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Barbara Gravendeel

Dr. B. (Barbara) Gravendeel, senior researcher and joint group leader Endless Forms, Biodiversity chair


Email: barbara.gravendeel@naturalis.nl
Phone: +31 (0)71 527 1910
Room number: 6.4.15 B, Sylvius Laboratory, Sylviusweg 72, 2333 BE, Leiden
“Museum collections help me to experience a world stretching far beyond my own life by recreating the past and launching dreams of the future.”

At a very young age, I was already stimulated to collect my first specimens for the local history museum of Vleuten - De Meern (now Leidsche Rijn) by my dad and mom who were passionate historians. They taught me that musea are magical places opening up a billion of years of earth’s history and keys to answers of challenges that we struggle with today rather than dusty attics to store things that must be kept even though no one uses them anymore.

Today, I continue in my parent's footsteps by applying modern techniques to historical collections. These analyses help me to answer questions about the origin and extinction of species including mankind. Maintaining a collection of over 37 million specimens is very expensive. I therefore increasingly collaborate with companies interested in the development of innovative products from museum specimens sometimes collected over 400 years ago!

One of the five million accessions in the herbarium of Naturalis 


I studied Biology at Utrecht University, where I graduated in 1997. At Leiden University, a PhD degree in molecular systematic research on Asian orchids was obtained in 2000. After securing a visiting lectureship at Harvard University with a FULBRIGHT grant, I became a researcher at the National Herbarium of The Netherlands in 2004 with a Netherlands Organisation for Scientific Research (NWO) MEERVOUD subsidy. From 2011 onwards, I have a joint Biodiversity chair at the University of Applied Sciences Leiden and Naturalis Biodiversity Center. Research for this chair focuses on the use of museum specimens for answering questions posed by companies and organisations. Major grants obtained so far were obtained from NWO (promotiebeurs voor docenten 023.003.015), Stichting Innovatie Alliantie (RAAK.MKB04.005 and RAAK.MKB07.021) and EU H2020 (Marie Curie Innovative Training Network Plant.ID 765000).  


Research interest

Recent advances in DNA sequencing technologies have initiated a new era of genomics. With the help of Next Generation Sequencing (NGS), more detailed data can now be produced in a much shorter timeframe. NGS has therefore taken several different biological disciplines to another level. I work in several of these.

In the field of genomics, we can now detect crucial low copy genes coding for adaptive phenotypic traits. Massive parallel sequencing for bioinventories of extant taxa reveal new details about ecological niches. In the field of palaeogenomics, NGS enables us to produce increasingly detailed reconstructions of both the palaeoenviroment and palaeobiology of fossil species to solve the riddle why they went extinct. The fields of plant conservation and wildlife forensics benefit from new phylogenetic and phylogeographic markers found. We can now trace the taxonomic identity and geographic origin of dis- or reappearing species illegally traded or expanding their range.


museomics, micro ct, next generation sequencing, ancient dna, wildlife forensics

Current research topics

Ancient Tomato Genomics

En Tibi herbarium of tomato 

Past processes that shaped the current genetic diversity of the domesticated tomato are still largely unknown. Using Illumina HiSeq, we sequenced and compared full genomes from tomato accessions stored worldwide in genebanks and herbaria up to 182 years ago.

The aim was to gain more insight in the bottlenecks that narrowed genetic diversity after domestication in the Andes and subsequent industrial breeding in Europe and the US.

As a result, chloroplast, mitochondrial and nuclear genomes were recovered with moderate to high read depth by mapping reads on the fully annotated genome of the inbred Heinz 1706 cultivar. Accelerated and relaxed levels of evolution of genes coding for pathogen resistance, shape, taste and ripening of tomatoes were recovered with HyPhy. Increasing patterns of de-amination over time were visualised with mapDamage.

The conclusion of this project was that museomics proved to be an effective tool for discovering genetic diversity lost during the breeding process.

Applied research

Vanilla pods analyzed in RAAK MKB project

Microbiome of Vanilla 

DNA extracts of fermented fruits of species of Vanilla are currently being sequenced to obtain an impression of their microbiome, i.e. the diversity of fungi and bacteria living inside. The diversity of the microbiome is very different among vanille fruits produced in tropical rainforests, plantations and under glass. This project is a collaboration between Naturalis Biodiversity Center, the University of Applied Sciences Leiden, Wageningen UR and several plant breeders, a soil improvement company, a chemistry company, a DNA sequencing company and a spice consultant.


Bio-inventories by massive parallel DNA barcoding

  • Bas van Geel (IBED, Amsterdam University)
  • Peter de Knijff, Letty de Weger (LUMC, Leiden University)
  • Dick Mol (Mammuthus Club International)

Genomics and Museomics

  • Mark Chase and Mike Fay (Royal Botanic Gardens Kew, U.K.)
  • Gunter Fischer (Kadoorie Farm & Botanic Garden, HongKong)
  • Peter Klinkhamer (IBL, Leiden University)
  • Philipp Schluter (University of Zurich, Switzerland)

Wildlife Forensics

  • Esther de Kok (RIKILT)
  • Tamara Peelen (Dutch Customs Laboratory, Amsterdam)

Supervision PhDs



Current promotores that I co-supervise PhD students with are:

  • prof. Sandra L. Baldauf 

  • prof. Erik F. Smets



Current PhD students and postdocs under my (co-)supervision are:




I teach the following topics at UL, UvA, Hogeschool Amsterdam and Hogeschool Leiden:

  • Bio-inventories by massive parallel DNA barcoding
  • Genomics and Museomics
  • Wildlife Forensics
Available student projects

Public outreach

Scientific activities

  • Nov. 12th 2013: Lectorale rede 'Biodiversiteit als bron van innovatie' at University of Applied Sciences, Leiden.

