biology and Conservation at Universiti Malaysia Sabah, where I worked as an associate professor from 2000 until 2006. (See: here, here, and here for more information on the collection, labs, and research programme I set up there.) A list of scientific publications is given below. In addition to my scientific work, I have been active as a popular science writer since 1994. I write on ecology and evolution for Science, ScienceNOW, and Natural History, the Malaysian national newspaper The Star and for several Dutch newspapers and magazines, such as NRC Handelsblad, Bionieuws, de Volkskrant, and Intermediair. I have also written a book on speciation, entitled: Frogs, Flies & Dandelions; the Making of Species (Oxford University Press; translated into French and Dutch) and a book on the ecology of biodiversity, entitled: The Loom of Life: Unravelling Ecosystems (Springer Verlag). Most of my articles and columns can be found on the Popular Science Writing page. For reviews of the latest book, see:
http://noorderlicht.vpro.nl/artikelen/40161066/ (Item on the web site of VPRO's Noorderlicht)
http://www.nature.com/nature/journal/v455/n7217/full/4551178b.html (book review in Nature)
http://cgi.omroep.nl/cgi-bin/streams?/radio1/vara/vroegevogels/20090104-08.wma?start=0:5:15.0 (VARA's Vroege Vogels Radio item)



Research interests
My chief scientific interest is in the origin and maintenance of biodiversity; i.e., speciation, phylogenetics, and species coexistence. I have attempted to be broadly oriented both taxonomically and in methods of primary investigation. I have studied terrestrial gastropods, Coleoptera, parasitic Hymenoptera, bacterial symbionts, and birds, in various parts of the world, namely Southeast Asia, the Netherlands, and the Mediterranean. I have done and published primary research in alpha-taxonomy, anatomy, morphometrics, a range of protein and DNA techniques, population genetics, phylogenetic reconstruction, behavioural observation, computer modelling, and field experiments on gene flow and selection. In addition, I have published intensively in popular science writing, which has forced me to broaden my knowledge outside of the groups and methods I use in my own research.
(Click here for an overview of the terrestrial gastropods of North Borneo.)
I am also the Dutch coordinator for the Evolution MegaLab, an interactive website to observe evolution in nature, as part of the international Darwin Year 2009. Click here to go to the official webpage for the Evolution MegaLab, or here for more information on the activities in the Netherlands.
Currently, I am working on the research projects as detailed below. Within each of these, smaller research projects ("stages") for students (Bachelor, Master, or "HBO") are available. I also occasionally help students design a project that is tailor-made to their wishes, provided it fits within one of my research interests. A list of available project synopsis can be found at Naturalis "Students" page.
The 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 as great a morphological diversity as do the instruments in a brass band, including some wonderful shapes, as shown by these Bornean species of Opisthostoma (Diplommatinidae), drawn by Jaap Vermeulen. Many species are endemic to single outcrops, and are sufficiently endangered to have been placed on the IUCN Red List. I have been working on the conservation ecology of these obligate limestone-dwelling species, including their population sizes, degree of inter-karst geneflow, and responses to habitat disturbance. I have also worked on the speciation and micro-evolution of shell shape in geographically isolated populations of Opisthostoma, which revealed that these snails and their main predator (a slug of the genus Atopos) are engaged in a Red Queen style arms race.
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Speciation where worlds collide
I am particularly interested in speciation-with-geneflow (‘parapatric speciation') across ecotones (sharp boundaries between adjacent, but ecologically very different habitats). Field situations that I am studying are speciation across the interface between the bright, warm, biodiverse, and productive tropical rainforest on limestone outcrops and the dark, cool, impoverished, and unproductive habitat inside the caves that are inside these outcrops. For example, I found that Georissa filiasaulae cave snails are direct descendants of G. saulae, which live on the outside of the outcrop, and are connected to the latter by a transition zone in the cave entrance. I also work on transition zones in snails, birds, and beetles along elevational gradients in Mount Kinabalu and other tropical mountains, and I am planning to expand this work to artificial ecotones in the Tropics, for example those between primary forest and plantations.
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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.
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Evolution after Invasion
I lead a project at the University of Groningen on adaptation of native herbivorous insects to introduced plants. Started in early 2008, the project aims to identify cases of normally oligophagous phytophagous insects that feed on woody plants without related native species (usually, trees and shrubs from other continents and/or genera with no European representatives). Once identified, the populations on the new host plant will be screened for morphometric or genetic differentiation from the population on the native host plant, to look for incipient sympatric speciation. This project is mainly carried out by PhD student Kim Meijer. An information sheet (in Dutch) on the project can be found here. I also have recently started a project on the evolution of invasiveness in the land snail family Succineidae. This project is run by PhD student Pongrat Dumrongrajwattatna.
