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Peter Hovenkamp

P.H. Hovenkamp, Senior researcher - Biodiversity Discovery


Email: peter.hovenkamp@naturalis.nl
Phone: +31 (0)71 7519213
Room number: 2.03, Nieuwenhuizenweg

Peter Hovenkamp (65) passed away in Malaysia

On Juli 13 we received very sad news about the death of our beloved colleague and valued researcher Peter Hovenkamp. Peter was part of a tour group visiting a cave in Gunung Mulu National Park when a flash flood hit unexpectedly. Unfortunately, he did not survive the accident. 
Peter was a long-time researcher. He started working at Leiden University in 1983 and has been a Naturalis employee since 2013. He was very dedicated to his work and passionate about his research, which mostly involved ferns. 
Our deepest sympathy goes out to Peters family and loved ones.


"Time spent on careful observation is never wasted."

I am first and foremost a systematist, and I want to find out what is really there.

I am convinced that we can only find meaningful answers if we ask meaningful questions. I think therefore that we need to know what species are there, and what their variability and distribution is, before we can investigate their evolution and interactions. 

The same attitude has also led me to consider some questions in the fields of phylogenetic reconstruction, biogeography and biodiversity informatics. It is fascinating to find time and again that answers to difficult questions suddenly become much easier when the questions are formulated clearly and carefully.

I graduated at Leiden University in 1986, on a thesis on the Polypodiaceous genus Pyrrosia. For a few years I then worked in the IT-support industry, and in 1990 I was appointed half-time at the Rijksherbarium, Leiden, where I worked on preparing a treatment of Polypodiaceae for Flora Malesiana. I am currently researcher at Naturalis Biodiversity Center, and Editor-in-Chief of the botanical scientific journal Blumea. 

ORCID: http://orcid.org/0000-0002-4124-2175

Publons: publons.com/a/703310/


Research interest

I have studied the taxonomy of number of fern genera (a.o. Pyrrosia, Selliguea, Nephrolepis, Oleandra), always with the goal to search for orderliness in the patterns displayed by the species. I mainly study herbarium specimens, because only well-documented and permanently preserved specimens provide reliable links to other studies.

PICT0354.JPG (2048×1536)

Oleandra neriifolia - one of the few ferns that have a shrubby habit.

In the course of these studies, I also was excited to find that it is still possible to discover entirely new structures or phenomena, such as the diversity of rhizome morphology in the Polypodiaceae, or the role of spore shape and structure in spore dispersal in Angiopteris (Marattiaceae).

A sporangium of Angiopteris (a eusporangiate fern) opening. Note the erratic movement of the spores. The movement is shown in real time. A single sporangium is approx. 1 mm wide.

In a more theoretical mood, I have tried to elucidate the meaning of permutation branch supports in phylogenetics, and the role of phylogeny in the reconstruction of the biogeographic history of taxa.


polypodiaceae, diplazium, nephrolepis, biogeography, phylogenetics, morphology and anatomy, taxonomy, oleandra

Current research topics

Affinities and origin of some ferns on Mount Kinabalu

During the Kinabalu-Crocker Range Expedition of 2012 I was able to collect sufficient representatives of some fern groups on Mount Kinabalu to start an investigation into the origin and affinities of the unique fern flora on this mountain. In a number of cases, it has become clear that this will only be the start of a larger project, e.g., a more wide-ranging revision of the genus Diplazium. For a start, see Diplazium in Malesia.

Elucidation of some species complexes in Nephrolepis

Taxonomic and phylogenetic research in Nephrolepis have identified a number of further questions regarding the origin of some species or species complexes in this genus. The structure of the complex of species around N. cordifolia, and the contributions of this complex to the formation of the cultivated N. “bostonii” are currently the topic of a phylogenetic study.

Taxonomy of Orthiopteris (Saccolomatacea)

Orthiopteris is a widespread genus of terrestrial ferns, of which the affinities are not quite clear. In most phylogenetic analyses, it occupies an isolated position at the basis of the higher leptosporangiate ferns. It occurs both in the Old and the New world tropics, but the exact relation between the species in these two regions are unclear. In an effort to clarify at least the Old-world species, we have revised these and their delimitations. Further work to relate these to the New-world species must now be undertaken.

