Dr Mark Carine

Mark Carine and the Azorean endemic Rumex azoricus. Photo: F. Rumsey
  • Researcher, Macronesian Flowering Plants
  • Life Sciences department
  • Plants Division
Natural History Museum
Cromwell Road


Employment history

April 2006-present: Researcher, Band 4, Dept. of Botany, NHM.

April 2001–April 2006: Researcher, Band 5, Department of Botany, The Natural History Museum, London.

April 2000–April 2001: Post-Doctoral Research Fellow, Department of Botany, The University of Reading.

July 1999–April 2000: Post-Doctoral Research Assistant, Department of Plant Sciences, University of Oxford.


1999: DPhil – ‘The Systematics of the southern Indian and Sri Lankan Strobilanthinae’. Department of Plant Sciences, University of Oxford. 

1993: Postgraduate Certificate in Education. Oxford University Department of Educational Studies. 

1991: BSc (Hons) Botany. The University of Reading.


Systematics of Convolvulaceae

Convolvulus vidalii

Convolvulus vidalii, endemic to the Western Rif mountains of Morocco © M. Carine

The Convolvulaceae (bindweed or morning glory family) comprises approximately 60 genera and more than 1500 species of herbaceous vines, trees, shrubs and herbs. My research on the family aims to address taxon delimitation problems in the family and investigate the processes of diversification in both temperate and tropical groups. I am particularly interested in the tribes Convolvuleae and Merremieae. 

Traditionally, pollen characters have been considered taxonomically valuable in the classification of the family. I am currently involved in a collaborative project with Dan Austin (University of Arizona) and George Staples (Singapore Botanic Garden) to develop an online 

Convolvulaceae pollen atlas


We are involved in the development of   , a scratchpad site for the family that is being developed by a worldwide network of researchers with interests in the family. 


Merremieae research

Hewittia malabarica

Hewittia malabarica © G. Staples

Ana Simões is a PhD funded by the Fundação para a Ciência e a Tecnologia, Govt. of Portugal, to work at the Natural History Museum on the systematics and evolution of Merremieae. The Merremieae is a taxonomically complex group of genera centred around the pantropical genus Merremia. Ana's project specifically aims to integrate morphology and DNA sequence data to address (i) tribal and generic circumscription, (ii) morphological evolution, (iii) biogeography and (iv) environmental niche occupancy through time.

Ana is registered for a PhD at the University of Reading. She is co-supervised by Alastair Culham (University of Reading) and George Staples (Singapore Botanic Garden).

Convolvuleae research

Convolvulus sabatius

Convolvulus sabatius growing in the Western Rif Mountains in northern Morocco. © M. Carine

The Convolvuleae as currently circumscribed includes the genera Convolvulus, Calystegia and Polymeria


Generic delimitation

Building on phylogenetic work that focussed on the Macaronesian taxa (see below), I have been working with Juana de Egea (NHM), Bob Johnson (Queensland Herbarium, Brisbane) and George Staples (Singapore Botanic Garden) to produce an integrated molecular and morphological study of the Convolvuleae, sampling broadly within the constituent genera (Convolvulus, Calystegia and Polymeria). The aim of this project is to rigorously test generic delimitation in the group and investigate character evolution. 

Maria Teresa Buril, a PhD student at the Universidade Federal de Pernambuco, Recife, Brazil recently joined the group for four months to work on the molecular systematics of Jacquemontia (Convolvulaceae). Teresa’s work at NHM is supported by grants from CAPES and IAPT.



Convolvulus floridus

The Canary Island endemic Convolvulus floridus © F. Rumsey

We have been investigating the evolution of Convolvulus species endemic to Macaronesia. The eleven species endemic to the region constitute a morphologically heterogeneous and taxonomically complex assemblage. Our work has focussed on the relationships and patterns of diversification of the island endemics and investigating the occurrence of hybridization between endemic taxa.

Further alpha taxonomic work on the genus has focussed on North Africa. A revision of the C. sabatius complex was recently published.

In collaboration with Robert Scotland and John Wood (Department of Plant Sciences, University of Oxford), David Harris (Royal Botanic Garden, Edinburgh) and David Mabberley (Royal Botanic Gardens, Kew), I was recently awarded a SynTax grant for a project entitled ‘Foundation Monographs - accelerating the pace of taxonomy’. The project aims to begin the process of developing novel and innovative solutions to address the lack of taxonomic knowledge of species-rich groups of plants and will result in a monograph of Convolvulus.


