Dr Paul Ramsay

Dr Paul Ramsay

Associate Professor in Ecology

School of Biological & Marine Sciences (Faculty of Science & Engineering)


Programme Leader of BSc (Hons) Conservation Biology


Deputy Head
School of Biomedical & Biological Sciences, University of Plymouth

Associate/Acting Head
School of Biological Sciences, University of Plymouth

1993- 2006

Lecturer/Senior Lecturer in Ecology

School of Biological Sciences, University of Plymouth


Visiting Lecturer in Ecology, Biogeography Laboratory, Department of Ecology and Natural Resources, Universidad Nacional Autónoma de México, Mexico City


Consultant Ecologist, Hunting Land & Environment Limited (now HTS Development Limited).


PhD in plant ecology, University of Wales, Bangor. (The Páramo Vegetation of Ecuador: the Community Ecology, Dynamics and Productivity of Tropical Grasslands in the Andes)


BSc (Hons) Biology, University of Wales, Bangor.

Professional membership

Fellow of the Society of Biology Member of the British Ecological Society

Teaching interests

My teaching interests reflect my research themes: biodiversity and conservation. During my time at the University of Plymouth, I have contributed a wide variety of subject matter on many different courses. I currently contribute to three modules in our postgraduate programmes and to nine undergraduate modules in our BSc (Hons) programmes in Environmental Biology, Marine Biology, Marine Biology & Coastal Ecology, and Biological Sciences.

I have taught on the following modules: Adaptations of Plants and Insect Herbivores; Biological Diversity; Biological Sciences Field Course (Azores); Coastal Ecosystems; Community Ecology; Concepts in Ecology; Conservation; Contaminated Land Remediation; Ecology Fieldcourse (UK, Spain, Borneo, Mexico); Ecology: Applications and Developments; Ecology and Conservation; Environmental Biology Field Course (Azores); Field Biology of Plants; Fundamentals of Ecology; Global Ecology; Habitat Restoration; Individuals, Populations and Communities; Introduction to Ecology and Evolution; Methods in Biology; Methods in Ecology; Restoration Ecology; Skills in Ecology; Skills in Marine Biology and Coastal Ecology; Research Skills and Methods in Biology; Terrestrial Ecosystems; Terrestrial Environments.

Research interests

My work is mostly focused on biodiversity and conservation management in two distinct geographical areas: the Andes and European uplands. However, I started off my career as a marine biologist and I have also been involved in several projects on aquatic ecology. Before joining the University of Plymouth, I worked for four years as an environmental consultant and I appreciate the need for practical advice on biodiversity conservation.

1. Biodiversity and sustainable use of tropical alpine grasslands in the Andes

Over 20 years ago, I carried out the first comprehensive survey of the vegetation of the high-altitude grasslands (páramos) of Ecuador. I also studied plant population dynamics in relation to agricultural fires. Subsequently, my interests broadened to include invertebrates and birds, and other Andean countries. Currently, I am involved in diverse projects in Venezuela, Colombia, Ecuador and Peru: plant community ecology, spatial relationships between plants, and hummingbird-plant interactions.

2. Biodiversity and sustainable use of high-altitude forest remnants in the Andes

Alongside the páramo grasslands are high-altitude forest patches. Since 1985 I have carried out investigations in these Polylepis woodlands—the highest woodlands in the world. Usually small and isolated, these woodlands are under threat from the gradual loss of trees at the woodland edge during grassland fires and from fuelwood collection by poor, high-altitude farmers. My current work is in Peru and Ecuador.

There are three themes to my work:

a. The population dynamics of Polylepis itself
How does Polylepis regenerate inside the woodland? Why does it fail to recolonise areas outside the woodland boundaries? How quickly do the trees grow? How old are they?

b. Biodiversity of Polylepis woodlands
What species are found in these patches? How does the biodiversity relate to “forest quality”?

c. Conservation of Polylepis woodlands
What are the main threats to Polylepis woodlands? How have the woodlands changed over time? How can we monitor the success of Polylepis woodland conservation and restoration? Part of this work involves working alongside local communities to provide alternatives to Polylepis fuelwood, and to minimise the impact of ongoing and past loss through replanting. This work also provides an opportunity for an occasional, humbling game of football at 4100 m!

I am also supervising a project in the Colombian Andes, looking at the ecology of two species of threatened parakeets (Leptosittaca branickii and Bolborhynchus ferrugineifrons) and evaluating different approaches to habitat assessment and management for these birds.

