Further information


Papers in peer-reviewed journals, book chapters


Van Rooy, J., N. E. Bell, T. A. Hedderson, J. Flores & J. Hyvönen. Notoligotrichum synonymised with Atrichopsis (Polytrichaceae, Bryophyta), with two new species from South Africa. Taxon (submitted).

Bell, N. E., I. U. Kariyawasam, J. Flores & J. Hyvönen. The diversity of the Polytrichopsida – a review. Bryophyte Diversity and Evolution (submitted).

Blockeel, T. L.,  N. E. Bell, M. O. Hill, N. G. Hodgetts, D.G. Long, S. L. Pilkington & G. P. Rothero. 2020. A new checklist of the bryophytes of Britain and Ireland, 2020. Journal of Bryology (in press).

Kariyawasam, I. U., M. J. Price, N. E. Bell, D. G. Long, R. R. Mill & J. Hyvönen. 2020. Unearthing a lectotype for Polytrichum commune Hedw. (Bryophyta, Polytrichaceae). Taxon (in press).

Flagmeier, M., J. Squirrell, M. Woodhead, D. G. Long, N. E. Bell, J. Russell, W. Powell & P. M. Hollingsworth. 2020. Globally rare oceanic-montane liverworts with disjunct distributions: evidence for long-distance dispersal. Biodiversity and Conservation 29: 3245-3264.

Hodgetts, N. G., L. Söderström, T. L. Blockeel, S. Caspari, M. S. Ignatov, N.A. Konstantinova, N. Lockhart, B. Papp, C. Schröck, M. Sim-Sim, D. Bell, N. E. Bell, H. H. Blom, M. A. Bruggeman-Nannenga, M. Brugués, J. Enroth, K. I.Flatberg, R. Garilleti, L. Hedenäs, D. T. Holyoak, V. Hugonnot, I. Kariyawasam, H. Köckinger, J. Kučera, F. Lara & R. D. Porley. 2020. An annotated checklist of bryophytes of Europe, Macaronesia and Cyprus. Journal of Bryology 42: 1-116.


Hodgetts, N. et al., (incl. N. E. Bell). 2019. A miniature world in decline: European Red List of Mosses, Liverworts and Hornworts. Brussels, Belgium: IUCN.

Liu Y., M. G. Johnson, C. J. Cox, R. Medina, N. Devos, A. Vanderpoorten, L. Hedenäs, N. E. Bell, J. R. Shevock, B. Aguero, D. Quandt, N. J. Wickett, A. J. Shaw & B. Goffinet. 2019. Resolution of the ordinal phylogeny of mosses using targeted exons from organellar and nuclear genomes. Nature Communications, 10: 1485, https://doi.org/10.1038/s41467-019-09454-w

Callaghan, D. A., N. E. Bell & L. L. Forrest. 2019. Taxonomic notes on Weissia subgenus Astomum, including Weissia wilsonii D.A.Callaghan, a new species from Europe. Journal of Bryology, 41: 135-148, https://doi.org/10.1080/03736687.2018.1551590

Bell, N. E. & M. Ignatov. 2019. Placing the regionally threatened moss Orthodontium gracile in the big picture – Phylogeny, genome incongruence and anthropogenic dispersal in the order Orthodontiales. Molecular Phylogenetics and Evolution, 134: 186-199, https://doi.org/10.1016/j.ympev.2018.12.024


Huttunen, S., N. E. Bell & L. Hedenäs. 2018. The evolutionary diversity of mosses – taxonomic heterogeneity and its ecological drivers. Critical Reviews in Plant Sciences, https://doi.org/10.1080/07352689.2018.1482434

Bell, N. E. & M. Ignatov. 2018. Orthodontiales. Pp. 518-523 in: Ignatov M.S. (ed.), Moss Flora of Russia. Volume 4. Bartramiales – Aulacomniales. KMK, Moscow, Russia.

Kariyawasam, U., D. G. Long & N. E. Bell. 2018. A Taxonomic Revision of Oligotrichum Lam. & DC. (Polytrichaceae) in the Sino-Himalaya. Journal of Bryology, https://doi.org/10.1080/03736687.2018.1478651


Coudert, Y, N. E. Bell, C. Edelin & C. J. Harrison CJ. 2017. Multiple innovations underpinned branching form diversification in mosses. New Phytologist, 10.1111/nph.14553.