Invited lectures

  • March 2018: The orchid genomic toolkit, 18th EOCCE, Paris, France.
  • August 2017: Added value of 'museomes': genomes preserved in musea, BioSystEU, Goteborg.
  • May 2017: Molecular phylogenetics of African Bulbophyllinae (Orchidaceae), AETFAT congress, Nairobi.
  • January 2017: Museomics: uncovering the secrets of Vanilla flavors, 1st Generade symposium, Leiden. 
  • December 2016: Next Generation Sequencing to unravel genome evolution, 1st WURomics conference, Wageningen.
  • August 2016: An integrative approach to orchidology: linking fossils, phylogenies and pollinators. Shanghai Botanical Garden, China.
  • May 2016: Below ground fungal diversity of helleborine orchids, 6th International Orchid Conservation Conference, HongKong.
  • April 2016: Determineren met DNA, STOWA congres, Arnhem.
  • October 2015: Metabarcoding for evolutionary studies, Nature of Life Seminars, VU Amsterdam.
  • February 2015: eDNA from fresh water, NAEM conference, Lunteren.
  • November 2014: Towards sustainable orchid cultivation in Africa, Dar es Salaam, Tanzania.
  • February 2014: Metabarcoding of Pleistocene dung, Tromso, Norway.
  • February 2014: Fighting wildlife crime with museum collections, Frontiers of Forensic Science, Amsterdam University.
  • December 2013: DNA barcoding helps monitoring illegal orchid trade, 5th International Orchid Conservation Conference, Reunion.

Public lectures

  • Evolutionaire oorsprong van orchideenbloemen, februari 2018, Werkgroep Europese Orchideeen, Maarn.
  • Overleeft DNA? MuseumJeugdUniversiteit, december 2017, Rotterdam.
  • Toepassing eDNA voor onderzoek aan invasieve plantensoorten, STOWA, juni 2017, Driebergen.
  • New species evolve, others go extinct. BeeVee symposium, maart 2017, Nijmegen.
  • Gaat eDNA het huidige waterkwaliteitsonderzoek veranderen? ILOW symposium, februari 2017, Utrecht.
  • Bospest door evolutie ingehaald. Floron dag, december 2016, Nijmegen.
  • DNA uit herbariumcollecties: wat kun je er allemaal mee? Gorteria symposium, november 2016, Leiden.
  • Orchidologie in de 21ste eeuw. Nederlandse Orchideeen Vereniging, september 2016, Utrecht.
  • Toegevoegde waarde van onderzoek aan natuurhistorische collecties. Rotterdams Natuurkundig Genootschap, maart 2016, Rotterdam.
  • DNA barcoding van nieuwe Schroeforchissen in Nederland en Belgie. Werkgroep Europese Orchideeen, februari 2016, Maarn.
  • DNA analyse van historische Gorilla collectes. Museum Boerhaave, september 2015, Leiden.
  • Bestuiving van Kleine keverorchis ingewikkelder dan gedacht. Studiegroep Europese en Mediterrane Orchideeen, november 2015, Meise.
  • Terug naar het Beekbergerwoud. Symposium Nieuwe Wildernis, september 2014, Lelystad.
  • Wildlife Forensics. Hanzehogeschool Groningen, maart 2014, Groningen.
  • Hoe behouden we Veenmosorchis voor Nederland? Vereniging Orchideeen Vermeerdering, maart 2014, Utrecht.
  • Van mestschimmel tot mammoetsteppe, Werkgroep Pleistocene Zoogdieren, april 2012, Den Haag. 