Publications
Naturalis RepositorySelected publications
1. Research papers
Sheldon, F.H., J. Nail, M. Lakim, A. Tuuga, P. Malim, J. Majuakim, A. Lo, M. Schilthuizen, P.A. Hosner & R.G. Moyle, 2009. Observations on the distribution, ecology, and systematics of forest birds in Sabah, Malaysia. Raffles Bulletin of Zoology, 57: 577-586.
Schilthuizen, M. & S. Looijestijn, 2009. The sexology of the chirally dimorphic snail species Amphidromus inversus. Malacologia 51: 379-387.
Clements, R., T.S. Liew, M. Schilthuizen & J.J. Vermeulen, 2008. Further twists in gastropod shell evolution. Biology Letters, 4: 179-182,
Clements, R., X.X. Lu, S. Ambu, M. Schilthuizen & C. Bradshaw, 2008. Using biogeographical patterns of endemic land snails to improve conservation planning for limestone karsts. Biological Conservation, 141: 2751-2764.
Liew, T.S., R. Clements & M. Schilthuizen, 2008. Sampling micromolluscs in tropical forests: one size does not fit all. Zoosymposia, 1: 271-280.
Gomes, S.R., J.B. Picanço, M. Schilthuizen & J.W. Thomé, 2008. An endemic Veronicellid from Sundaland: redescription of Valiguna flava (Heynemann, 1885) (Gastropoda, Soleolifera). The Veliger, 50: 163-170.
Haase, M. & M. Schilthuizen, 2007. A new Georissa (Gastropoda, Neritopsida, Hydrocenidae) from a limestone cave in Malaysian Borneo. Journal of Molluscan Studies, 73: 215-221.
Benedick, S., T.A. White, J.B. Searle, K.C. Hamer, Nazirah Mustaffa, Chey Vun Khen, Maryati Mohamed, M. Schilthuizen & J.K. Hill, 2007. Impacts of habitat fragmentation on genetic diversity in a tropical forest butterfly on Borneo. Journal of Tropical Ecology, 23: 623-634, doi: 10.1017/S0266467407004543
Blakemore, R.J., C. Csuzdi, M.T. Ito, N. Kaneko, T. Kawaguchi, T. & M. Schilthuizen, 2007. Taxonomic status and ecology of Oriental Pheretima darnleiensis (Fletcher, 1886) and other earthworms (Oligochaeta : Megascolecidae) from Mt Kinabalu, Borneo. Zootaxa, 1613: 23-44.
Schilthuizen, M., P.G. Craze, A.S. Cabanban, A. Davison, E. Gittenberger, J. Stone & B.J. Scott, 2007. Sexual selection maintains whole-body chiral dimorphism. Journal of Evolutionary Biology, 20: 1941-1949.
Schilthuizen, M., A. van Til, M. Salverda, T.-S. Liew, S.S. James, B. Elahan & J.J. Vermeulen, 2006. Micro-allopatric divergence in a snail associated with behavioural differences in its predator. Evolution, 60: 1851-1858.
Craze, P.G., B. Elahan & M. Schilthuizen, 2006. Spatial ecology of opposite shell-coiling morphs in a tropical land snail. Ecography, 29: 477-486.
Benedick, S., J.K. Hill, Nazirah Mustaffa, Chey Vun Khen, Maryati Mohamed, J.B. Searle, M. Schilthuizen & K. C. Hamer, 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.
Uit de weerd, D.R., D.S.J. Groenenberg, M. Schilthuizen & E. Gittenberger, 2006. Reproductive character displacement by inversion of coiling in clausiliid snails (Gastropoda, Pulmonata). Biological Journal of the Linnean Society, 88:155-164.
Schilthuizen, M. B.J. Scott, A.S. Cabanban & P.G. Craze, 2005. Population structure and coil dimorphism in a tropical land snail. Heredity, 95: 216-220.
Moyle, R., M. Schilthuizen, M. Abdul Rahman, & F.H. Sheldon, 2005. Molecular phylogenetic analysis of the White-crowned forktail (Enicurus leschenaulti) in Borneo. Journal of Avian Biology, 36: 96-101.
Schilthuizen, M., T.-S. Liew, B. Elahan & I. Lackman-Ancrenaz, 2005. Effects of karst forest degradation on Pulmonate and Prosobranch land snail communities in Sabah, Malaysian Borneo. Conservation Biology, 19: 949-954.
Schilthuizen, M., A.S. Cabanban & M. Haase, 2005. Possible speciation with gene flow in tropical cave snails. Journal of Zoological Systematics and Evolutionary Research, 43: 133-138.
