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Ton de Winter

Dr. A.J. (Ton) de Winter, Senior researcher


Email: ton.dewinter@naturalis.nl
Phone: +31 (0)71-56 87 567
Room number: C03.07, Darwinweg 2


Research interest

My research interests are the evolution, systematics, biogeography and ecology of non-marine gastropods (snails and slugs). Within this broad field, I presently focus on the land snails inhabiting the Afrotropical region (the western part in particular), but other geographic areas are also occasionally touched, for phylogenetic and other comparative reasons, or by stochastic causes. The western Palaearctic non-marine Molluscan fauna has been and remains for me an area of interest.


ecology, evolution, biogeography, non-marine gastropods, snails, slugs, systematics

Current research topics


Systematics of African land snail taxa from Senegal to Northern Angola, and from the Gulf of Guinea Islands to 15° Eastern longitude. Preparation of annotated and illustrated list of taxa is the first step towards a faunal overview. Numerous new taxa are awaiting description, but this can only be done after evaluation of the taxa described in the past.

Revisionary work on African land snails is often largely based on shell characters, since material with conserved soft parts is often not available. But careful study of shells is also indispensible since virtually all type specimens are empty shells. The classification of many, if not most, African snails is thus somewhat tentative. A more thorough classification requires study of the morphology of the soft parts ánd molecular data, for which sustained collection effort is required. Apart from revisionary work on partcular taxa, also faunistic inventories are being prepared, for which species IDs are often based on ad-hoc taxonomic decisions, many species being assigned to informal morphospecies. Matching land snail distribution patterns with those of plant species (Dr. J.J. Wieringa) may assist in answering biogeographic questions.

The following Afrotropical groups are currently being studied:

1. Helicarionoidea of western Africa (fully-shelled taxa, semislugs and slugs).

The systematic position of shelled helicarioid snails is still at its infancy. A first revision of Thapsia (De Winter, 2008) showed that even large shells that are very similar in size, shape and sculpture, belong to different genera with very different reproductive anatomies. This work is far from finished. The larger group of taxa referred to as Trochozonites is anatomically at least as heterogeneous, and a revision of this group would be desirable. For the slugs and semislugs of West Africa two large revisions are available (Van Mol, 1970; Van Goethem, 1976), but a supervicial scrutiny of recently collected material indicates that extensive updates are necessary in most groups.


2. Western African Streptaxidae (Gulella s.l., Ptychotrema s.l., Streptostele s.l., Gonaxis s.l., Edentulina).

This family in Western Africa embraces many hundrerds of species that are all carnivorous, and represents one of the two most speciose radiations in tropical Africa (with the vegetarian Subulinidae). Their shells are often quite distinctive in shape, sculpture, size and complexity of apertural barriers. Although quite a large numbers of species have been named in the past, the number of recently discovered species may be at least as large or greater in some groups. Also at generic level much revisionary work needs to be done, many taxa are only known by their shells. Here micro CT-scanning is used for revealing internal shell structures that are otherwise not readily visible from an intact shell.


3. Other Western African groups

Almost every land snail family is in need of extensive taxonomic revision. These include the Subulinidae, possibly the largest family in Western Africa, of which the genus level taxonomy is rather confused and species are difficult to delimit. But also smaller groups like Maizaniidae, Cyclophoridae, Euconulidae, Veronicellid slugs, Cerastidae, and various endodontoid groups, among others, harbour numerous novelties at all levels, and are dealt with when an appropriate occasion arises. Even the giant African snail family Achatinidae is still insufficiently known.


Land snails of the Netherlands (With E. Gittenberger)

A new systematic overview and identification tool of the land snails occuring in the Netherlands is long overdue. The last book covering the Dutch land snail fauna dates back to 1984. The Dutch fauna presently includes a substantial number of species that were discovered since 1984, mostly non-native species, like Arion lusitanicus/vulgaris, Xerotricha apicina, Testacella haliotidea, but also overlookedindigeneous species like Balea heydeni and Arion fuscus. New taxa, mostly of southern origin, continue to be discovered. Not all taxonomic issues relating to the Dutch land snail fauna are solved as yet, taxa like Euconulus sp., Arion s.s, Deroceras sp. and Oxyloma sp., among others, require further study.