  • Parool mei 2018: Hoop op Hollandse hop via moleculaire veredeling
  • OVT VPRO april 2018: Herboren mammoet
  • Nieuwe oogst februari 2018: Groene planten en innovatieve teelten
  • Dag van de vrouwen maart 2017: Barbara Gravendeel in de voetsporen van Johanna Westerdijk
  • Mare december 2016: Vleeseters als vlaggenschip
  • Algemeen Dagblad 'Vincent luistert' oktober 2016: Hondsbrutale exoot
  • Vroege Vogels oktober 2016: Nieuwe schroeforchissen
  • Mare september 2016: Waarom zijn zij nog geen prof?
  • Vroege Vogels mei 2016: Amerikaanse vogelkers aantrekkelijk voor kevers
  • Bionieuws mei 2016: Insecten houden van Bospest
  • Kennislink april 2016: In de voetsporen van Hannibals olifanten
  • Volkskrant februari 2016: 14 tot 45 miljoen jaar oud plantenfossiel ontdekt
  • Volkskrant januari 2016: Pestbacterie veel hardnekkiger dan gedacht
  • NPO1 januari 2016: Stropers misbruiken wetenschappelijke publicaties
  • EenVandaag december 2015: Vandaag 25 jaar geleden - stier Herman  
  • EenVandaag oktober 2015: Het nut van ijstijdpoep
  • Mare september 2015: Steeds vaker onderzoek naar museaal DNA
  • Leids Dagblad augustus 2015: Aap van papier-mache leefde ooit echt
  • Noordhollands Dagblad augustus 2015: Nieuw leven voor uitstervend fruit
  • Telegraaf juli 2015: Uitgestorven diersoorten: uit de dood herrezen?
  • NPO radio interview juni 2015: De Kennis van Nu
  • Volkskrant mei 2015: Gorilla-mache
  • Discovery News april 2015: orchid lip formation
  • Onderzoek aan mammoetbloed: live NOS radio Eenvandaag uitzendig 24 december 2014
  • Pieter in de Prehistorie oktober 2014: bijdrage aan VPRO TV uitzending over de mammoet
  • RTV Noord Holland juni 2014: radio interview forensisch DNA onderzoek
  • Leidraad mei 2014: Survival of the fittest - klimaatverandering en wilde plantensoorten
  • NTR (radio) februari 2014: Ontwikkelen van medische hulpmiddelen uit inhoudsstoffen
  • Trouw december 2013: Leren van oude drollen
  • Vara’s Vroege Vogels (radio) 24 november 2013: Fossiele drol voor Mol
  • ZAAAMspraak november 2013: Holland Rijnland Biobased inspireert via regional TV
  • Volkskrant augustus 2013: terug uit de dood: een mammoet klonen, kan het nu wel?
  • Vara’s Vroege Vogels augustus 2013: de wolf is terug
  • Omroep Flevoland (tv): Wolf op eigen kracht uit Oost-Europa
  • NOS 18.00 en 20.00 uur Journaal, Nieuwsuur en Een vandaag  (tv)
  • RTL nieuws (tv): Hoe lang wolf hier was blijft onduidelijk
  • EenVandaag: De wolf is terug. Waar komt hij vandaan?
  • Volkskrant: Wolf Luttelgeest komt uit Oost-Europa
  • NRC: Wolf kwam op eigen houtje uit Oost-Europa naar Nederland
  • Trouw: Als er één wolf is volgen er wel meer
  • Telegraaf: Meer duidelijkheid over Nederlandse wolf
  • Resource juni 2013: herbariumplant geeft geheimen prijs
  • BNR radio april 2013: uitgestorven soorten tot leven wekken met DNA uit museumcollecties.
  • Leids Dagblad jan 2013: Schatkamer: de oudste tomaat.
  • Hortus Vriendennieuws 2012: Moderne biologie - experimenten en teamwerk
  • NatuurWetenschap&Techniek online 12 okt 2012: Houdbaarheidsdatum DNA bepaald 
  • Caught - A Small Marvel at the Smithsonian Institution's Orchid Exhibit. Orchids 2012
  • NatuurWetenschap&Techniek mei 2012: taxonomie is booming business
  • NRC 4 feb 2012: dode mammoet komt tot leven
  • Leids Dagblad 2011: nieuwe Leidse lector Biodiversiteit aan de slag 
  • UnityFM 2011: radio interview over lectoraat Biodiversiteit
  • VPRO Labyrinth 2011: pitch LeveDNA!
  • Mare 2010: Forensische houtkunde
  • Cahier Stichting Bio-Wetenschappen Darwin & Genomics 2009: kader over phylogenomics
  • Mare 2008: Mammoetmenu bestaat uit poep en madeliefjes 
  • Pre-University College 2008: buiten de hokjesgeest durven denken
  • Jaarverslag 2007 Universiteit Leiden: Biologen ontdekken fossiele orchidee
  • BBC News 2007: Orchids date to time of the dinos
  • Discovery Channel 2007: Orchid Fossil Solves Age-Old Mystery
  • Bionieuws 2007: Fossiel toont leeftijd van orchidee
  • NRC Handelsblad 2007: Barnsteenbij dateert orchidee
  • Bionieuws 2007: Mammoetmaag toont hoe steppe toendra werd
  • NRC Handelsblad 2005: de Heukels gehusseld met DNA
  • Mare 2004: Genen schrapen uit fossielen
  • Volkskrant 2004: De stokoude vrouw is een joekel
  • Volkskrant 2002: Een woud van regels
  • Bionieuws 2001: Darwins triomf is Linnaeus’ failliet


Hogeschool Leiden, lectoraten (in Dutch)

Europese orchideeën, KNNV (in Dutch)

LeveDNA (in Dutch)



Journals SCI, peer-reviewed

Langeveld B.W., Mol D., Zazula G.D., Gravendeel B., Eurlings M., McMichael C.N.H., Groenenberg D., Reenen G.B.A. van, Palmeira M., Vogel J., Geel B. van 2018. A multidisciplinary study of a late Pleistocene arctic ground squirrel (Urocitellus parryii) midden from Yukon, Canada. Quaternary Research 89: 333-351.
Go to website (DOI)

Bogarin D., Fernandez M., Borkent A., Heemskerk A., Pupulin A., Ramirez S., Smets E.F., Gravendeel B. 2018. Pollination of Trichosalpinx (Orchidaceae: Pleurothallidinae) by biting midges (Diptera: Ceratopogonidae). Botanical Journal of the Linnean Society 186: 510-543.
Go to website (DOI)

Bogarín D., Perez-Escobar O.A., Groenenberg D., Holland S.D., Karremans A.P., Lemmon E.M., Lemmon A.R., Pupulin F., Smets E., Gravendeel B. 2018. Anchored hybrid enrichment generated nuclear, plastid and mitochondrial markers resolve the Lepanthes horrida (Orchidaceae: Pleurothallidinae) species complex. Molecular Phylogenetics and Evolution 129: 27-47.
Go to website (DOI)

Lee Y.I., Yap J.W., Shairul I., Leitch I.J., Fay M.F., Lee Y.C., Hidalgo O., Dodsworth S., Smulders M.J.M., Gravendeel B., Leitch A.R. 2018. Satellite DNA in Paphiopedilum subgenus Parvisepalum as revealed by high-throughput sequencing and fluorescent in situ hybridization. BMC Genomics 19: 578.
Go to website (DOI)