Phytochemical characterization of trophopods

Trophopods are underground storage organs in ferns, of which the distribution and function is still poorly understood. In collaboration with Young Hae Choi and Shi-Kai Yan (IBL, Leiden) and the Hortus botanicus Leiden we have studied the starch contents and its seasonal changes of trophopods in Matteucia struthiopteris.

Towards a checklist of the ferns of Sulawesi

This project originated because I was able to work on a number of large collections made on Sulawesi (Celebes): the collections made by Jermy and Walker in 1969 (thanks to a Synthesys fellowship at the BM in London) and the collections of Hennipman in 1979. I have collected the available information on a Scratchpad-website, and it is now a “backburner” project, where I occasionally spend some time to updating the information. Go to the website Ferns of Sulawesi.

The species from the Dryopteris affinis-complex in The Netherlands

The Dryopteris affinis complex is represented in The Netherlands by an unknown number of taxa and hybrids, at different ploidy levels. In collaboration with Ben Zonneveld, for the flowcytometric analysis of nuclear weights, and a number of Dutch amateur experts we are now trying to find out how many distinct taxa there are, and how they can be distinguished morphologically.

Students that have contributed or are still contributing to his research recently include Boon-Chuan Ho, Nor Hidayah Binti Jahaya (Malaysia), A.J. Boot (Leiden), T. T. Luong (Vietnam) and Nurul Hazlina Binti Haji Zaini (Brunei).

The Taxonomic Mindmapper

Through my involvement in the pro-iBiosphere project I have also become interested in novel ways to connect different types of specimen metadata. In a joint project with Royal Botanic Gardens Kew and Royal Botanic Gardens Edinburgh we are now trying to develop  this approach into a formal description. 


  • Natural Products Laboratory, Institute of Biology, Leiden University.
  • Hortus Botanicus, Leiden University.
  • Biodiversity Informatics, Royal Botanic Gardens, Kew.



From the beginning of my appointment as researcher, I have been involved in teaching theory and methods in phylogeny reconstruction to BSc and MSc students in Leiden, and occasionally to MSc and PhD students in international courses organized in Bogor, Leiden and Wageningen. I also teach Ferns in courses in Cryptogamic Botany and Plant identification. 

Supervision MSc Students

2014 Supervisor, Zaini - Taxonomy, morphology and anatomy of several unusual Diplaziums
2012 Supervisor - Thien Tam Luong - Revision of the fern Genus Orthiopteris in Malesia and adjacent regions
2009 Supervisor - Nor Hidayah Binti Yahaya – Taxonomic and Phylogenetic studies in Nephrolepis cordifolia and related species
2006 Supervisor - Agung Sedayu - Revision and ecology of the fern genus Acrostichum in Malesia
2005 Supervisor - Muhammad Iqbal – Zonation of the fern flora on Gunung Lumut, East Kalimantan
2003 Supervisor - Ho Boon-chuan – Revision of the species group Oleandra sibbaldii (Oleandraceae)
2002 Supervisor - Ho Boon-Chuan – Revision of Calyptrochaete (Hookeriaceae)
1999 Supervisor - A.A.P. Leonardia – Revision of the fern genus Arthropteris
Available student projects

Public outreach

I am a more or less active member of the Nederlandse Varenvereniging (Dutch Fern Society), and I have been Editor of their journal VarenVaria for a couple of years. 