Supervision MSc Students


co-supervisor - P.A.J. Bakker - Evolution and diversification in euconulid-shaped snails

2013. supervisor - Björn Elzer - Radula morphology of the land snail Arianta arbustorum in relation to function, habitata and geographic distribution.

2012. supervisor - Dominique Derogee - Conchological variation of Nesopupa bisulcata in relation to environmental factors.

2011 supervisor - Nienke Vastenhout - Systematics and biogeography of the Afrotropical land snail genus Avakubia (Gastropoda, Streptaxidae).

Supervision MSc Students

2014 Supervisor - P.A.J. Bakker - Evolution and diversification in euconulid-shaped snails
Available student projects



Journals SCI, peer-reviewed

winter A.J. de 2017. “Dwarfs among pygmies”: the genus Punctum in western Africa and the Congo Basin, with descriptions of two new species and notes on other Afrotropical punctoids (Gastropoda, Eupulmonata: Punctidae) . Archiv für Molluskenkunde 146: 203-211.
Go to website (DOI)

Donovan S.K., Fearnhead F.E., Winter A.J. de 2017. Two rare taxa from the type area of the Devonian, south-west England. Proceedings of the Geologists' Association 128: 675-678.
Go to website (DOI)

Journals non-SCI, peer-reviewed

Gittenberger E., Leda P., Winter A.J. de, Jochum A. 2017. First record of Carychium in Bhutan (Gastropoda, Ellobioidea). Basteria 81: 107-110.


Journals non-SCI, peer-reviewed

Winter A.J. de, Leeuwen S. van, Hovestadt A. 2016. A new species of Glyphyalus (Gastropoda, Pulmonata, Oxychilidae) from the Dutch Caribbean island of St. Eustatius. Basteria 80: 39-46.

Breure A.S.H., Gittenberger E., Winter A.J. de 2016. In memoriam Dr. A.C. van Bruggen 1929-2016. Basteria 80: 161-170.


Journals SCI, peer-reviewed

Jochum A., Winter A.J. de, Weigand A.M., Gómez B., Prieto C. 2015. Two new species of Zospeum Bourguignat, 1856 from the Basque-Cantabrian Mountains, Northern Spain (Eupulmonata, Ellobioidea, Carychiidae). ZooKeys 483: 81-96.
Go to website (URI)


Journals SCI, peer-reviewed

Winter A.J. de, Vastenhout N. 2013. Revision of the Afrotropical land snail genus Avakubia Pilsbry, 1919, with description of Pseudavakubia gen.n. and eleven new species (Gastropoda: Pulmonata: Streptaxidae). African Invertebrates 54: 605-663.
Go to website (URI)


Journals non-SCI, peer-reviewed

Winter A.J. de, Nieulande F.A.D. van 2011. Testacella haliotidea Draparnaud, 1801 in The Netherlands (Gastropoda Pulmonata, Testacellidae). Basteria 75: 11-21.


Journals SCI, peer-reviewed

Soes D.M., Glöer P., Winter A.J. de 2009. Viviparus acerosus (Bourguignat, 1862) (Gastropoda: Viviparidae), a new exotic snail species for the Dutch fauna. Aquatic Invasions 4: 373-375.

Journals non-SCI, peer-reviewed

Winter A.J. de, Cremers H.J.W.M., Soes D.M. 2009. The Asian tramp snail Bradybaena similaris in a tropical greenhouse in Arnhem, The Netherlands. Basteria 73: 61-64.