Journals SCI, peer-reviewed

Dirks-Mulder A., Butot R., Schaik P. van, Wijnands J.W.P.M., Berg R. van den, Krol L., Doebar S., Kooperen K. van, Boer H. de, Kramer E.M., Smets E.F., Vos R.A., Vrijdaghs A., Gravendeel B. 2017. Exploring the evolutionary origin of floral organs of Erycina pusilla, an emerging orchid model system. BMC Evolutionary Biology 17: 89.
Go to website (DOI)

Ghorbani A., Gravendeel B., Selliah S., Zarre S., Boer, H. de 2017. DNA barcoding of tuberous Orchidoideae: a resource for identification of orchids used in Salep. Molecular Ecology Resources 17: 342-352.

Schiebold J.M.I., Bidartondo M.I., Karasch P., Gravendeel B., Gebauer G. 2017. You are what you get from your fungi: nitrogen stable isotope patterns in Epipactis species. Annals of Botany 119: 1085-1095.
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Schwallier R., Gravendeel B., de Boer H., van Heuven B.J., Sieder A., Sumail S., van Vugt R., Nylinder S., Lens F. 2017. Evolution of wood anatomical characters in Nepenthes and close relatives of Caryophyllales. Annals of Botany 119: 1179-1193.
Go to website (DOI)

Scopece G., Gravendeel B., Cozzolino S. 2017. The effect of different chiral morphs on visitation rates and fruit set in the orchid Spiranthes spiralis. Plant Ecology & Diversity 10: 97-104.
Go to website (DOI)

Boer H.J. de, Ghorbani A., Manzanilla V., Raclariu A.C., Kreziou A., Ounjai S., Osathanunkul M., Gravendeel B. 2017. DNA metabarcoding of orchid-derived products reveals widespread illegal orchid trade. Proceedings of the Royal Society B: Biological Sciences 284
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Veldman S., Gravendeel B., Otieno J.N., Lammers Y., Duijm E., Nieman A., Bytebier A., Ngugi G., Martos F., Andel T. van, Boer H.J. de 2017. High-throughput sequencing of African orchid cake highlights conservation challenges in orchids. Biodiversity and Conservation 26: 2029-2046.
Go to website (DOI)

Gravendeel B. 2017. Tales of a plant twitcher. Nature Plants 3: 908.
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Journals non-SCI, peer-reviewed

Gravendeel B., Noort F. van, Vugt R. van 2017. Duurzame productie van salep. Orchideeën 79: 90-92.


Journals SCI, peer-reviewed

Bakker F.T., Lei D., Yu J., Mohammadin S., Wei Z., van de Kerke S., Gravendeel B., Niewenhuis M., Staats M., Alguezar-Planas D.E., Holmer R. 2016. Herbarium genomics: plastome sequence assembly from a range of herbarium specimens using an Interactive Organelle Genome Assembly (IOGA) pipeline. Biological Journal of the Linnean Society 117: 33-43.
Go to website (DOI)

Schwallier R., Raes N., de Boer H.J., Vos R.A., van Vugt R., Gravendeel B. 2016. Phylogenetic analysis of niche divergence reveals distinct evolutionary histories and climate change implications for tropical carnivorous pitcher plants . Diversity and Distributions 22: 97-110.
Go to website (DOI)

Horn I.R., Rijn M. van, Zwetsloot T.J.J., Basmagi S., Dirks-Mulder A., Leeuwen W.B. van, Ravensberg W.J., Gravendeel B. 2016. Development of a multiplex Q-PCR to detect Trichoderma harzianum Rifai strain T22 in plant roots. Journal of Microbiological Methods 121: 44-49.
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Staats M., Arulandu A.J., Gravendeel B., Holst-Jensen A., Scholtens I., Peelen T., Prins T.W., Kok E. 2016. Advances in DNA metabarcoding for food and wildlife forensic species identification. Analytical and Bioanalytical Chemistry 408: 4615-4630.
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Haarsma A.J., Siepel H., Gravendeel B. 2016. Added value of metabarcoding combined with microscopy for evolutionary studies of mammals. Zoologica Scripta 45: 37-49.
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Karremans A.P., Albertazzi F.J., Bakker F.J., Bogarin D., Eurlings M.C.M., Pridgeon A., Pupulin F., Gravendeel B. 2016. Phylogenetic reassessment of Specklinia and its allied genera in the Pleurothallidinae (Orchidaceae). Phytotaxa 272: 1-36.
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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.
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Journals non-SCI, peer-reviewed

Claessens J., Ulenberg S., Gravendeel B. 2016. Torymus arcticus (Torymidae), ein neuer Bestäuber für Chamorchis alpina. Journal Europaischer Orchideen 48: 3-10.

Gravendeel, B. 2016. Nieuwe schroeforchissen in Nederland en Belgie. De Levende Natuur 117: 203-206.