3. Conservation and habitat management

Another research theme involves the study of particular organisms of conservation importance. Particular plants and animals have a greater significance for research than others because of their ecological, economic or cultural importance. I am particularly interested in spatial relationships between species, and the conservation importance of these relationships. Some of this work involves relatively complex spatial analysis and modelling.

Current projects with plants involve Espeletia and Puya (high-altitude giant rosette plants in Ecuador, Colombia and Venezuela), Euphrasia vigursii (an endemic plant of south-western Britain), and Berberis holstii (a medicinal shrub in Malawi). Projects with animals include Bolborhynchus ferrugineifrons and Leptosittaca branickii (two vulnerable parakeets in Colombia), Ctenosaura pectinata (the Black Iguana, eaten by people in Mexico), and Danaus plexippus (the monarch butterfly, whose overwintering sites in Mexico are under threat).

Conservation management is another element of my research interests. In particular, I am investigating the relationship between land use and conservation. I have been involved in surveys of Mexican mountain forests, some with regard to the monarch butterflies mentioned earlier. In the UK, I have worked with postgraduate students on grazing management and its effects on plant communities. We are evaluating the impact of mowing as a surrogate for grazing in Dartmoor National Park. In the Andes, I have studied the effects of fires on plant and animal communities.

Restoration of damaged ecosystems is another area of interest for me. I am currently involved in forest restoration projects in Ecuador and Peru, both working with local people to replant native trees for conservation and ecosystem services.

4. Approaches to studying biodiversity that do not involve species identification

I often work in places where the taxonomy of the organisms I meet is poorly understood. This has obvious implications for the kinds of studies that are possible within reasonable time frames. One solution to this problem is the use of functional types: groups of organisms with similar resource use and responses to environmental change. With this approach, it is possible to compare different ecosystems with few, or no, species in common. It also permits ecological studies with groups that have not been described adequately in taxonomic terms. However, the obvious question with such studies is, “Do functional types provide useful information, or information comparable to that obtained using traditional taxonomic approaches?” Part of my interest, therefore, is to explore this basic question. In a similar way, I have looked at the suitability of using supra-specific identifications for studying biodiversity.

I am tackling more specific issues, too. I have defined plant functional types using morphological and ecophysiological traits in Ecuador, Spain and Mexico. Using this approach I have tested ideas about evolutionary convergence at high altitudes (Ecuador) and the detection of plant communities with functional types (Picos de Europa, Spain). I have also worked with biomass-size spectra (how biomass is divided up between organisms of different sizes) to describe communities of aquatic invertebrates in river, estuary and marine benthos. Again, I have used this approach to consider similarities and differences in systems with completely different taxa to test hypotheses about community structure.

Research degrees awarded to supervised students

Current graduate students

Bejarano-Bonilla DA (in preparation) The relationship of an endemic and threatened parakeet, Bolborhynchus ferrugineifrons Lawrence 1880, with habitat quality and land use in the Colombian Andes. PhD, University of Plymouth.

Maliwichi-Nyirenda C (in preparation) Population variation and conservation status of Berberis holstii in Nyika National Park, Malawi. PhD, University of Plymouth.

Ng S (in preparation) Aquatic macrophyte complexity and pond invertebrate diversity. MSc thesis, University of Plymouth.

White T (in preparation) Nectar production in a giant rosette plant (Puya hamata) in the Andes. MSc thesis, University of Plymouth.
Completed graduate students

Aguirre V (2008) Demography and genetic diversity of the Mexican black iguana, Ctenosaura pectinata, PhD, University of Plymouth.

Avila V (2004) The use of forest authenticity to determine conservation and restoration priorities in the upper Río Magdalena catchement, México. Masters in Restoration Ecology, Universidad Nacional Autónoma de México.

Amblin E (2005) Common dolphin distribution in relation to environmental features in the Celtic Deep. MSc thesis, University of Plymouth.

Biggi GF (2002) Characterisation of Emiliana huxleyi host-virus dynamic. MSc thesis, University of Plymouth.

Brunet M (2005) Habitat use by otters in Loe Pool, Cornwall. MSc thesis, University of Plymouth.

Clarke E (2006) Age-size relationships, growth rates and sustainable management of high-altitude Polylepis trees in the Cordillera de Vilcanota, Perú. MSc thesis, University of Plymouth.

Denning TMW (2003) The effects of the invasive terrestrial Amphipod, Arcitalitrus dorrieni, on native leaf litter invertebrate communities and organism diversity in the Plymouth area. MSc thesis, University of Plymouth.

Doidge K (2006) Fine-scale habitat preferences of Vigur’s Eyebright (Euphrasia vigursii) on Lydford High Down, Dartmoor. MSc thesis, University of Plymouth.