Journals SCI, peer-reviewed

Kartonegoro A., Veldkamp J.F., Hovenkamp P., Welzen P. van 2018. A revision of Dissochaeta (Melastomataceae, Dissochaeteae). PhytoKeys 107: 1-178.
Go to website (URI)

Baas P., Hovenkamp P.H. 2018. In memoriam Jan-Frits Veldkamp (31 March 1941 – 12 November 2017). Blumea: Journal of Plant Taxonomy and Plant Geography: 1-10.
Go to website (DOI)

He L., Schneider H., Hovenkamp P., Marquardt J., Wei R., Wei X., Zhang X., Xiang Q. 2018. A molecular phylogeny of selligueoid ferns (Polypodiaceae): Implications for a natural delimitation despite homoplasy and rapid radiation. Taxon 67: 237-249.
Go to website (DOI)

Thomson S.A., Pyle R.L., Ahyong S.T., [...] Byng J.W., Dekker R., Dijkstra K.-D., Duistermaat H., Geml J., Hovenkamp P., et al. 2018. Taxonomy based on science is necessary for global conservation. PLoS Biology 16: e2005075.
Go to website (DOI)

Hovenkamp P., Hendrikx B., Roskam H., Winter W. de. 2018. Het Dryopteris affinis-complex in Nederland. Gorteria 40: 42-54.


Journals non-SCI, peer-reviewed

Lindsay S., Hovenkamp P.H., Middleton D.J. 2017. New combinations and typifications in Aglaomorpha (Polypodiaceae). Gardens’ Bulletin Singapore 69: 149-155.
Go to website (DOI)


Journals SCI, peer-reviewed

Khine P.K., Lindsay S., Fraser-Jenkins C., Kluge J., Kyaw M., Hovenkamp P. 2016. Selliguea kachinensis (Polypodiaceae), a new fern species of uncertain affinity from Northern Myanmar. PhytoKeys 62: 73-81.
Go to website (URI)

Yahaya N.H., Michael Stech, Zonneveld B.J.M., Hovenkamp P.H. 2016. What is Nephrolepis ‘bostoniensis’?: Unravelling the origin of Nephrolepis hybrids and cultivars with molecular data. Scientia Horticulturae 204: 153-160.

Hovenkamp P., Yan S.-K., Young H.-C. 2016. Seasonal changes in starch content in trophopods of Matteuccia struthiopteris (L) Todaro . American Fern Journal 16: 153-160.
Go to website (URI)

PPG I 2016. A community-derived classification for extant lycophytes and ferns. PPG I. Journal of Systematics and Evolution 54: 563-603.
Go to website (URI)


Journals SCI, peer-reviewed

Rothfels C.J., Johnson A.K., Hovenkamp P.H., Swofford D.L., Roskam H.C., Fraser-Jenkins C.R., Windham M.D., Pryer K.M. 2015. Natural hybridization between genera that diverged from each other approximately 60 million years ago. The American Naturalist 185: 433-442.
Go to website (URI)

Luong T.T., Hovenkamp P.H., Sosef M.S.M. 2015. Revision of the fern genus Orthiopteris (Saccolomataceae) in Malesia and adjacent regions. PhytoKeys 53: 39-71.
Go to website (URI)

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.
Go to website (DOI)

Journals non-SCI, peer-reviewed

Hovenkamp P.H. 2015. Two new Diplazium (Woodsiaceae) species from East Malesia. The Fern Gazette 20: 49-54.


Journals SCI, peer-reviewed

Hovenkamp P. 2014. A too modest proposal. Cladistics 30: 232-233.
Go to website (DOI)

Journals non-SCI, peer-reviewed

Vos R.A., Biserkov J.V., Balech B., Beard N., Blissett M., Brenninkmeijer C., Dooren T. van, Eades D., Gosline G., Groom Q.J., Hamann T.D., Hettling H., Hoehndorf R., Holleman A., Hovenkamp P., Kelbert P., King D., Kirkup D., Lammers Y., DeMeulemeester T., Mietchen D., Miller J.A., Mounce R., Nicolson N., Page R., Pawlik A., Pereira S., Penev L., Richards K., Sautter G., Shorthouse D.P., Tähtinen M., Weiland C., Williams A.R., Sierra S. 2014. Enriched biodiversity data as a resource and service. Biodiversity Data Journal 2: 1-15.
Go to website (URI)

Edited books

Hovenkamp P. (eds.). 2014. Planten uit de oertijd. 20 p. Hortus botanicus Leiden, Leiden.