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

Bogarín D., Pupulin F., Smets E., Gravendeel B. 2016. Evolutionary diversification and historical biogeography of the Orchidaceae in Central America with emphasis on Costa Rica and Panama. Lankesteriana 16: 189-200.
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Journals SCI, peer-reviewed

Schwallier R., Boer H.J. de, Visser N., Vugt R.R. van, Gravendeel B. 2015. Traps as treats: a traditional sticky rice snack persisting in rapidly changing Asian kitchens. Journal of Ethnobiology and Ethnomedicine 11: 1-9.
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Karremans A.P., Bogarin D., Gravendeel B. 2015. Specklinia lugduno-batavae (Orchidaceae), a new species in the S. digitalis group. Blumea: Journal of Plant Taxonomy and Plant Geography 59: 180-184.
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Kreziou A., Boer H. de, Gravendeel B. 2015. Harvesting of salep orchids in north-western Greece continues to threaten natural populations. Oryx 49: 1-4.
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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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Kok E.T., Jong M.J., Gravendeel B., van Leeuwen W.B., Baars E.W. 2015 2015. Resistance of Antibiotics and Antifungal Medicinal Products: Can Complementary and Alternative Medicine Help Solve the Problem in Common Infection Diseases? The introduction of a Dutch Research Consortium. Evidence-Based Complementary and Alternative Medicine: ID 52158.

Karremans A.P., Pupulin F., Gravendeel B. 2015. Specklinia dunstervillei, a new species long confused with Specklinia endotrachys (Orchidaceae: Pleurothallidinae). PLOS ONE 10: e0131971.

Karremans A.P., Bogarin D., Pupulin F., Luer C.A., Gravendeel B. 2015. The glandulous Specklinia: morphological convergence versus phylogenetic divergence. Phytotaxa 218: 101-127.
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Sulistyo B.P., Boos R., Cootes J.E., Gravendeel B. 2015. Dendrochilum hampelii (Coelogyninae, Epidendroideae, Orchidaceae) traded as ‘Big Pink’ is a new species, not a hybrid: evidence from nrITS, matK and ycf1 sequence data. PhytoKeys 56: 83-97.
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De Boer H.J., Cross H.B., de Wilde W.J.J.O., Duyfjes-de Wilde B.E.E. & Gravendeel B. 2015. Molecular phylogenetic analyses of Cucurbitaceae tribe Benincaseae urge for merging of Pilogyne with Zehneria. Phytotaxa 236: 173-183.

Karremans A.P., Pupulin F., Grimaldi D., Beentjes K.K., Butôt R., Fazzi G.E., Kaspers K., Kruizinga J., Roessingh P., Smets E.F., Gravendeel B. 2015. Pollination of Specklinia by nectar-feeding Drosophila: the first reported case of a deceptive syndrome employing aggregation pheromones in Orchidaceae. Annals of Botany 116: 437-455.
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Journals non-SCI, peer-reviewed

Gravendeel B., Dirks-Mulder A. 2015. Floral development: Lip formation in orchids unravelled. Nature Plants 1: 15056.
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Claessens J., Beentjes K.K., Heijerman T., Miller J., Gravendeel, B. 2015. Beobachtungen von Miarus campanulae als Bestäuber von Cephalanthera rubra. Journal Europaischer Orchideen 47: 77-87.


Journals SCI, peer-reviewed

Welker F., Duijm E., Gaag K. van der, Geel B. van, Knijff P. de, Leeuwen J. van, Mol D., Plicht J. van der, Raes N., Reumer J., Gravendeel B. 2014. Analysis of coprolites from the extinct mountain goat Myotragus balearicus. Quaternary Research 81: 106-116.
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Geml J., Gravendeel B., Neilen M., Lammers Y., Raes N., Semenova T.A., Noordeloos M.E. 2014. The Contribution of DNA Metabarcoding to Fungal Conservation: Diversity Assessment, Habitat Partitioning and Mapping Red-Listed Fungi in Protected Coastal Salix repens Communities in the Netherlands. PLOS ONE 9: e99852.
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Lammers Y., Peelen T., Vos R.A., Gravendeel B. 2014. The HTS barcode checker pipeline, a tool for automated detection of illegally traded species from high-throughput sequencing data. BMC Bioinformatics 15: 44.
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Geel B. van, Protopopov A., Bull I., Duijm E., Gill F., Nieman A., Rudaya N., Trofimova S., Tikhonov A., Vos R.A., Zhilich S., Gravendeel B. 2014. Multiproxy diet analysis of the last meal of an early Holocene Yakutian Bison. Journal of Quaternary Science 29: 261-268.
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Aflitos S., Schijlen E., Finkers R., Smit S., Wang J., Zhang G., Li N., Mao L., Jong H. de, Bakker F., Gravendeel B., Breit T., Dirks R., Huits H., Struss D., Swanson-Wagner R., Leeuwen H. van, Ham R.C.H.J. van, Fito L., Guignier L., Sevilla M., Ellul P., Ganko E., Kapur A., Reclus E., Geus B. de, Geest H. van de, Lintel Hekkert B. te, Haarst J. van, Smits L., Koops A., Sanchez-Perez G., Heusden A.W. van, Visser R., Quan Z., Min J., Liao L., Wang X., Wang G., Yue Z., Yang X., Xu N., Schranz E., Smets E., Vos R.A., Rauwerda J., Ursem R., Schuit C., Kerns M., Berg J. van den, Vriezen W., Janssen A., Jahrmann T., Moquet F., Bonnet J., Peters S. 2014. Exploring genetic variation in the tomato (Solanum section Lycopersicon) clade by whole-genome sequencing. Plant Journal 80: 136-148.
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Gravendeel B., Protopopov A., Bull I., Duijm E., Gill F., Nieman A., Rudaya N., Tikhonov A.N., Trofimova S., Reenen G.B.A. van, Vos R., Zhilich., Geel B. van 2014. Multiproxy study of the last meal of a mid-Holocene Oyogos Yar horse, Sakha Republic, Russia. Holocene 24: 1-9.
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Ghorbani A., Gravendeel B., Naghibi F., Boer H.J. de 2014. Wild orchid tuber collection in Iran: a wake-up call for conservation. Biodiversity and Conservation 23: 2749-2760.
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Journals non-SCI, peer-reviewed

Weeda E.J., Bijlsma R.J., Gravendeel B. 2014. Wat voor oerwoud was het Beekbergerwoud. Nieuwe Veluwe 3: 28-30.