Breure A.S.H., Gittenberger E., Maassen W.J.M., Winter A.J. de 2009. In and out of Africa: Dr A.C. van Bruggen, keen educator and eminent biologist. Zoologische Mededelingen 83: 509-524.

Winter A.J. de 2009. A new species of the operculate land snail genus Maizaniella from Liberia (Gastropoda, Caenogastropoda). Zoologische Mededelingen 83: 661-666.

Edited books

Breure A.S.H., Smeenk C., Winter A.J. de (eds.). 2009. Festschrift for Dr. A.C. van Bruggen on the occasion of his 80th birthday. Zoologische Mededelingen 83: 267 p. Naturalis, Leiden, The Netherlands.


Journals non-SCI, peer-reviewed

Winter A.J. de 2008. Redefinition of Thapsia Albers, 1860, and the description of three new helicarionoid genera from western Africa (Gastropoda, Stylommatophora). Zoologische Mededelingen 82: 441-478.

Winter A.J. de 2008. Costigulella primennilus spec. nov., a new minute western African terrestrial snail, with remarks on the genus Costigulella (Gastropoda Pulmonata: Streptaxidae). Zoologische Mededelingen 82: 245-252.

Winter A.J. de 2008. Notes on Parennea species from western Africa, including descriptions of two new species (Gastropoda, Pulmonata, Streptaxidae). Basteria 72: 215-222.


Journals non-SCI, peer-reviewed

Winter A.J. de 2006. Gulella (Silvigulella?) kuiperi spec. nov., a new minute land snail from Cameroon (Gastropoda, Pulmonata, Streptaxidae). Basteria Supplement 3: 69-72.


Journals non-SCI, peer-reviewed

Soes D.M., Winter A.J. de 2005. Xerotricha apicina (Lamarck, 1822), een nieuwe immigrant in Nederland (Gastropoda, Pulmonata, Hygromiidae). Xerotricha apicina (Lamarck, 1822), a new immigrant land snail in the Netherlands. Basteria 69: 81-86.


Journals SCI, peer-reviewed

Pinceel J., Jordaens K., Houtte N. van, Winter A.J. de, Backeljau T. 2004. Molecular and morphological data reveal cryptic taxonomic diversity in the terrestrial slug complex Arion subfuscus/fuscus (Mollusca, Pulmonata, Arionidae) in continental north-west Europe. Biological Journal of the Linnean Society 83: 23-38.

Journals non-SCI, peer-reviewed

Boesveld A., Winter A.J. de 2004. Oxychilus (Ortizius) navarricus helveticus (Blum, 1881), een nieuwe landslak voor de Nederlandse fauna (Gastropoda, Pulmonata, Zonitidae). Oxychilus (Ortizius) navarricus helveticus (Blum, 1881), a new land snail for Dutch fauna (Gastropoda, Pulmonata, Zonitidae. Basteria 69: 1-6.


Journals non-SCI, peer-reviewed

Bruggen A.C. van, Winter A.J. de 2003. Studies on the Streptaxidae (Mollusca, Gastropoda Pulmonata) of Malawi 8. A revision of ‘Marconia’ hamiltoni (Smith), the largest local streptaxid, with the description of a new genus. Zoologische Verhandelingen 345: 59-78.


Chapters in books

Winter A.J. de 2002. Notes on land operculates from Cameroon (Gastropoda: Cyclophoridae and Maizaniidae), including the description of two new species. 237-251:


Chapters in books

Winter A.J. de 2001. Grauwe wegslak/Bos-wegslak Arion circumscriptus (complex); Zwarte wegslak/Donkere wegslak Arion hortensis (complex); Egel-wegslak Arion intermedius. 82-84:


Journals non-SCI, peer-reviewed

Gittenberger E., Goud J., Maassen W.J.M., Menkhorst H.P.M.G., Ripken Th.E.J., Winter A.J. de, Bank R.A. 2000. Comment on the proposed conservation of Trichia Hartmann, 1840 (Mollusca, Gastropoda) and proposed emendation of spelling of Trichiinae Lozek, 1956 (Mollusca) to Trichiainae, so removing the homonymy with Trichiidae Fleming, 1821 (Insecta, Coleoptera). Bulletin of Zoological Nomenclature 57: 166-167.