Calystegia relationships

Reticulate relationships within Calystegia inferred from ITS data. After Brown et al (in press) © M. Carine

Jackie Brown worked on Calystegia for her Masters thesis and demonstrated complex reticulate patterns of relationships among those taxa of the C. sepium complex that occur in Britain. Her work was supported by the Botanical Society of the British Isles and a paper describing the results of her project is now in press. Jackie is now extending this work to determine whether similar processes may explain the taxonomically challenging patterns of variation observed more widely in Calystegia.

Other collaborators involved in this project are Mark Spencer (NHM) and Dick Brummitt (Royal Botanic Gardens, Kew).


Plant diversity of the Macaronesian Islands

My work on island biodiversity focuses on the  Macaronesian region - the archipelagos of the Azores, Madeira, Salvage Islands, Canaries and Cape Verdes, located in the northern Atlantic Ocean. 



Relationships of the Macaronesian flora


Pico Island in the Azores archipelago © M. Carine

I am interested in a range of questions relating to the biogeography of the Macaronesian region:

  • How are the island floras related to continental floras and to what extent have the islands acted as source areas for continental floras?
  • When did Macaronesian lineages diverge from their continental relatives and when did they radiate?
  • Do different plant groups show congruent patterns of relationships?

Current research on the relationships of the flora focuses on investigating phylogenetic and phylogeographic patterns in a number groups to address specific biogeographic questions (multiple congeneric introductions; trans-continental disjunctions). Groups currently being studied include: Azorean endemic Viburnum Macaronesian endemic Helianthemum, Erica arborea and Erica scoparia. 

My collaborators in this research are Fred Rumsey (NHM),  Mónica Moura and Luis Silva (Departamento de Biologia, Universidade dos Açores); Alfredo Reyes-Betancort (Jardín de Aclimatación de La Orotava (ICIA), Tenerife) and Alain Vanderpoorten and Aurélie Désamoré (University of Liège, Institute of Botany)



Endemism and evolution of the Canary Islands endemic flora

Worldmap analysis of endemic diversity in the Canary Islands

Endemic species richness per 10 x 10km grid cell for the Canary Islands. After Reyes-Betancort et al. (2008).

We recently completed a WORLDMAP analysis that investigated patterns of diversity across the Canary Islands. For a copy of the paper click here: Reyes-Betancort et al (2008) PDF (775.3 KB)





A related analysis investigated the delimitation of areas of endemism within the archipelago. Areas of endemism are the fundamental units of cladistic biogeographic analysis. However, there remains no consensus on the most appropriate method for their delimitation. Our work demonstrated that NDM outperforms hierarchical clustering methods in the Canary Islands and using this method we delimited seventeen areas of endemism that provide a framework for further biogeographic analyses.

We are now using our framework of areas of endemism to investigate the role of vicariance in generating disjunct distribution patterns in the Canary Islands endemic flora.  With support from The Royal Society, we are investigating the role of vicariance in the evolution of the endemic flora of the eastern islands of Fuerteventura and Lanzarote. 

Collaborators in this work are: Chris Humphries and Malcolm Penn (NHM); Alfredo Reyes-Betancort, Arnoldo Santos-Guerra and Rosana Guma (Jardín de Aclimatación de La Orotava (ICIA), Tenerife); Juli Caujapé Castells and Ruth Jaén Molina (Jardin Botanico Canario "Viera y Clavijo", Gran Canaria).



Diversity patterns at the regional level: the Azores diversity enigma

Angelica lignescens

Angelica lignescens, endemic to the Azores and distributed on all three sub-archipelagos © M. Carine

The Azorean endemic flora is distinctive when compared to other oceanic archipelagos both in terms of the widespread distribution of endemics across the archipelago and the paucity of evolutionary radiations in the flora.

Island age, lineage age and habitat diversity have been proposed as explanations for these distinctive characteristics but it appears that such explanations are inadequate and we recently suggested that Pleistocene climatic regimes may have contributed to the patterns observed (Carine & Schaefer, in press). Specifically, we hypothesised that climatically-induced episodes of habitat fragmentation and population contractions in the Canaries may have driven the rapid evolution of single island endemics, a process largely absent from the Azores flora. 

However,  the Azorean flora remains relatively poorly known and we are therefore working with colleagues in the Azores and elsewhere to document diversity patterns in the Azores flora and further investigate differences in diversity and the underlying processes responsible for generating that diversity in the Canaries and Azores. In so doing, we aim to address the question: what drives the origin of morphological and molecular diversity in oceanic islands?