Veldman S., Otieno J., Andel T.R. van, Gravendeel B., Boer H.J. de 2014. Efforts urged to tackle thriving illegal orchid trade in Tanzania and Zambia for chikanda production. Traffic Bulletin 26: 47-50.

Ervynck A., Strydonck M. van, Boudin M., Gravendeel B., Randi E., Fabbri E., Caniglia R., Ottoni C., Thieren E., Vuyst W. de, Lievois D., Neer W. van 2014. De poot van de wolf aan de poort van het Steen (deel 2). Radiokoolstofdatering en genetisch onderzoek van een historisch curiosum. Handelingen der Maatschappij voor Geschiedenis & Oudheidkunde 68: 3-11.

Gravendeel B., Noordeloos M., Geml J. 2014. Nieuwe inzichten in de samenstelling van de paddestoelenflora in Kruipwilgstruweel dankzij DNA analyse. Holland’s Duinen 64: 22-25.

Ghorbani A., Gravendeel B., Zarre S., Boer H. de 2014. Illegal wild collection and international trade of CITES-listed terrestrial orchid tubers in Iran. Traffic Bulletin 26: 18-24.

Chapters in books

Gravendeel B., Fischer G., Camargo Smidt E.S. de, Sieder A., Vugt R.R. van, Vermeulen J.J. 2014. Dendrobieae: 621. Bulbophyllum. Genera Orchidacearum 6: ISBN 9780199646517.

Veldman S., Otieno J., Gravendeel B., Andel T. van, Boer H. de 2014. Conservation of Endangered Wild Harvested Medicinal Plants: Use of DNA Barcoding. Novel Plant Bioresources: Applications in Food, Medicine and Cosmetics Wiley & Sons, London.

Gravendeel B., Gischer F., Schmidt E.S., Sieder A., Vugt R.R. van, Vermeulen J. 2014. Bulbophyllinae.


Journals SCI, peer-reviewed

Eurlings M.C.M., Lens F., Pakusza C., Peelen T., Wieringa J.J., Gravendeel B. 2013. Forensic identification of Indian Snakeroot (Rauvolfia serpentina Benth. ex Kurz) using DNA barcoding. Journal of Forensic Sciences 58: 822-830.
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Bogarin D., Karremans A.P., Rincon R., Gravendeel B. 2013. A new Specklinia (Orchidaceae: Pleurothallidinae) from Costa Rica and Panama. Phytotaxa 115: 31-41.

Subedi A., Kunwar B., Choi Y., Dai Y., Andel T. van, Chaudhary R.P., Boer H.J. de, Gravendeel B. 2013. Collection and trade of wild-harvested orchids in Nepal. Journal of Ethnobiology and Ethnomedicine 9: 64.
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Welker F., Gravendeel B. 2013. Dung reveals goat's last days. Nature 503: 440.

Hof K. van den, Marcussen T., Berg R.G. van den, Weeda E.J., Gravendeel B. 2013. Fenotypische plasticiteit en nomenclatuur van Melkviooltje (Viola stagnina - V. persicifolia nom.ambig. propos.). Gorteria: Tijdschrift voor de Floristiek 36: 60-84.

Gravendeel B., Groot A. de, Kik M., Beentjes K.K., Bergman H., Caniglia R., Cremers H., Fabbri E., Groenenberg D., Grone A., Groot Bruinderink G., Font L., Hakhof J., Harms V., Jansman H., Janssen R., Lammertsma D., Laros I., Linnartz L., Marel D. van der, Mulder J.L., Mije S. van der, Nieman A.M., Nowak C., Randi E., Rijks M., Speksnijder A., Vonhof H.B. 2013. The first wolf found in the Netherlands in 150 years was the victim of a wildlife crime. Lutra 56: 93-109.

Gathier G., Peelen T., Vugt R.R. van, Niet T. van der, Gravendeel B. 2013. Forensic identification of CITES protected slimming Cactus (Hoodia gordonii) in traditional African Medicines (TAMs) using DNA barcoding. Journal of Forensic Sciences 58: 1467-1471.
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Journals SCI, peer-reviewed

Sirichamorn Y., Adema F.A.C.B., Gravendeel B., Welzen P.C. van 2012. Phylogeny of palaeotropic Derris-like taxa (Fabaceae) based on chloroplast and nuclear DNA sequences shows reorganization of (infra)generic classifications is needed. American Journal of Botany 99: 1793-1808.
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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.

Pupulin F., Karremans A.P., Gravendeel B. 2012. A reconsideration of the empusellous species of Specklinia (Orchidaceae: Pleurothallidinae) in Costa Rica. Phytotaxa 63: 1-20.

Przybylowicz T., Roessingh P., Groot A.T., Biesmeijer J.C., Oostermeijer J.G.B., Chittka L., Gravendeel B. 2012. Possible chemical mimicry of the European lady's slipper orchid (Cypripedium calceolus). Contributions to Zoology 81: 103-110.

Chapters in books

Gravendeel B., Przybylowicz T. 2012. Bijen en orchideeen: waarom bedriegen zo slecht nog niet is.