Journals SCI, peer-reviewed

Liu H., Jiang R., Guo J., Hovenkamp P., Perrie L.R., Shepherd L., Hennequin S., Schneider H. 2013. Towards a phylogenetic classification of the climbing fern genus Arthropteris. Taxon 62: 668-700.

Chapters in books

Hovenkamp P. 2013. Flora of Peninsular Malaysia: Oleandraceae. 2: 67-71: Forest Research Institute Malaysia, Kepong, Malaysia.

Shugang L, Hovenkamp P, Gilbert M 2013. Flora of China: Selliguea. 2-3: 773-785: Science Press, Beijing and MBG Press, St. Louis, Beijing and St. Louis. ISBN 9781935641117.

Shugang L., Hovenkamp P. 2013. Flora of China: Goniophlebium, Metapolypodium, Polypodiastrum, Polypodiodes. 2-3: 797-804: Science Press, Beijing and MBG Press, St. Louis, Beijing and St. Louis. ISBN 9781935641117.

Shugang L., Hovenkamp, P. 2013. Flora of China: Pyrrosia. Flora of China 2-3: 786-796: Science Press and MBG Press, Beijing and St. Louis. ISBN 9781935641117.

Shugang L., Hovenkamp P. 2013. Flora of China: Arthromeris. 2-3: 768-772: Science Press, Beijing and MBG Press, St. Louis, Beijing and St. Louis. ISBN 9781935641117.

Xinping Q., Xianchun Z., Youxing L., Gilbert M., Hovenkamp P. 2013. Flora of China: Lepisorus. 2-3: 808-824: Science Press, Beijing and MBG Press, St. Louis, Beijing and St. Louis. ISBN 9781935641117.

Youxing L., Hovenkamp P., Gilbert M. 2013. Flora of China: Lemmaphyllum. Flora of China 2-3 Flora of China 2-3 824-826: Science Press, Beijing and MBG Press, St. Louis, Beijing and St. Louis. ISBN 9781935641117.

Shugang L., Hovenkamp P. 2013. Flora of China: Christopteris. 2-3: 772-773: Science Press, Beijing and MBG Press, St. Louis, Beijing and St. Louis. ISBN 9781935641117.

Hovenkamp P. 2013. Flora of Peninsular Malaysia: Polypodiaceae. 2: 97-198: Forest Research Institute Malaysia, Kepong, Malaysia.

Hovenkamp P. 2013. Flora of Peninsular Malaysia: Nephrolepidaceae. 2: 51-66: Forest Research Institute Malaysia, Kepong, Malaysia.

Zhang X.-C., Hovenkamp P.H. 2013. Oleandraceae (Flora of China). Oleandraceae 747-748:

Zhang X.-C., Hovenkamp P. 2013. Polypodiaceae (Flora of China). Polypodiaceae 758-850:

Xing F.-W., Wang F.-G., Hovenkamp P.H. 2013. Nephrolepidaceae (Flora of China). Nephrolepidaceae 727-729:

Wang F.-G., Xing F.-W., Hovenkamp P.H. 2013. Arthropteris (Flora of China). Arthropteris 730-731:


Journals SCI, peer-reviewed

Hovenkamp P.H., Ho B.C. 2012. A revision of the fern genus Oleandra (Oleandraceae) in Asia. PhytoKeys 11: 1-37.
Go to website (DOI)

Rakotondrainibe F., Hovenkamp P.H. 2012. A New Species of Pyrrosia (Polypodiaceae) from Madagascar. Novon: A Journal for Botanical Nomenclature 22: 75-77.
Go to website (DOI)

Hovenkamp P. 2012. Syncretism and corroboration. Cladistics 28: 115-116.
Go to website (DOI)

Chapters in books

Hovenkamp P.H., Miyamoto F. 2012. Nephrolepidaceae (Flora Malesiana).