I am a collaborator in the Azores Government-funded Veronica project that is investigating relationships and diversity of Azorean endemic taxa of conservation concern using molecular tools (project led by Mónica Moura, Maria Graciete Belo Maciel and Luis Silva (Departamento de Biologia, Universidade dos Açores). As part of this project, we are investigating the taxonomy and evolution of Azorean endemic Leontodon. With Hanno Schaefer (Imperial College, London) I am also working on a CEE-funded project to examine molecular diversity in a range of Azorean endemic taxa.  




The discovery of the Macaronesian flora

Sir Hans Sloane's collections in the Special Collections Room

Charlie Jarvis, Arnoldo Santos Guerra and Mark Carine in the Botany Department's Special Collections Room examining the collections made by Sir Hans Sloane in Madeira in 1687. © M. Carine

The BM herbarium is a unique resource for the Macaronesian flora and it is particularly rich in historical material, including many types.

Working with colleagues in Madeira, the Canaries and USA, we are documenting and researching our historical Macaronesian collections. At present, we are focussing particularly on Sir Hans Sloane, who visited Madeira in 1687, James Cuninghame who collected on the island of La Palma in 1697 - 1698 and Frances Masson, who spent three years plant hunting in the region between 1776 and 1779. 

I am collaborating in this work with Charlie Jarvis (NHM), Miguel Menezes de Sequeira (Departamento de Biologia Universidade da Madeira), Arnoldo Santos Guerra (Jardín de Aclimatación de La Orotava, Tenerife) and Javier Francisco-Ortega and Mike Maunder (Department of Biological Sciences, Florida International University & Center for Plant Conservation, Fairchild Tropical Botanic Garden).

Other interests

Systematics and ecology of Strobilanthes s.l.

Strobilanthes lupulina

Strobilanthes lupulina photographed in the Silent Valley, Kerala. © M. Carine

Strobilanthes s.l. comprises approximately 450 species worldwide. More than seventy are distributed in the peninsular Indian region and the systematics of these taxa was the focus of my doctoral research. I maintain an interest in the taxonomy and evolution of the group. 

One species of Strobilanthes, S. kunthiana is well known throughout India for its remarkable plietesial life history strategy even though it occurs only in the southern Western Ghats. Every twelve years, these large and woody shrubs flower, set seed and then die en masse, to be replaced by seedlings of the next generation. Where it occurs, S. kunthiana dominates large tracts of vegetation and both flowering events and subsequent die-back are spectacular. The regular and predictable twelve-year flowering cycle of this species has been well documented since 1838. The most recent occurred in 2005—2006 and flowering events are still widely reported in the Indian national press: 

The Hindu


Most other species of Strobilanthes in peninsular India are also thought to be plietesial. However, in marked contrast to the extremely well known and well-characterised flowering cycle of S. kunthiana, there is an almost total absence of robust field data on the flowering of other species. Indeed, many species are very poorly known with no fewer than 23 species (~30%) each known from less than 5 herbarium specimens, most of which were collected more than 60 years ago. 


DNA barcoding

DNA barcoding is the use of a short DNA sequence or sequences from a standardized locus (or loci) as a species identification tool. I have recently been involved in two projects related to DNA barcoding:

Establishing a standard DNA barcode for land plants

I was recently involved in a project funded by the Alfred P. Sloan and Gordon and Betty Moore Foundations  to investigate DNA regions for their potential as barcodes for all land plant species. More details about the project can be found here: 

Establishing a standard DNA barcode for land plants

More information about DNA barcoding in plants can be found here: 

CBOL Plant Working Group



Taxonomy at a Crossroads: Science, Publics and Policy in Biodiversity

I was also a researcher on a 3 year interdisciplinary research project funded by the UK Economic and Social Research Council that brought together social and natural scientists to explore contemporary innovation in the taxonomic sciences, focusing specifically on DNA barcoding. The project was undertaken by a team of social scientists at Lancaster University in collaboration with a team of natural scientists at the Natural History Museum, London. More details about the project can be found here: 

Taxonomy at a crossroads



Schaefer, H., Moura, M., Belo Maciel, M. G. Silva, L., Rumsey, F.J. Carine, M.A. (2011). The Linnean shortfall in oceanic island biogeography: a case study in the Azores. Journal of Biogeography 38: 1345-1355.

Vanderpoorten, A, Laenen, B., Gabriel, R., Gonzalez-Mancebo, R., Rumsey, F. J. & Carine, M. A. (2011). Dispersal, diversity and evolution of the Macaronesian cryptogamic floras. In Plant and Islands (ed. D. Bramwell and J. Caujape-Castells): 338-364.

Désamoré, A., Laenen, B., Devos, N., Popp, M., Gonzaléz-Mancebo, J-M., Carine, M.A. Vanderpoorten, A. (2011). Out of Africa: north-westwards Pleistocene expansions of the heather Erica arborea. Journal of Biogeography 38: 164—176.