Edwards CJ (2002) China clay sand waste-tip restoration to lowland heath using a water tyreatment works sludge as a soil substitute: seedling recruitment and development. MSc thesis, University of Plymouth.

Fotherby RM (2004) The spatial distribution of Vigur’s Eyebright (Euphrasia vigursii) on Lydford High Down, Dartmoor. MSc thesis, University of Plymouth.

Green JJ (2003) A critical examination of managed realignment with particular focus on the Southwest of the United Kingdom. MSc thesis, University of Plymouth.

Harris R (2005) Does structural complexity affect spider diversity and assemblage structure? MSc thesis, University of Plymouth.

Haynes T (2006) The response of Vigur’s Eyebright (Euphrasia vigursii) to a management experiment on Lydford High Down, Dartmoor. MSc thesis, University of Plymouth.

Humphries N (2006) Spatial interactions between limpets and fucoid seaweeds: relationships feedbacks and climate change. MSc thesis, University of Plymouth.

Jameson J (2005) An investigation into the changes in Polylepis forest cover in the Cordillera Vilcanota, Peru, 1956–2005 based on remote sensing data. MSc thesis, University of Plymouth.

Kidney D (2002) The effects of anthropogenic disturbance on lepidopteran communities in high-altitude temperate forest in Mexico. MSc thesis, University of Plymouth.

Mata-Zayas E (2007) The Distribution of Phylogenetic Diversity of Mammals in Mexico and its Implications for Conservation. PhD, University of Plymouth.

Sands D (2005) Does the fractal complexity of Laminaria hypoborea holdfasts have an influence on species diversity? MSc thesis, University of Plymouth.

Stewart G (2002) Grazing Management on Bodmin Moor. PhD thesis, University of Plymouth.

Summers, S (2006) Climate change and Laminaria interactions in the south-west of England. MSc thesis, University of Plymouth.

Wheeldon RC (2002) The impacts of forest disturbance on bird species richness and composition in a Mexican montane forested catchment. MSc thesis, University of Plymouth.

Postdoctoral Research Fellows

Duckworth JC (1998-1999) The role of plant functional types in the description of mountain ecosystem biodiversity for conservation. Jointly with Prof. Martin Kent, School of Geography, University of Plymouth. Funding from University of Plymouth.