Hovenkamp P.H., Leonardia A.A.P. 2012. Tectaria-group: Arthropteris (Flora Malesiana). 2 4: 145-149:

Hovenkamp P.H., Ho B.-C. 2012. Oleandraceae (Flora Malesiana). 2 4: 123-136:


Journals SCI, peer-reviewed

Hovenkamp P. 2011. Stability, replication, pseudoreplication, support and consensus-a reply to Brower. Cladistics 27: 4-5.
Go to website (DOI)

Journals non-SCI, peer-reviewed

Roos M.C., Berendshohn W.G., Dessein S., Hamann T., Hoffmann N., Hovenkamp P., Janssen T., Kirkup D., Kok R. de, Sierra S.E.C., Smets E., Webb C., Welzen P.C. van 2011. e-Flora Malesiana: state of the art and perspectives. Gardens’ Bulletin Singapore 63: 189-195.

Hovenkamp P.H., Fraser-Jenkins C.R., Schneider H., Zhang X-C. 2011. Proposal to conserve the name Lepisorus against Belvisia, Lemmaphyllum, Paragramma, Drymotaenium & Neocheiropteris (Pteridophyta, Polypodiaceae). Taxon 60: 591-592.


Journals SCI, peer-reviewed

Hennequin S., Hovenkamp P. Christenhusz M.J.M., Schneider H. 2010. Phylogenetics and biogeography of Nephrolepis- a tale of old settlers and young tramps. Botanical Journal of the Linnean Society 164: 113-127.
Go to website (DOI)

Hovenkamp P., Raes N. 2010. Indicators of endemism. Cladistics 26: 213-213.


Journals SCI, peer-reviewed

Roos M.C., Hovenkamp P. 2009. Flora Malesiana in the coming decade. Blumea: Journal of Plant Taxonomy and Plant Geography 54: 3-5.
Go to website (DOI)

Hovenkamp P. 2009. Support and stability. Cladistics 25: 107-108.
Go to website (DOI)

Hovenkamp P.H, Ham R.W.J.M. van der, Uffelen G.A. van, Hecke M. van, Dijksman J.A., Star W. 2009. Spore movement driven by the spore wall in an eusporangiate fern. Grana 48: 122-127.
Go to website (DOI)

Edited books

Hovenkamp, P. (eds.). 2009. Proceedings of the 7th Flora Malesiana Symposium. Blumea 54 1-3: 299 p.


Journals SCI, peer-reviewed

Schneider H. Kreier H.P., Hovenkamp P., Janssen T. 2008. Phylogenetic relationships of the fern genus Christiopteris shed new light onto the classification and biogeography of drynarioid ferns. Botanical Journal of the Linnean Society 157: 645-656.


Journals SCI, peer-reviewed

Hovenkamp P. 2006. Can taxon-sampling effects be minimized by using branch supports?. Cladistics 22: 264-275.


Journals SCI, peer-reviewed

Hovenkamp P.H., Miyamoto F. 2005. A conspectus of the native and naturalized species of Nephrolepis (Nephrolepidaceae) in the world. Blumea: Journal of Plant Taxonomy and Plant Geography 50: 279-322.


Journals SCI, peer-reviewed

Schneider H., Janssen T., Hovenkamp P., Smith A.R., Cranfill R., Haufler C.H., Ranker T.A. 2004. Phylogenetic relationships of the enigmatic malesian fern Thylacopteris (Polypodiaceae, Polypodiidae). International Journal of Plant Sciences 165: 1077-1087.

Hovenkamp P. 2004. In support of support. Cladistics 20: 85-85.

Hovenkamp P. 2004. Review of: TNT - Tree analysis using new technology. Version 1.0, by P. Goloboff, J. S. Farris and K. Nixon. Available from the authors and from http://www.zmuc.dk/public/phylogeny. Cladistics 20: 378-383.

Hovenkamp P. 2004. No areas. Cladistics 20: 85-85.


Journals SCI, peer-reviewed

Welzen P.C. van, Turner H., Hovenkamp P. 2003. Historical biogeography of Southeast Asia and the West Pacific, or the generality of unrooted area networks as historical biogeographic hypotheses. Journal of Biogeography 30: 181-192.