Bebber, D., Carine, M.A., Wortley, A.,  Harris, D., Word, J.R.I., Prance, G., Davidse, G. Paige, J., Pennington, T., Robson, N.K.B. & Scotland, R.W. (2010). Herbaria are a major frontier for species discovery. Proceedings of the National Academy of Sciences USA 107: 22169-22171.

Carine, M. A. & Robba, L. (2010). A revision of the Convolvulus sabatius (Convolvulaceae) complex. Phytotaxa 14: 1—21.

Laenen, B., Désamoré, A., Devos, N., Shaw, A. J., González-Mancebo, J-M., Carine, M. A. & Vanderpoorten, A. (2010). Macaronesia: a source of hidden genetic diversity for post-glacial recolonization of western Europe in the leafy liverwort Radula lindenbergiana. Journal of Biogeography 38: 631-639.

Carine, M.A., Santos Guerra, A., Guma, I.R. & Reyes-Betancort, J.A. (2010). Endemism and evolution of the Macaronesian Flora. In: Beyond Cladistics: The Branching of a Paradigm (ed. D.M. Williams & S. K. Knapp): 101-124

Francisco-Ortega, J., Santos Guerra, A., Jarvis, C. E., Carine, M.A., Menezes de Sequeira, M. & Maunder, M. (2010). Early British collectors and observers of the Macaronesian flora: from Sloane to Darwin. In: Beyond Cladistics: The Branching of a Paradigm (ed. D.M. Williams & S. K. Knapp): 125-144.

Carine, M.A. & Schaefer, H. (2010). The Azores diversity enigma: why are there so few Azorean endemic flowering plants and why are they so widespread? Journal of Biogeography 37: 77–89

Vanderpoorten, A., Carine, M.A., Gradstein, R., Devos, N. (2010). The ghosts of Gondwana and Laurasia in modern liverwort distributions. Biological Reviews 85: 471—487.

Menezes de Sequeira, M., Santos-Guerra, A., Jarvis, C.E., Oberli, A., Carine, M.A., Maunder, M. & Francisco-Ortega, J. (2010). Pre-Linnaean descriptions of Macaronesian plants: Sir Hans Sloane’s collections and his account of a brief visit to Madeira in 1687. Taxon 59: 598-612


CBOL Plant Working Group [Hollingsworth, P., Forrest, L., Spouge, J., Hajibabaei, M., Ratnasingham, S. , Van der Bank, M., Chase, M., Cowan, R. , Erickson, D., Fazekas, A., Graham, S., James, K.E. , Kim, K-J., John, W., Schneider, H., Van Alphen Stahl, J., Barrett, S., Van den Berg, C., Bogarin, D., Burgess, K., Cameron, K., Carine, M., Chacón, J., Clark, A., Clarkson, J., Conrad, F., Devey, D., Ford, C., Hedderson, T., Hollingsworth, M., Husband, B., Kelly, L., Kesanakurti, P., Kim, J., Kim, Y., Lahaye, R., Lee, H-L., Long, D., Madriñán, S., Maurin, O., Meusnier, I., Newmaster, S., Park, C-W., Percy, D., Petersen, G., Richardson, J., Salazar, G., Savolainen, V., Seberg, O., Wilkinson, M., Yi, D-K., Little, D.] (2009). A DNA Barcode for Land Plants. Proceedings of the National Academy of Sciences USA 106: 12794—12797.

Brown, J.M., Brummitt, R.K., Spencer, M. & Carine M.A. (2009). Disentangling the bindweeds: hybridization and taxonomic diversity in Calystegia R.Br. Botanical Journal of the Linnean Society 160: 388–401.

Martínez Rodríguez, J., Vargas Gómez, P., Carine, M.A., Jury, S.L. (2009). Taxonomic relationships of a new species of Iris subgenus Xiphium (Iridaceae) from Mediterranean Morocco. Candollea 64: 127-132.

Carine, M.A., Humphries, C.J, Guma, I.R., Reyes-Betancort, J.A. & Santos Guerra, A. (2009). Areas and algorithms: numerical approaches to the delimitation of areas of endemism in the Canary Islands archipelago. Journal of Biogeography 36: 593—611.


Francisco-Ortega, J., Santos Guerra, A., Carine, M.A. & Jarvis, C. E. (2008). Plant hunting in Macaronesia by Francis Masson: the plants sent to Linnaeus and Linnaeus fil. Botanical Journal of the Linnean Society 157: 393—428.