Journals SCI, peer-reviewed

Winter A.J. de, Gomez B.J., Prieto C.E. 1999. Sinistrexcisa, a new genus of land snail from Central West Africa with four new species (Gastropoda : Pulmonata : Streptaxidae). Journal of Molluscan Studies 65: 209-221.

Journals non-SCI, peer-reviewed

Boeters H.D., Falkner G., Gittenberger A., Winter A.J. de, Proschwitz T. von, Ripken Th.E.J. 1999. Comments on the proposed conservation of Hydrobia Hartmann, 1821 (Mollusca, Gastropoda) and Cyclostoma acuta Draparnaud, 1805 (currently Hydrobia acuta) by the replacement of the lectotype of H. acuta with a neotype; proposed designation of Turbo ventrosus Montagu, 1803 as the type species of Ventrosia Radoman, 1977; and propsed emendation of spelling of Hydrobiina Mulsant, 1844 (Insecta, Coleoptera) to Hydrobiusina, so removing the homonymy with Hydrobiidae Troschel, 1857 (Mollusca). Bulletin of Zoological Nomenclature 56: 56-62.


Journals SCI, peer-reviewed

Winter A.J. de, Gittenberger E. 1998. The land snail fauna of a square kilometer patch of rainforest in southwestern Cameroon, high species richness, low abundance and seasonal fluctuations. Malacologia 40: 231-250.

Journals non-SCI, peer-reviewed

Winter A.J. de, Vermeulen J.J. 1998. A first record of Truncatellina (Pulmonata: Vertiginidae) in SE. Asia: Truncatellina cf. insulivaga on Java, Indonesia. Basteria 62: 269-271.


Journals non-SCI, peer-reviewed

Winter A.J. de 1997. A giant specimen of Archachatina marginata (Gastropoda Pulmonata: Achatinidae). Basteria 61: 41-42.

Winter A.J. de 1997. Limax flavus L. synanthropical in Madagascar (Gastropoda Pulmonata: Limacidae). Basteria 61: 40.


Journals non-SCI, peer-reviewed

Winter A.J. de 1996. Six new species of Streptaxidae (Gastropoda Pulmonata) from West Africa. Basteria 60: 139-148.

Winter A.J. de 1996. Miscellaneous notes on West African Maizaniella (Prosobranchia: Maizaniidae). Basteria 59: 91-95.


Journals SCI, peer-reviewed

Winter A.J. de 1995. Genetic control and evolution of acoustic signals in planthoppers (Homoptera, Delphacidae). Researches on Population Ecology 37: 99-104.

Journals non-SCI, peer-reviewed

Bruggen A.C. van, Winter A.J. de 1995. Notes on Micractaeon, a monotypic genus of African land snails (Gastropoda Pulmonata: Ferussaciidae?). Zoologische Mededelingen 69: 79-92.


Journals SCI, peer-reviewed

Backeljau T., Winter A.J. de, Martin R., Rodriguez T., Bruyn L. de 1994. Genital and allozyme similarity between Arion urbiae and A. anguloi (Mollusca, Pulmonata). Zoological Journal of the Linnean Society 110: 1-18.


Journals SCI, peer-reviewed

Winter A.J. de, Rollenhagen T. 1993. Differences in preference for species-specific female calls between acoustically experienced and acoustically naive male Ribautodelphax planthoppers (Homoptera, Delphacidae). Journal of Insect Behavior 6: 411-419.


Journals SCI, peer-reviewed

Winter A.J. de 1992. The genetic basis and evolution of acoustic mate recognition signals in a Ribautodelphax planthopper (Homoptera, Delphacidae): 1. The female call. Journal of Evolutionary Biology 5: 249-265.