Bell, N. E., J. Hyvönen, K.-Y. Yao & W.-Z. Ma. 2017. Description and phylogenetic investigation of Pogonatum shevockii N.E.Bell & Hyvönen (Polytrichaceae), a new East Asian species with a unique leaf morphology. Journal of Bryology 39: 235-246.

Ellis, L. T., O. M. Afonina, R. L. Andriamiarisoa, H. Bednarek-Ochyra, B. Cykowska-Marzencka, M. Stryjak-Bogacka, N. E. Bell, M. Boiko, D. A. Callaghan, P. Campisi, M. G. Dia, M. L. Marino, F. Provenzano, J. Eckstein, J. Enroth, P. Erzberger, T. Ezer, M. L. Gargano, E. Ginzburg, P. Górski, S. R. Gradstein, C. Reeb, C. Hannoire, M. Infante, I. Jukonienė, E. V. Kushnevskaya, M. Lebouvier, J. Nagy, A. Opmanis, V. Plášek, Z. Skoupá, M. S. Sabovljević, A. D. Sabovljević, J. R. Shevock, D. K. Singh, S. Majumdar, M. Skudnik, A. Uselienė, G. Venturella, M. Węgrzyn, P. Wietrzyk, Y.-J. Yoon, J. H. Kim & E. Yücel. 2017. New national and regional bryophyte records, 53. Journal of Bryology 39: 368-387.

Bell, N. E.. 2017. Editorial. Edinburgh Journal of Botany 74: 109-110.


Daniels, E. D., K. C. Kariyappa, J.Hyvönen & N. E. Bell. 2016. The first Indian record of Pogonatum marginatum Mitt. (Polytrichaceae) from the Western Ghats. Bryophyte Diversity and Evolution 38: 41-46.


Bell, N. E., I. U. Kariyawasam, T. A. J. Hedderson & J. Hyvönen. 2015. Delongia gen. nov., a new genus of Polytrichaceae (Bryophyta) with two disjunct species in East Africa and the Himalaya. Taxon 64: 893–910.I.

Ivanova, E. I., N. E. Bell, O. I. Kuznetsova, E. A. Ignatova & M. S. Ignatov. 2015. The genus Polytrichum sect. Aporotheca (Polytrichaceae) in Russia. Arctoa 24: 67-78.


Bell, N. E., J. L. Boore, B. D. Mishler & J. Hyvönen. 2014. Organellar genomes of the four-toothed moss, Tetraphis pellucida. BMC Genomics 15:383.

Ivanova, E. I., N. E. Bell, O. I. Kuznetsova, H. Lee, M. Park & M. S. Ignatov. 2014. The genus Polytrichastrum (Polytrichaceae) in Russia. Arctoa 23: 164-184.

Poczai, P., N. E. Bell & J. Hyvönen. 2014. Imre Festetics and the Sheep Breeders’ Society of Moravia: Mendel’s Forgotten “Research Network”. PloS Biology 12: e1001772.


Poczai, P., I. Varga, M. Laos, A. Cseh, N. E. Bell, J. Valkonen & J. Hyvönen. 2013. Advances in plant gene-targeted and functional markers: a review. Plant Methods 9:6.


Bell, N. E., A. E. Newton & J. Hyvönen. 2012. Epiphytism and generic endemism in the Hypnodendrales: Cyrtopodendron, Franciella and macro-morphological plasticity. Taxon 61: 498-514.

Bell, N. E., A. E. Newton & J. Hyvönen. 2012. New observations on Pterobryella (Müll.Hal.) A.Jaeger and Cyrtopodendron M.Fleisch. In New Caledonia. Journal of Bryology 34: 306-308.

Huttunen, S., N. E. Bell, M. Stech & D. Quandt. 2012. Bryophyte Tree Of Life – the current state of phylogenetic reconstruction in the mosses (Bryophyta). Journal of Bryology 34: 157-159.

Bell, N. E. & J. Hyvönen. 2012. Gametophytic simplicity in Laurasian and Gondwanan Polytrichopsida – the phylogeny and taxonomy of the Oligotrichum morphology. Journal of Bryology 34: 160-172.