Journals SCI, peer-reviewed

Turner H., Hovenkamp P., Welzen P.C. van 2001. Biogeography of Southeast Asia and the West Pacific. Journal of Biogeography 28: 217-230.

Hovenkamp P. 2001. A direct method for the analysis of vicariance patterns. Cladistics 17: 260-265.


Journals SCI, peer-reviewed

Hovenkamp P.H. 2000. Paraselliguea, a new genus of Malesian Polypodiaceae, a correction. Blumea: Journal of Plant Taxonomy and Plant Geography 45: 376-376.


Journals SCI, peer-reviewed

Hovenkamp P. 1999. Unambiguous data or unambiguous results?. Cladistics 15: 99-102.


Journals SCI, peer-reviewed

Hovenkamp P. 1998. An account of the Malay-Pacific species of Selliguea (Polypodiaceae). Blumea: Journal of Plant Taxonomy and Plant Geography 43: 1-108.

Hovenkamp P. 1998. Lepisorus in Malesia. Blumea: Journal of Plant Taxonomy and Plant Geography 43: 109-115.

Chapters in books

Hovenkamp et al. 1998. Polypodiaceae. 3 In: Kalkman, C., Nooteboom, H.P. (eds.). Flora Malesiana 2 (3): Polypodiaceae, Davalliaceae, Azollaceae, Cheiropleuriaceae, Equisetaceae, Matoniaceae, Plagiogyriaceae: 1-234 ISBN 90-71236-39-0.


Journals SCI, peer-reviewed

Hovenkamp P.H. 1997. Paraselliguea, a new genus of Malesian Polypodiaceae. Blumea: Journal of Plant Taxonomy and Plant Geography 42: 485-487.

Hovenkamp P. 1997. Vicariance events, not areas, should be used in biogeographical analysis. Cladistics 13: 67-79.


Journals SCI, peer-reviewed

Hovenkamp P.H. 1996. Selliguea sri-ratu, a new species in Polypodiaceae. Blumea: Journal of Plant Taxonomy and Plant Geography 41: 19-20.

Chapters in books

Balgooy M.M.J. van, Hovenkamp P.H., Welzen P.C. van 1996. Phytogeography of the Pacific - floristic and historical distribution patters in plants. SPB Academic Publishing, Amsterdam. ISBN 905103136X.


Journals SCI, peer-reviewed

Hovenkamp P.H., Franken N.A.P. 1993. An account of the fern genus Belvisia-mirbel (Polypodiaceae). Blumea: Journal of Plant Taxonomy and Plant Geography 37: 511-527.


Journals SCI, peer-reviewed

Hovenkamp, P. 1990. The significance of rhizome morphology in the systematics of the polypodiaceous ferns (Sensu-stricto). American Fern Journal 80: 33-43.


Journals SCI, peer-reviewed

Hovenkamp, P.H., Joncheere G.J. de 1988. Additions to the fern flora of Sulawesi. Blumea: Journal of Plant Taxonomy and Plant Geography 33: 395-409.


Edited books

Hovenkamp P., Gittenberger E., Hennipman E., Jong R. de, Roos M.C., Sluys R., Zandee M. (eds.). 1987. Systematics and Evolution: A Matter of Diversity. 341 p. Utrecht University, ISBN 9789090019673.


Authored books

Hovenkamp P. 1986. A monograph of the fern genus Pyrrosia (Polypodiaceae). Leiden Botanical Series 9 280 p. Brill, Leiden. ISBN 9789004080652.


Journals SCI, peer-reviewed

Hovenkamp P. 1984. Some new names and combinations in Pyrrosia mirbel (Polypodiaceae). Blumea: Journal of Plant Taxonomy and Plant Geography 30: 207-208.


Journals SCI, peer-reviewed

Venverloo C.J., Hovenkamp P.H., Weeda A.J., Libbenga K.R. 1980. Cell-division in Nautilocalyx explants - 1: Phragmosome, preprophase band and plane of division. Zeitschrift für Pflanzenphysiologie 100: 161-174.