Key publications are highlighted

Smith AM & Ramsay PM 2018 'A comparison of ground-based methods for estimating canopy closure for use in phenology research' Agricultural and Forest Meteorology 252, 18-26 Author Site , DOI PEARL
Zomer MA & Ramsay PM 2018 'Espeletia giant rosette plants are reliable biological indicators of time since fire in Andean grasslands' Plant Ecology 1-10 , DOI PEARL
Ramsay PM 2015 'Handbook of spatial point-pattern analysis in ecology' JOURNAL OF APPLIED STATISTICS 42, (4) 916-917 Author Site , DOI
Barnett AA, Silva WS, Shaw PJA & Ramsay PM 2015 'Inundation duration and vertical vegetation zonation: a preliminary description of the vegetation and structuring factors in borokotóh (hummock igapó), an overlooked, high-diversity, Amazonian vegetation association' Nordic Journal of Botany 33, (5) 601-614 , DOI
Peng Y, Lachmuth S, Gallegos SC, Kessler M, Ramsay PM, Renison D, Suarez R & Hensen I 2015 'Pleistocene climatic oscillations rather than recent human disturbance influence genetic diversity in one of the world's highest treeline species' American Journal of Botany 102, (10) 1676-1684 , DOI
Peyre G, Balslev H, Martí D, Sklenár P, Ramsay P, Lozano P, Cuello N, Bussmann R, Cabrera O & Font X 2015 'VegPáramo, a flora and vegetation database for the Andean páramo' Phytocoenologia 45, (1) 195-201 , DOI
Ramsay PM 2014 'Giant rosette plant morphology as an indicator of recent fire history in Andean páramo grasslands' Ecological Indicators 45, 37-44 , DOI
Pitman RT, Mulvaney J, Ramsay PM, Jooste E & Swanepoel LH 2014 'Global Positioning System-located kills and faecal samples: A comparison of leopard dietary estimates' Journal of Zoology 292, (1) 18-24 , DOI
García-Meneses PM & Ramsay PM 2014 'Puya hamata demography as an indicator of recent fire history in the páramo of El Ángel and Volcán Chiles, Ecuador-Colombia' Caldasia 36, (1) 53-69 , DOI
Pitman RT, Kilian PJ, Ramsay PM & Swanepoel LH 2013 'Foraging and habitat specialization by female leopards (Panthera pardus) in the Waterberg Mountains of South Africa' SOUTH AFRICAN JOURNAL OF WILDLIFE RESEARCH 43, (2) 167-176 Author Site , DOI
García-Meneses PM & Ramsay PM 2012 'Pollinator response to within-patch spatial context determines reproductive output of a giant rosette plant' Basic and Applied Ecology 13, (6) 516-523 , DOI
Pitman RT, Swanepoel LH & Ramsay PM 2012 'Predictive modelling of leopard predation using contextual Global Positioning System cluster analysis' JOURNAL OF ZOOLOGY 288, (3) 222-230 Author Site , DOI
Santini G, Ramsay PM, Tucci L, Ottonetti L & Frizzi F 2011 'Spatial patterns of the ant Crematogaster scutellaris (Hymenoptera, Formicidae) in a model ecosystem' Ecological Entomology 36, 625-634 , DOI
Bilton DT, Mcabendroth LC, Nicolet P, Bedford A, Rundle SD, Foggo A & Ramsay PM 2009 'Ecology and conservation status of temporary and fluctuating ponds in two areas of southern England' Aquatic Conservation: Marine and Freshwater Ecosystems 19, (2) 134-146 , DOI
Jameson JS & Ramsay PM 2007 'Changes in high-altitude Polylepis forest cover and quality in the Cordillera de Vilcanota, Perú, 1956-2005' Biological Conservation 138, (1-2) 38-46 , DOI
Ramsay PM & Fotherby RM 2007 'Implications of the spatial pattern of Vigur's Eyebright (Euphrasia vigursii) for heathland management' Basic and Applied Ecology 8, (3) 242-251 , DOI
Abellán P, Bilton DT, Millán A, Sánchez-Fernández D & Ramsay PM 2006 'Can taxonomic distinctness assess anthropogenic impacts in inland waters? A case study from a Mediterranean river basin' Freshwater Biology 51, (9) 1744-1756 , DOI
Bilton DT, Mcabendroth L, Bedford A & Ramsay PM 2006 'How wide to cast the net? Cross-taxon congruence of species richness, community similarity and indicator taxa in ponds' Freshwater Biology 51, (3) 578-590 , DOI
Ramsay PM, Kent M, Reid CL & Duckworth JC 2006 'Taxonomic, morphological and structural surrogates for the rapid assessment of vegetation' JOURNAL OF VEGETATION SCIENCE 17, (6) 747-754 Author Site , DOI
McAbendroth L, Ramsay PM, Foggo A, Rundle SD & Bilton DT 2005 'Does macrophyte fractal complexity drive invertebrate diversity, biomass and body size distributions?' Oikos 111, (2) 279-290 , DOI
Aucca C & Ramsay PM 2005 'Management of biodiversity and land use in Southern Peru: ECOAN's activities to help conserve Polylepis woodlands' Mountain Research and Development 25, (3) 287-289 , DOI
Ramsay PM 2004 'Consequences of being a plant' Journal of Biogeography 31, (1) 174-175 , DOI
Ramsay PM & Oxley ERB 2001 'An assessment of aboveground net primary productivity in Andean grasslands of central Ecuador' MOUNTAIN RESEARCH AND DEVELOPMENT 21, (2) 161-167 Author Site , DOI
Sklenár P & Ramsay PM 2001 'Diversity of zonal páramo plant communities in ecuador' Diversity and Distributions 7, (3) 113-124 , DOI