Huttunen, S., N. E. Bell, V. K. Bobrova, V. Buchbender, W. R. Buck, C. J. Cox, B. Goffinet, L. Hedenäs, B. Ho, M. S. Ignatov, O. Kuznetsova, I. A. Milyutina, A. E. Newton, S. Olsson, L. Pokorny, J. A. Shaw, M. Stech, A. Troitsky, A. Vanderpoorten & D. Quandt. 2012. Disentangling knots of rapid evolution: origin and diversification of the moss order Hypnales. Journal of Bryology 34: 187-211.


Poczai, P., I. Varga, N. E. Bell & J. Hyvönen. 2011. Genetic diversity assessment of bittersweet (Solanum dulcamara, Solanaceae) germplasm using conserved DNA-derived polymorphism and intron-targeting markers. Annals of Applied Biology 159: 141-153.


Bell, N. E. & J. Hyvönen. 2010. A phylogenetic circumscription of Polytrichastrum (Polytrichaceae): reassessment of sporophyte morphology supports molecular phylogeny. American Journal of Botany 97: 566-578.

Bell, N. E. & J. Hyvönen. 2010. Phylogeny of the moss class Polytrichopsida (Bryophyta): generic-level structure and incongruent gene trees. Molecular Phylogenetics and Evolution 55: 381-398.

Bell, N. E. & J. Hyvönen. 2010. Molecular evidence for the phylogenetic position of the New Caledonian endemic genus Parisia. Journal of Bryology 32: 303-305.

Bell, N. E. & J. Hyvönen. 2010. A dendroid form of Pogonatum cirratum (Polytrichopsida, Bryophyta) from Mt. Kinabalu (Sabah, Malaysia). Acta Bryolichenologica Asiatica3: 39-45.


Bell, N. E. & J. Hyvönen. 2008. Rooting the Polytrichopsida: the phylogenetic position of Atrichopsis and the independent origin of the polytrichopsid peristome. Pp. 227-239 in: H. Mohamed and B. B. Baki (eds.) Bryology in the New Millennium. University of Malaya, Kuala Lumpur.


Bell, N. E., N. Pedersen & A. E. Newton. 2007. Ombronesus stuvensis, a new genus and species of the Ptychomniaceae (Bryophyta) from south west Chile. Taxon 56: 887-896.

Bell, N. E., D. Quandt, T. J. O’Brien & A. E. Newton. 2007. Taxonomy and phylogeny in the earliest diverging pleurocarps: square holes and bifurcating pegs. The Bryologist 110: 533-560.

Bell, N. E. & P. V. York. 2007. Vetiplanaxis pyrrhobryoides, a new fossil moss genus and species from Middle Cretaceous Burmese amber. The Bryologist 110: 514-520.

Quandt, D., N. E. Bell & M. Stech. 2007. Unravelling the knot: the Pulchrinodaceae fam. nov. (Bryales). Nova Hedwigia Beiheft 131: 21-39

Bell, N. E. & A. E. Newton. 2007. Pleurocarpy in the rhizogoniaceous grade. Pp. 41-64 in: A. E. Newton & R. Tangney (eds.) Pleurocarpous Mosses: Systematics and Evolution. CRC Press, Boca Raton, FL.

Newton, A. E., N. Wikstrom & N. E. Bell. 2007. Dating the diversification of the pleurocarpous mosses. Pp. 337-366 in: A. E. Newton & R. Tangney (eds.) Pleurocarpous Mosses: Systematics and Evolution. CRC Press, Boca Raton, FL.


Hill, M. O., N. E. Bell, M. A. Bruggeman-Nannenga, M. Brugués, M. J. Cano, J. Enroth, K. I. Flatberg, J.-P. Frahm, M. T. Gallego, R. Garilleti, J. Guerra, L. Hedenäs, D. T. Holyoak, J. Hyvönen, M. S. Ignatov, F. Lara, V. Mazimpaka, J. Muñoz & L. Söderström. 2006. An annotated checklist of the mosses of Europe and Macaronesia. Journal of Bryology 28:198-267.


Bell, N. E. & A. E. Newton. 2005. The paraphyly of Hypnodendron and the phylogeny of related non-hypnanaean pleurocarpous mosses inferred from chloroplast and mitochondrial sequence data. Systematic Botany 30: 34-51


Bell, N. E. & A. E. Newton. 2004. Systematic studies of non-hypnanaean pleurocarps: establishing a phylogenetic framework for investigating the origins of pleurocarpy. In B. Goffinet, V. C. Hollowell & R. E. Magill (eds.), Molecular Systematics of Bryophytes. Monographs in Systematic Botany from the Missouri Botanical Garden 98: 290-319.