Journals SCI, peer-reviewed

Smidt E.C., Borba E.L., Gravendel B., Fischer G.A., Berg C. van den 2011. Molecular phylogeny of the Neotropical sections of Bulbophyllum (Orchidaceae) using nuclear and plastid spacers. Taxon 60: 1050-1064.

Subedi A., Chaudhary R.P., Vermeulen J.J., Gravendeel B. 2011. Panisea panchaseensis sp nov (Orchidaceae) from central Nepal. Nordic Journal of Botany 29: 361-365.
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Subedi A., Chaudhary R.P., Achterberg C. van, Heijerman Th., Lens F., Dooren T.J.M. van, Gravendeel B. 2011. Pollination and protection against herbivory of Nepalese Coelogyninae (Orchidaceae). American Journal of Botany 98: 1095-1103.
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Geel B. van, Guthrie R.D., Altmann J.G., Broekens P., Bull I.D., Gill F.L., Jansen B., Nieman A.M., Gravendeel B. 2011. Mycological evidence of coprophagy from the feces of an Alaskan Late Glacial mammoth. Quaternary Science Reviews 30: 2289-2303.
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Doorduin L., Gravendeel B., Lammers Y., Ariyurek Y., Chin-A-Woeng T., Vrieling K. 2011. The complete chloroplast genome of 17 individuals of pest species Jacobaea vulgaris: SNPs, microsatellites and barcoding markers for population and phylogenetic studies. DNA Research 18: 93-105.
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Geel B. van, Fisher D.C., Rountrey A.N., Arkel J. van, Duivenvoorden J.F., Nieman A.M., Reenen G.B.A. van, Tikhonov A.N., Buigues B., Gravendeel B. 2011. Palaeo-environmental and dietary analysis of intestinal contents of a mammoth calf (Yamal Peninsula, northwest Siberia). Quaternary Science Reviews 30: 3935-3946.
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Journals SCI, peer-reviewed

Sedayu A., Eurlings M.C.M., Gravendeel B., Hetterscheid W.L.A. 2010. Morphological character evolution of Amorphophallus (Araceae) based on a combined phylogenetic analysis of trnL, rbcL, and LEAFY second intron sequences. Botanical Studies 51: 473-490.

Eurlings M.C.M., Heuveling van Beek H., Gravendeel B. 2010. Polymorphic microsatellites for forensic identification of agarwood (Aquilaria crassna). Forensic Science International 197: 30-34.
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Journals SCI, peer-reviewed

Geel B. van, Aptroot A., Baittinger C., Birks H.H., Bull I.D., Cross H.B., Evershed R.P., Gravendeel B., Kompanje E.J.O., Kuperus P., Mol D., Nierop K.G.J., Pals J.P., Tikhonov A.N., Reenen G. van, Tienderen P.H. van 2008. The ecological implications of a Yakutian mammoth's last meal. Quaternary Research 69: 361-376.
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Hof K. van den, Berg R.G. van den, Gravendeel B. 2008. Chalcone synthase gene lineage diversification confirms allopolyploid evolutionary relationships of European rostrate violets. Molecular Biology and Evolution 25: 2099-2108.
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Kocyan A., Vogel E.F. de, Conti Elena, Gravendeel B. 2008. Molecular phylogeny of Aerides (Orchidaceae) based on one nuclear and two plastid markers: A step forward in understanding the evolution of the Aeridinae. Molecular Phylogenetics and Evolution 48: 422-443.
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Journals non-SCI, peer-reviewed

Gravendeel B. 2008. Fossiele bestuiver dateert orchideeënfamilie. Entomologische Berichten 68: 2-4.


Journals SCI, peer-reviewed

Fischer G.A., Gravendeel B., Sieder A., Andriantiana J., Heiselmayer P., Cribb P.J., Camargo Smidt E. de, Samuel R., Kiehn M. 2007. Evolution of resupination in malagasy species of Bulbophyllum (Orchidaceae). Molecular Phylogenetics and Evolution 45: 358-376.
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Ramirez S.R., Gravendeel B., Singer R.B., Marshall C.R., Pierce N.E. 2007. Dating the origin of the Orchidaceae from a fossil orchid with its pollinator. Nature 448: 1042-1045.
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Journals non-SCI, peer-reviewed

Gravendeel B. 2007. Fossiele bestuiver dateert orchideeënfamilie. Eurorchis 19: 16-19.

Gravendeel B., Baas P., Smets E. 2007. Herbaria in Nederland: van blijvend belang voor het behoud van biodiversiteit. Gewina: Tijdschrift voor de Geschiedenis der Geneeskunde, Natuurwetenschappen, Wiskunde en Techniek 30: 153-161.


Journals SCI, peer-reviewed

Gravendeel B., Schuiteman A., Vogel E.F. de 2005. Molecular dating and vicariance analysis of Coelogyninae (Orchidaceae). Regnum Vegetabile 143: 131-148.

Hsu S.C., Gravendeel B., Vogel E.F. de 2005. Taxonomic revision of geesinkorchis (Coelogyninae; Epidendroideae; Orchidaceae). Blumea: Journal of Plant Taxonomy and Plant Geography 50: 505-517.

Eurlings M.C.M., Gravendeel B. 2005. TrnL-trnF sequence data imply paraphyly of Aquilaria and Gyrinops (Thymelaeaceae) and provide new perspectives for agarwood identification. Plant Systematics and Evolution 254: 1-12.
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Edited books

Bakker F., Chatrou L.W., Gravendeel B. (eds.). 2005. Plant Species-level Systematics: New perspectives on pattern & process. Regnum Vegetabile 143 348 p. ISBN 9783906166391.