Duckworth J, Kent M & Ramsay P 2000 'Plant functional types: an alternative to taxonomic plant community description in biogeography?' Progress in Physical Geography 24, (4) 515-542 , DOI
Woods S, Ortiz-Crespo F & Ramsay PM 1998 'Presence of the Giant Hummingbird, Patagona gigas, and the Ecuadorian Hillstar, Oreotrochilus chimborazo jamesoni, at the Ecuador-Colombia Border' Cotinga 10, 37-40
Owen N, Kent M, Dale P & Ramsay PM 1998 'The vegetation of Kildonan machair, Isle of South Uist, Outer Herbrides, Scotland' Scottish Geographical Magazine 114, 34-41
Ramsay PM, Rundle SD, Attrill MJ, Uttley MG, Williams PR, Elsmere PS & Abada A 1997 'A rapid method for estimating biomass size spectra of benthic metazoan communities' CANADIAN JOURNAL OF FISHERIES AND AQUATIC SCIENCES 54, (8) 1716-1724 Author Site , DOI
Ramsay PM 1997 'Flexible student-centred learning in the field' Ethos 13,
Rundle SD & Ramsay PM 1997 'Microcrustacean communities in streams from two physiographically contrasting regions of Britain' Journal of Biogeography 24, (1) 101-111
Ramsay PM & Oxley ERB 1997 'The growth form composition of plant communities in the Ecuadorian paramos' Plant Ecology 131, (2) 173-192 , DOI
Attrill MJ, Ramsay PM, Thomas RM & Trett MW 1996 'An estuarine biodiversity hot-spot' Journal of the Marine Biological Association of the United Kingdom 76, (1) 161-175
Ramsay PM & Oxley ERB 1996 'Fire temperatures and postfire plant community dynamics in Ecuadorian grass páramo' Vegetatio 124, (2) 129-144
Fisher GG, Clark L & Ramsay PM 1991 'Pesticides in a chalk catchment: inputs and aquatic residues' British Crop Protection Council Monographs 47, 193-200
Ramsay PM 2008 'Gradients in a tropical mountain ecosystem of Ecuador' Mountain Research & Development 28, (3/4) 340-341 , DOI
Ramsay PM 2001 The Ecology of Volcán Chiles: high-altitude ecosystems on the Ecuador-Colombia border. Plymouth Pebble & Shell
Hare J & Ramsay PM 2001 'Aquatic invertebrate communities of Ecuadorian waterfall pools: the effects of altitude and associated variables' in Ramsay PM The Ecology of Volcán Chiles: high-altitude ecosystems on the Ecuador-Colombia border Plymouth Pebble & Shell 193-199
Ramsay PM 2001 'Diurnal temperature variation in the major growth forms of an Ecuadorian páramo plant community' in Ramsay PM The Ecology of Volcán Chiles: high-altitude ecosystems on the Ecuador-Colombia border Plymouth Pebble & Shell 101-112
Smithers P, Ramsay PM, Bond AN & Burne ME 2001 'Macro-arthropod communities of the giant rosette plant, Espeletia pycnophylla subsp. angelensis' in Ramsay PM The Ecology of Volcán Chiles: high-altitude ecosystems on the Ecuador-Colombia border Plymouth Pebble & Shell 169-175
Oxbrough AG & Ramsay PM 2001 'Páramo spiders of Volcán Chiles, Ecuador' in Ramsay PM The Ecology of Volcán Chiles: high-altitude ecosystems on the Ecuador-Colombia border Plymouth Pebble & Shell 159-168
Ramsay PM 2001 'Páramo vegetation recovery in the first two years after a fire on Volcán Chiles, Ecuador' in Ramsay PM The Ecology of Volcán Chiles: high-altitude ecosystems on the Ecuador-Colombia border Plymouth Pebble & Shell 65-73
Ramsay PM & Mitchell P 2001 'The causes of leaning in Espeletia pycnophylla subsp. angelensis' in Ramsay PM The Ecology of Volcán Chiles: high-altitude ecosystems on the Ecuador-Colombia border Plymouth Pebble & Shell 119-124
Ramsay PM 2001 'The zonal páramo vegetation of Volcán Chiles' in Ramsay PM The Ecology of Volcán Chiles: high-altitude ecosystems on the Ecuador-Colombia border Plymouth Pebble & Shell 27-38
Woods S & Ramsay PM 2001 'Variability in nectar supply: implications for high-altitude hummingbirds' in Ramsay PM The Ecology of Volcán Chiles: high-altitude ecosystems on the Ecuador-Colombia border Plymouth Pebble & Shell 209-217
Coombes L & Ramsay PM 2001 'Vegetation of a cushion mire at 3600 m on Volcán Chiles, Ecuador' in Ramsay PM The Ecology of Volcán Chiles: high-altitude ecosystems on the Ecuador-Colombia border Plymouth Pebble & Shell 47-54
Ramsay PM 1999 'La ecología de los páramos ecuatorianos' in Velázquez A Ecología, conservaciٌón y uso sustentable de los ecosistemas montanos neotropicales: memorias de la primera reunión de la red Mexico City Universidad Nacional Autónoma de México 32-46
Ramsay PM 1999 'Landscape mosaics in the High Andes: the role of fire in páramo communities' in Kovář P Nature and Culture in Landscape Ecology: Experiences for the 3rd Millennium Prague The Karolinum Press 192-199
Ramsay PM 1999 'Student ecology project work in Ecuador' in Williams C; Griffiths J; Chalkley B Fieldwork in the Sciences Plymouth Science Education Enhancement and Development and the University of Plymouth 37-39
Ramsay PM & Davis MJ 1992 'Contaminated land remediation' in Final S; Taylor C Environment Solutions: The Directory of Environmental Technology York University of York 92-94