Winter A.J. de, Bruggen A.C. van 1992. Systematic position and distribution of the African land snails Afropunctum seminium (Morelet) and 'Guppya' rumrutiensis (Preston) (Mollusca, Gastropoda Pulmonata: Euconulidae), with the description of two new taxa. Proceedings of the Koninklijke Nederlandse Akademie van Wetenschappen: Biological, Chemical, Geological 95: 515-533.

Journals non-SCI, peer-reviewed

Winter A.J. de, Maassen W.J.M. 1992. Two new Hygromiidae from Crna Gora and Kosovo (Gastropoda Pulmonata), with a checklist of the hygromiid species described from the western Balkan. Basteria 56: 173-192.

Winter A.J. de 1992. Genetic studies on acoustic differentiation in Ribautodelphax planthoppers (Homoptera, Delphacidae). Proceedings of the section Experimental and Applied Entomology of the Netherlands Entomological Society (NEV) 3: 116-120.


Journals non-SCI, peer-reviewed

Winter A.J. de 1991. Deroceras rodnae in eastern France (Pulmonata, Agriolimacidae). Basteria 55: 138.

Winter A.J. de, Ripken Th.E.J. 1991. New land snails from the Iberian Peninsula. Basteria 55: 3-7.


Journals SCI, peer-reviewed

Bruggen A.C. van, Winter A.J. de 1990. Carinazingis regalis nov. gen. nov. spec. from Malawi, an urocyclid snail with a frontal organ and not belonging to Gymnarion (Mollusca, Gastropoda, Pulmonata, Urocyclidae). Proceedings of the Koninklijke Nederlandse Akademie van Wetenschappen: Biological, Chemical, Geological 93: 235-252.

Winter A.J. de 1990. On some small West African land snails (Gastropoda, Pulmonata). Journal of Conchology 33: 305-309.

Winter A.J. de, Rollenhagen T. 1990. The importance of male and female acoustic behaviour for reproductive isolation in Ribautodelphax planthoppers (Homoptera: Delphacidae). Biological Journal of the Linnean Society 40: 191-206.

Journals non-SCI, peer-reviewed

Winter A.J. de 1990. Little known land snails from the French Alpes (Pulmonata). Basteria 54: 227-237.


Journals non-SCI, peer-reviewed

Winter A.J. de 1989. Remarks on the non-marine molluscan fauna of the Azores. 3. A new species of Drouetia from the Isle of Sao Miguel (Pulmonata: Zonitidae). Basteria 53: 63-67.

Winter A.J. de, Ripken Th.E.J. 1989. Leiostyla anglica (Wood) on Madeira (Gastropoda Pulmonata, Pupillidae). Basteria 53: 47-48.

Winter A.J. de 1989. Arion lusitanicus Mabille in Nederland (Gastropoda, Pulmonata, Arionidae). Basteria 53: 49-51.

Winter A.J. de 1989. New records of Achatina fulica Bowdich from the Côte d'Ivoire. Basteria 53: 71-72.


Journals non-SCI, peer-reviewed

Winter A.J. de 1988. Remarks on the non-marine molluscan fauna of the Azores, 1. A new Punctum (Pulmonata, Punctidae) from the Azores. 2. Notes on Deroceras (Pulmonata, Agriolimacidae) from the Azores. Basteria 52: 105-109.

Winter A.J. de 1988. Achatina fulica in West Africa. Basteria 52: 2.


Journals SCI, peer-reviewed

Backeljau T., Winter A.J. de 1987. An electrophoretic characterization of 3 paratypes of Arion fagophilus De Winter, 1986, with notes on the subgeneric division of the genus Arion ferussac, 1819 (Mollusca, Pulmonata). Zeitschrift für Zoologische Systematik und Evolutionsforschung 25: 169-180.