Charlat, S., E.A. Hornett, E.A. Dyson, P.H.Y. Ho, N.T. Loc, M. Schilthuizen, N. Davies, G.K. Roderick & G.D.D. Hurst, 2005. Is extreme male-killer prevalence a local or common event in the butterfly Hypolimnas bolina? A survey across Indo-Pacific populations. Molecular Ecology, 14: 3525-3530.
Schilthuizen, M., E. Gutteling, F.W. Welter-Schultes, C.H.M. van Moorsel, M. Haase & E. Gittenberger, 2004. Phylogeography of the land snail Albinaria hippolyti from Crete, based on ITS-1 sequences. Biological Journal of the Linnean Society, 83: 317-326.
Schilthuizen, M., H.-N. Chai, T.E. Kimsin & J.J. Vermeulen, 2003. Abundance and diversity of land snails on limestone in Borneo. Raffles Bulletin of Zoology 51: 35-42.Schilthuizen, M., M.I.F. Teräväinen, Noor Faezamoltha Kartini Tawith, Hamidah Ibrahim, Sim Mei Chea, Chong Pow Chuan, Leonardo Jeffery Daim, Amirzaidi Jubaidi, Mohammed Juni Madjapuni, Muhafizar Sabeki & Adnan Mokhtar, 2002. Microsnails at microscales in Borneo: distributions of Prosobranchia versus Pulmonata. Journal of Molluscan Studies, 68: 259-262.
Schilthuizen, M. & H.A. Rutjes, 2001. Land snail diversity in a square kilometre of tropical rainforest in Sabah, Malaysian Borneo. Journal of Molluscan Studies, 67: 417-423.
Schilthuizen, M., R.F. Hoekstra & E. Gittenberger, 2001. The “rare allele phenomenon” in a ribosomal spacer. Molecular Ecology, 10: 1341-1345.
Schilthuizen, M., R.F. Hoekstra & E. Gittenberger, 1999. Selective increase of a rare haplotype in a land snail hybrid zone. Proceedings of the Royal Society of London B, 266: 2181-2185.
Schilthuizen, M., J.J. Vermeulen, G.W.H. Davison & E. Gittenberger, 1999. Population structure in a snail species from isolated Malaysian limestone hills, inferred from ribosomal DNA sequences. Malacologia 41: 271-284.
Schilthuizen, M. & R. Stouthamer, 1998. Distribution of Wolbachia among an assemblage associated with the parthenogenetic gall wasp Diplolepis rosae. Heredity, 81: 270-274.
Schilthuizen, M., J. Honda & R. Stouthamer, 1998. The parthenogenesis‑inducing Wolbachia in a Californian Trichogramma species originates from a single infection. Annals of the Entomological Society of America, 91: 410-414.Hurst, G.D.D. & M. Schilthuizen, 1998. Selfish genetic elements and speciation. Heredity, 80: 2-8.
Schilthuizen, M. & E. Gittenberger, 1998. Screening mollusks for Wolbachia infection. Journal of Invertebrate Pathology, 71: 268-270.
Schilthuizen, M., G. Nordlander, J.J.M. van Alphen & R. Stouthamer, 1998. Morphological and molecular phylogenetics in the genus Leptopilina (Hymenoptera: Cynipoidea: Eucoilidae). Systematic Entomology, 23: 253-264.
Schilthuizen, M. & R. Stouthamer, 1997. Horizontal transmission of parthenogenesis‑inducing symbionts in Trichogramma. Proceedings of the Royal Society of London, series B, 264: 361-366.
Schilthuizen, M. E. Gittenberger & A. Gultyaev, 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.
Schilthuizen, M. & M. Lombaerts, 1995. Life on the edge: a hybrid zone in Albinaria hippolyti 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 (Pulmonata: Clausiliidae). Netherlands Journal of Zoology, 45: 261‑290.Schilthuizen, M. & E. Gittenberger, 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. Biological Journal of the Linnean Society, 52: 317‑324.Schilthuizen, M. & M. Lombaerts, 1994. Population structure and levels of gene flow in the Mediterranean land snail Albinaria corrugata (Pulmonata: Clausiliidae). Evolution, 48: 577‑586.
2. Book chapters
Schilthuizen, M. & B.J. Scott, 2003. Allopatric speciation: not so simple after all. Book chapter in Evolution: From Molecules to Ecosystems (A. Moya & E. Font, eds.). Oxford University Press.
3. Opinion papers and (book) reviews
Clements, R., N.S. Sodhi, M. Schilthuizen & P. Ng, 2006. Karsts of Southeast Asia: neglected and imperiled arks of biodiversity. BioScience, 56: 733-742.
Schilthuizen, M. & A. Davison, 2005. The convoluted evolution of snail chirality. Naturwissenschaften, 92: 504-515.