Chapters in books

Gravendeel B., Vogel E.F. de, Schuiteman A. 2005. Subtribe Coelogyninae. Genera Orchidacearum 4 4: ISBN 0198507127.

Gravendeel B., Mabberley D., Meijden R. van der 2005. Andere indeling plantenfamilies: nieuwe inzichten uit DNA-onderzoek. Heukels' Flora van Nederland ISBN 9789075000894.


Journals SCI, peer-reviewed

Pedersen H.A., Gravendeel B., Mudiana D. 2004. Three new species of dendrochilum (Orchidaceae) and their phylogenetic positions according to plastid and nuclear ribosomal its sequences. Blumea: Journal of Plant Taxonomy and Plant Geography 49: 351-360.

Gravendeel B., Eurlings M.C.M., Berg C. van den, Cribb P.J. 2004. Phylogeny of Pleione (Orchidaceae) and parentage analysis of its wild hybrids based on plastid and nuclear ribosomal ITS sequences and morphological data. Systematic Botany 29: 50-63.

Pelser P.B., Hof K. van den, Gravendeel B., Meijden R. van der 2004. The systematic value of morphological characters in Senecio sect. Jacobaea (Asteraceae) as compared to DNA sequences. Systematic Botany 29: 790-805.

Bakker H.C. den, Gravendeel B., Kuyper T.W. 2004. An ITS phylogeny of Leccinum and an analysis of the evolution of minisatellite-like sequences within ITS1. Mycologia 96: 102-118.

Mols J.B., Gravendeel B., Chatrou L.W., Pirie M.D., Bygrave P.C., Chase M.W., Kessler P.J.A. 2004. Identifying clades in Asian Annonaceae: Monophyletic genera in the polyphyletic Miliuseae. American Journal of Botany 91: 590-600.

Gravendeel B., Smithson A., Slik J.W.F., Schuiteman A. 2004. Epiphytism and pollinator specialization: drivers for orchid diversity?. Philosophical Transactions of the Royal Society B: Biological Sciences 359: 1523-1535.
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Grob G.B.J., Gravendeel B., Eurlings M.C.M. 2004. Potential phylogenetic utility of the nuclear FLORICAULA/LEAFY second intron: comparison with three chloroplast DNA regions in Amorphophallus (Araceae). Molecular Phylogenetics and Evolution 30: 13-23.
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Journals SCI, peer-reviewed

Pelser P.B., Gravendeel B., Meijden R. van der 2003. Phylogeny reconstruction in the gap between too little and too much divergence: the closest relatives of Senecio jacobaea (Asteraceae) according to DNA sequences and AFLPs. Molecular Phylogenetics and Evolution 29: 613-628.
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Journals non-SCI, peer-reviewed

Wilde W.J.J.O. de, Duyfjes B.E.E., Gravendeel B., Ham R.W.J.M. van der, Ho B.C., Rugayah, Vo G.T.P. 2003. Callitriche nana (Callitrichaceae), a new species from Java, Indonesia. Floribunda 2: 57-66.


Journals SCI, peer-reviewed

Grob G.B.J., Gravendeel B., Eurlings M.C.M., Hetterscheid W.L.A. 2002. Phylogeny of the tribe Thomsonieae (Araceae) based on chloroplast matK and trnL intron sequences. Systematic Botany 27: 453-467.

Gravendeel B., Vogel E.F. de 2002. Revision of coelogyne section Moniliformes (Orchidaceae) based on morphology, plastid and nrDNA ITS sequences. Blumea: Journal of Plant Taxonomy and Plant Geography 47: 409-462.

Pelser P.B., Gravendeel B., Meijden R. van der 2002. Tackling speciose genera: species composition and phylogenetic position of Senecio sect. Jacobaea (Asteraceae) based on plastid and nrDNA sequences. American Journal of Botany 89: 929-939.


Journals SCI, peer-reviewed

Gravendeel B., Chase M.W., Vogel E.F. de, Roos M.C., Mes T.H.M., Bachmann K. 2001. Molecular phylogeny of Coelogyne (Epidendroideae; Orchidaceae) based on plastid RFLPS, matK, and nuclear ribosomal its sequences: Evidence for polyphyly. American Journal of Botany 88: 1915-1927.


Journals SCI, peer-reviewed

Pelser P.B., Gravendeel B., Vogel E.F. de 2000. Revision of Coelogyne section Fuliginosae (Orchidaceae). Blumea: Journal of Plant Taxonomy and Plant Geography 45: 253-+.

Sierra S.E.C., Gravendeel B., Vogel E.F. de 2000. Revision of Coelogyne section Verrucosae (Orchidaceae): A new sectional delimitation based on morphological and molecular evidence. Blumea: Journal of Plant Taxonomy and Plant Geography 45: 275-318.

Authored books

Gravendeel B. 2000. Reorganising the orchid genus Coelogyne - a phylogenetic classification based on molecules and morphology. Doctoral thesis 208 p. Nationaal Herbarium Nederland, Leiden, The Netherlands. ISBN 9789071236488.


Journals SCI, peer-reviewed

Gravendeel B., Vogel E.F. de 1999. Revision of Coelogyne section Speciosae (Orchidaceae). Blumea: Journal of Plant Taxonomy and Plant Geography 44: 253-320.


Journals SCI, peer-reviewed

Gravendeel B. 1998. Phylogeny of Coelogyne Lindl. (Orchidaceae) based on morphology and cpDNA RFLP data. Acta Botanica Neerlandica 47: 23-264.