Journals non-SCI, peer-reviewed

Winter A.J. de 1987. Verrucarion demeryi, a new semi-slug from Sierra Leone (Mollusca: Pulmonata: Urocyclidae). Basteria 51: 87-93.

Winter A.J. de, Visser G.J.M. 1987. Deroceras agreste op Terschelling (Gastropoda, Pulmonata, Agriolimacidae). Basteria 51: 53-56.


Journals non-SCI, peer-reviewed

Winter A.J. de, Butot L.J.M. 1986. Three new Deroceras species from Greece (Gastropoda, Pulmonata, Agriolimacidae). Basteria 50: 35-44.

Winter A.J. de 1986. Little known and new South-West European slugs (Pulmonata: Agriolimacidae, Arionidae). Zoologische Mededelingen 60: 135-158.


Journals non-SCI, peer-reviewed

Winter A.J. de 1985. Two new semi-slugs from Gabon, western Africa (Gastropoda Pulmonata: Urocyclidae). Zoologische Mededelingen 59: 209-220.

Winter A.J. de 1985. A new rapid method for the relaxation and killing of slugs. Basteria 49: 71-72.

Gittenberger E., Winter A.J. de 1985. A Pyrenean Monacha species, Monacha (Metatheba) atacis spec. nov. (Mollusca: Gastropoda: Pulmonata). Zoologische Mededelingen 59: 197-207.

Winter A.J., Butot L.J.M. 1985. On some slugs from the Isle of Andros, Greece, including the descriptions of two new species (Gastropoda: Pulmonata). Bulletin Zoölogisch Museum, Universiteit van Amsterdam 10: 77-85.

Winter A.J. de 1985. A new Deroceras species from north-western Spain (Gastropoda: Pulmonata: Agriolimacidae). Zoologische Mededelingen 59: 69-77.


Journals non-SCI, peer-reviewed

Winter A.J. de 1984. The Arion hortensis complex: Designation of types, descriptions, and distribution patterns, with special reference to the Netherlands. Zoologische Mededelingen 59: 1-17.


Journals SCI, peer-reviewed

Heinstra P.W.H., Eisses K.T., Schoonen W.G.E.J., Aben W., Winter A.J. de, Horst D.J. van der, Marrewijk W.J.A. van, Beenakkers A.M.T., Scharloo W., Thorig G.E.W. 1983. A dual function of alcohol dehydrogenase in Drosophila. Genetica 60: 129-137.

Boeters H.D., Winter A.J. de 1983. Neues über Avenionia Nicolas 1882 (Prosobranchia: Hydrobiidae). Archiv für Molluskenkunde 114: 25-30.

Journals non-SCI, peer-reviewed

Notenboom J., Winter A.J. de 1983. Avenionia brevis roberti Boeters (Prosobranchia, Hydrobiidae) in the Netherlands, with notes on its habitat. Basteria 47: 149-153.

Winter A.J. de 1983. Euphaedusa cumingiana (L. Pfeiffer) (Pulmonata, Clausiliidae) new to the fauna of Java (Indonesia). Basteria 47: 59-60.


Journals non-SCI, peer-reviewed

Gittenberger E., Winter A.J. de 1980. Nieuwe gegevens over in Nederland gevonden naaktslakken. Basteria 44: 71-76.

Thapsia sp., copula. (Helicarionoidea, shell diameter 25 mm)

Trochozonites talcosus (Gould) (Helicarionoidea, shell height 9 mm)

"Gulella" kuiperi De Winter, 2006 (Streptaxidae) (shell height  2.5 mm; drawing I. van Noortwijk)

Urocyclid slug  from Liberia (photo Carel Jongkind)

Avakubia acuminata (Thiele) (Streptaxidae, shell height 5.3 mm; drawing Inge van Noortwijk)

Gulella” jongkindi  de Winter (Streptaxidae, shell height 5.2 mm; drawing Bas Blankenvoort)

Testacella haliotidea