Schilthuizen, M., 2005. The darting game in snails and slugs. Trends in Ecology and Evolution, 20: 581-584.
Schilthuizen, M., 2005. On the origin of reproductive isolation. BioEssays, 27: 669-670.Schilthuizen, M., 2003. Sexual selection on land snail shell ornamentation: a hypothesis that may explain shell diversity. BMC Evolutionary Biology, 2003, 3: 13.
Schilthuizen, M., 2002. A grand old synthesizer's overview. Science, 295: 50.
Schilthuizen, M., 2000. Dualism and conflicts in understanding speciation. BioEssays, 22: 1134-1141.
Schilthuizen, M., 2000. Ecotone--speciation prone. Trends in Ecology and Evolution, 15: 130-131.
Schilthuizen, M., 1999. Cloning Odysseus and the seed of speciation. Trends in Ecology and Evolution, 14: 90-91.
Hurst, G.D.D. & M. Schilthuizen, 1998. Selfish genetic elements and speciation. Heredity, 80: 2-8.
4. Taxonomic and faunistic papers and books
Schilthuizen, M. & H.J. Vallenduuk, 1998. Kevers op Kadavers. Wetenschappelijke Mededelingen KNNV, 222: 1-150.
Gittenberger, E. & M. Schilthuizen, 1992. The earliest name in Albinaria Vest clarified after two centuries. Basteria, 56: 159-161.
Schilthuizen, M. & E. Gittenberger, 1991. A new Albinaria species from Central Crete (Gastropoda Pulmonata: Clausiliidae). Basteria, 55: 111‑114.
Schilthuizen, M. & E. Gittenberger, 1990. Zur Formenbildung und Verbreitung der Albinaria‑Arten von Kreta; neue Taxa der "Gruppe der candida" (Gastropoda Pulmonata: Clausiliidae). Basteria, 54: 131‑142.
5. Popular articles
Below are links to a small selection of popular science articles I have written. See my personal website for full-text content of my more than 220 newspaper and magazine articles in Dutch and in English.
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Synchronised sex; how a jolt to the body clock can drive evolution (16 January 1999)
When the biological clocks of males and females are out of sync, their sex lives suffer. Or at least it does if they are melon flies, Japanese entomologists have found. They say that differences in daily rhythms might even promote the evolution of new species...
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A Paradox to Everyone but Himself (September 2004)
Last month my students and I took a field trip to a small forest reserve a couple of miles from our university campus in Malaysian Borneo. Slip-sliding down a steep jungle path, clutching the soggy stems of wild yams in a futile attempt to stay upright, we collapsed into a pebbly streambed...
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Dutch ecology under attack (Science, 292: 1055)
In a cost-saving move, Leiden University in the Netherlands has proposed excising five sections--including two internationally prominent research groups...
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The race for solid semen (24 November 2003)
Chimps are the most notorious swingers among the great apes. Their wanton sex lives, in which males compete to impregnate females, have led males to evolve huge testicles, three times the size of humans'...
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Miljoenen torren wachten aan Zwitserse grens (17 augustus 1996)
Eén document moet er nog worden getekend, dan verhuist de befaamde kevercollectie-Frey van Duitsland naar Zwitserland. De strijd in de familie Frey is beslecht...
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Gepassioneerd zoeker naar de waarheid (19 januari 1996)
Hij geldt als een van de grootste biologen van deze eeuw en op zijn vijfenzeventigste heeft hij nog niets van zijn scherpte verloren: de Britse evolutiebioloog John Maynard Smith. 'Sinds mijn vijftiende besef ik dat ik slechts het produkt van natuurlijke selectie ben. En ik ben daar gelukkig mee.'...
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Wormen (16 februari 1995)
'Mensen, houdt die wurm goed onder water, anders droogt uw preparaat uit en dan kunt u het weggooien', schalt de stem van Gerrit Anker door de luidsprekers...
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Menno Schilthuizen is ziek van geesten (15 oktober 2004)
Een doordeweekse namiddag een tijdje geleden. Het is buiten al aan het schemeren en dan ineens: grote consternatie op de gang van het instituut. Firdaus heeft een spook gezien!...
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De ring gebroken (20 november 2004)
De ringsoort van het zilvermeeuw-complex. Hele generaties biologen zijn ermee opgegroeid, maar na DNA-analyse blijkt dit schoolvoorbeeld van de evolutiebiologie niet te kloppen. "Er blijft helemaal niks van over."...
Giants of the mountain (17 August 2004)
Legend has it that a giant dragon dwells on the summit of Mount Kinabalu. Kinabalu, at 4,095 m the tallest peak in Southeast Asia, is climbed by thousands of hikers each year, and the dragon remains undiscovered...
Thursday, January 7, 2010