Reyes-Betancort, J.A., Santos Guerra, A., Guma, I.R., Humphries, C.J. & Carine, M.A. (2008) Diversity, rarity and the evolution and conservation of the Canary Islands endemic flora. Anales del Jardín Botánico de Madrid 65: 25—45.


Chase, M.W., Cowan, R.S., Hollingsworth, P.M., van den Berg, C., Madriñán, S.,; Petersen, G.; Seberg, O., Jørgsensen, T., Cameron, K.M., Carine, M.A., Pedersen, N., Hedderson, T.A.J., Conrad, F., Salazar, G., Richardson, J.E.,  Hollingsworth, M.L., Barraclough, T.G., Kelly, L., Wilkinson, M. (2007). A proposal for a standardised protocol to barcode all land plants. Taxon 56: 295—299.

Carine, M.A., Robba, L., Little, R., Russell, S.J. and Santos Guerra, A. (2007). Molecular and morphological evidence for hybridization between endemic Canary Island Convolvulus. Botanical Journal of the Linnean Society 154: 187-204.

Vanderpoorten, A., Rumsey, F. J. and Carine, M.A. (2007). Does Macaronesia exist? Conflicting signal in the bryophyte and pteridophyte floras. American Journal of Botany 94: 625-639.


Carine, M.A., Rumsey, F. J., Ait-Lafkih, M., Rejdali, M., Rutherford, R. W. and Jury, S. L.  (2006). New plant collections from the North Morocco Checklist Area. Lagascalia 26: 196—219.


Carine, M. A. (2005). Spatio-temporal relationships of the Macaronesian endemic flora: a relictual series or window of opportunity? Taxon 54: 895-903.
L. Robba, M. A. Carine, S. J. Russell & F. M. Raimondo (2005) The monophyly and evolution of Cynara L. (Asteraceae) sensu lato: evidence from the Internal Transcribed Spacer region of nrDNA and morphology. Plant Systematics and Evolution 253 (1-4): 53-64.

Silvertown, J., Francisco-Ortega, J. & Carine, M. (2005). The monophyly of island radiations: an evaluation of niche pre-emption and some alternative explanations. Journal of Ecology 93: 653-657.


Carine, M. A., Alexander, J. M. and Scotland, R. W. (2004). A revision of the Strobilanthes kunthiana group (Phlebophyllum sensu Bremekamp). Kew Bulletin 59: 1—25.
Carine, M. A., Francisco-Ortega, J., Santos-Guerra, A. and Russell. S. J. (2004). Relationships of island and continental floras: molecular evidence for multiple colonisations into Macaronesia and subsequent back-colonisation of the continent in Convolvulus L. American Journal of Botany 91: 1070-1085.

Moylan, E. C., Bennett, J. R., Carine, M. A., Olmstead, R. G. and Scotland, R. W. (2004). Phylogenetic relationships among Strobilanthes s.l. (Acanthaceae): evidence from ITS nrDNA, trnL-F cpDNA, and morphology. .American Journal of Botany 91: 724—735.


Carine, M. A., Alexander, J. M. and Russell, S. J. (2003). Evolution of spines and the taxonomic status of Convolvulus L. section Acanthocladi Boiss.: preliminary results from the ITS 2 region of nrDNA. Bocconea 16: 703—710.


Carine, M. A. and Scotland, R. W. (2002). Classification of the Strobilanthinae: trying to classify the unclassifiable? Taxon 51: 259—279.


Scotland, R. W. and Carine, M. A. (2000). Classification or phylogeny estimates? Cladistics 16: 411—419.

Carine, M. A. and Scotland, R. W. (2000). 68 taxa and 32 characters: resolving species relationships using morphological data. In: Harley, M. M., Morton, C. M. and Blackmore, S. (eds). Pollen and Spores: Morphology and Biology. Royal Botanic Gardens, Kew, pp 365—384. Royal Botanic Gardens Kew.

Carine, M. A. and Scotland R. W. (2000). The taxonomy and biology of Stenosiphonium Nees (Acanthaceae). Botanical Journal of the Linnean Society 133: 101—128.

Carine, M. A., Jayasekara, P. and Scotland, R. W. (2000). A new species of Strobilanthes (Acanthaceae) from Sri Lanka. Kew Bulletin 55: 971—976.


Carine, M. A. and Scotland, R. W. (1999). Taxic and transformational homology: different ways of seeing. Cladistics 15: 121—129.


Carine, M. A. and Scotland, R. W. (1998). Pollen morphology of Strobilanthes Blume (Acanthaceae) from southern India and Sri Lanka. Review of Palaeobotany and Palynology 103: 143—165.