Hodgetts, N. (et al.) 2019. A miniature world in decline European Red List of Mosses, Liverworts and Hornworts. IUCN, Brussels, Belgium.

Callaghan, D., & N. E. Bell. 2017. Identity and taxonomy ofWeissia multicapsularis. Commissioned report.

Bell, N. E. & J. Cuvertino. 2003. Distribution and Diversity of Bryophytes in the Katalalixar National Reserve, Region XI, Chile. For the Biodiversity Aysén Project, a joint initiative between Raleigh International and Corporación Nacional Forestal (CONAF), Chile.

Bell, N. E. 2002. Potential Impact on the Bryophyte Flora of Maplehurst Wood SSSI of Current and Future Activities at Woodlands Farm, Hastings. Commissioned report.


Articles in non-peer reviewed periodicals

Bell, N E. 2018. The threatened Moss Orthodontium gracile. Much more than meets the eye (or even the microscope). The Botanics Magazine 71: 6-7.

Bell, N. E. 2017. Orthodontium in the UK: conservation value in a global context. Field Bryology 118: 24-25.

Bell, N. E. & J. Hyönen. 2014. Lehtisammalten makromorfologia – konvergenssia yksinkertaisissa alkiollisissa kasveissa. (Translastion: “Macromorphology in mosses: architectural convergence and phylogeny in simple land plants”). Luonnon Tutkija 4: 135-140.

Piippo, S., N. E. Bell, R. Fagersten, X. He, N. Kalinauskaite & T. Koponen. 2013. Suomen Sammalseuran syysretki Kalkkikalliolle ja Roosinmäelle Helsingin ja Vantaan rajalla 2012. Bryobrotherella 16: 23-36.

Piippo, S., N. E. Bell, R. Fagersten, X. He, N. Kalinauskaite & T. Koponen. 2013. Suomen Sammalseuran kevätretki Sipoonkorven Bakunkärriin 2012. Bryobrotherella 16: 1-7.

Enroth, J., N. E. Bell, M. von Cräutlein, X. He, S. Huttunen, J. Hyvönen, H. Korpelainen, S. Laaka-Lindberg, T. Laukka, M. Lehtonen, S. Olsson, S. Piippo, R. Ryömä, J. Valkonen, H. Vasander & R. Virtanen. 2011. Sammalten valtakunta. Luonnon Tutkija 115: 120-145.

Bell, N. E. 2010. The giant mosses of New Caledonia: a collecting trip, October-November 2008. Bryobrotherella 13: 32-40.

Selected current projects

Phylogeny of the genus Orthodontium and the status of the threatened UK priority conservation species Orthodotium gracile.

Phylogenetic structure, biogeography and taxonomy of Scotland's oceanic liverwort heath community ("mixed northern hepatic mat").

Evolutionary development of branching forms in pleurocarpous mosses (in collaboration with Yoan Coudert at the Ecole Normale Supérieure in Lyon).

Taxonomy, phylogeny and species discovery in the Polytrichaceae (multiple collaborators).

Research group and teaching


Isuru Kariyawasam (current) - Rebooting Systematics for Morphologically Cryptic Taxa in the Genomic Age – A Molecular Taxonomic Revision of Polytrichum Hedw. Sect. Polytrichum (Polytrichaceae). University of Edinburgh.


Diego Sánchez-Ganfornina (current) - Dating the diversification of Australasian dendroid mosses in the order Hypnodendrales (and addressing unresolved taxonomic questions). University of Edinburgh.

MSc (2nd supervisor)

Isuru Kariyawasam (2013) - Taxonomic Revision of Oligotrichum DC. in Lam. & DC. (Polytrichaceae) in the Sino-Himalaya.

Current teaching

MSc Plant Taxonomy and Biodiversity - Bryophytes.

Membership and editorial

Editorial Roles

Editor-in-Chief, Edinburgh Journal of Botany

Scientific Editor, Journal of Bryology

Societies and Committees

Member of Council, British Bryological Society

Chair of Publications Committee, British Bryological Society