Publications by CIRCLE staff

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Measuring and modelling above-ground carbon and tree allometry along a tropical elevation gradient


Emerging international policy aimed at reducing carbon emissions from deforestation and forest degradation (REDD+) in developing countries, has resulted in numerous studies on above-ground live carbon (AGC) in tropical forests. Data presented in this paper shows that forest AGC estimates from East Africa differ by 24% (55t/ha) depending on whether they incorporate height. Regression models in this paper show that the reason for this discrepancy is that tree stature varies with elevation. High AGC was also strongly associated with shallow slopes, followed by intermediate elevation, elephant absence, low potential evapotranspiration and low soil pH. Further regression models also show that the overall AGC is more related to stem size and shape than to the wood density of species present. The paper also discusses the relative benefits of different measurements of topography, soil, and physical disturbance, and recommends a tiered approach to AGC monitoring, depending on available resources.

Also see the publication list, where these data were incorporated into a global analysis showing that our observed 50t/ha difference in biomass estimates when excluding height is typical of tropical forests (Feldpausch et al., 2012).

Hambly, N., Marshall, A.R. (2013) Zoo BAPs: Biodiversity Action Plans for conserving native wildlife in and around zoological gardens. Journal of Zoo and Aquarium Research 2.

Lewis et al. [multi-author] (in press) Aboveground biomass and structure of 259 African tropical forests. Philosophical Transactions of the Royal Society.

Gereau, R.E., Burgess, N.D., Fjeldså, J., Hall, J., Hemp, A., Jump, A.S., Kajuni, A.R., Marchant, R.A., Marshall, A.R., Platts, P.J., Taylor, C.M., Tibazarwa, F.I. (2013) Phylogenetic patterns of extinction risk: the need for critical application of appropriate datasets. PeerJ preprints doi: 10.7287/peerj.preprints.55v1.

Willcock, S., Phillips, O.L., Platts, P.J., Balmford, A., Burgess, N.D., Lovett, J.C., Ahrends, A., Bayliss, J., Doggart, N., Doody, K., Fanning, E., Green, J., Hall, J., Howell, K.M., Marchant, R., Marshall, A.R., Mbilinyi, B., Munishi, P.K.T., Owen, N., Swetnam, R.D., Topp-Jorgensen, E.J., Lewis, S.L. (2012) Towards regional, error-bounded landscape carbon storage estimates for data-deficient areas of the world. PLoS One 7.

Laurance, W.F. [multi-author] (2012) Averting biodiversity collapse in tropical forest protected areas. Nature 489 290-294.

Howlett, C., Marshall, A.R., Hughes, W.O.H. (2012) Digit ratios and dominance in female baboons. International Journal of Primatology 33(6) 1439-1452.

Greve, M., Lykke, A.M., Fagg, C.W., Bogaert, J., Friis, I., Marchant, R., Marshall, A.R., Ndayishimiye, J., Sandel, B.S., Sandom, C., Schmidt, M., Timberlake, J.R., Wieringa, J.J., Zizka, G., Svenning, J.-C. (2012) Continental-scale variability in browser diversity is a major driver of diversity patterns in acacias across Africa. Journal of Ecology 100 1093-1104.

Marshall, A.R., Willcock, S., Platts, P.J., Lovett, J.C., Balmford, A., Burgess, N.D., Latham, J.E., Munishi, P.K.T., Salter, R., Shirima, D.D., Lewis, S.L. (2012) Measuring and modelling above-ground carbon and tree allometry along a tropical elevation gradient. Biological Conservation 154 20-33.

Marshall, A.R., Platts, P.J., Gereau, R.E., Kindeketa, W., Kang’ethe, S., Marchant, R. (2012) The genus Acacia (Fabaceae) in East Africa: distribution, biodiversity and the protected area network. Plant Ecology and Evolution 145(3) 289–301.

Feldpausch, T.R. [multi-author] (2012) Tree height integrated into pantropical forest biomass estimates. Biogeosciences 9 3381–3403.

Deere, N.J. (2011) Exploitation or conservation? Can the hunting tourism industry in Africa be sustainable? Environment: Science and Policy for Sustainable Development 53 20-32.

Marshall, A.R., Deere, N. (2011) Field conservation by BIAZA zoos: how well are we doing? Zoo Research Newsletter 12(4) 2-3.

Fisher, B., Turner, R.K., Burgess, N.D., Swetnam, R.D., Green, J., Green, R.E., Kajembe, G., Kulindwa, K., Lewis, S.L., Marchant, R., Marshall, A.R., Madoffe, S., Munishi, P.K.T., Morse-Jones, S., Mwakalila, S., Paavola, J., Naidoo, R., Ricketts, T., Rouget, M., Willcock, S., White, S., Balmford, A. (2011) Measuring, modeling and mapping ecosystem services in the Eastern Arc Mountains of Tanzania. Progress in Physical Geography 35(5) 595–611.

Shirima, D.D., Munishi, P.K.T., Lewis, S.L., Burgess, N.D., Marshall, A.R., Balmford, A., Swetnam, R., Zahabu, E. (2011) Carbon storage, structure and composition of Miombo woodlands in Tanzania’s Eastern Arc Mountains. African Journal of Ecology 49(3) 332-342.

Platts, J., Burgess, N., Gereau, R., Lovett, J., Marshall, A.R., McClean, C., Pellika, P., Swetnam, R., Marchant, R. (2011) Delimiting tropical mountain ecoregions for conservation. Environmental Conservation 38(3) 312-324.

Feldpausch, T.R. [multi-author] (2011) Height-diameter allometry of tropical forest trees. Biogeosciences 8 1081-1106.

Laurance, W.F. [multi-author] (in press) Global warming, elevational ranges and the vulnerability of tropical biota. Biological Conservation 144 548-557.

Ahrends, A., Rahbek, C., Bulling, M.T., Burgess, N.D., Platts, P.J., Lovett, J.C., Wilkins Kindemba, V., Owen, N., Ntemi-Sallu, A., Marshall, A.R., Mhoro, B.E., Fanning, E., Marchant, R. (2011) Conservation and the botanist effect. Biological Conservation 144 131-140.

Swetnam, R. D., Fisher, B., Mbilinyi, B. P., Munishi, P. K., Willcock, S., Ricketts, T., Mwakalila, S., Balmford, A., Burgess, N. D., Marshall, A. R., Lewis, S. L. (2011) Mapping socio-economic scenarios of land cover change: a GIS method to enable ecosystem service modelling. Journal of Environmental Management 92(3) 563-574.

Burgess, N.D., Bahane, B., Clairs, T., Danielsen, F., Dalsgaard, S., Funder, M., Hagelberg, N., Harrison, P., Haule, C., Kabalimu, K., Kilahama, F., Kilawe, E., Lewis, S.L., Lovett, J.C., Lyatuu, G., Marshall, A.R., Meshack, C., Miles, L., Munishi, P.K.T., Nashanda, E., Shirima, D., Swetnam, R.D., Willcock, S., Williams, A., Zahabu, E. (2010) Getting ready for REDD+ in Tanzania: Progress and Challenges. Oryx 44(3) 339-351.

Platts, P.J., Ahrends, A., Gereau, R.E., McClean, C.J., Lovett, J.C., Marshall, A.R., Pellikka, P.K.E., Mulligan, M., Fanning, E., Marchant, R. (2010) Can distribution models help refine inventory-based estimates of conservation priority? A case study in the Eastern Arc forests of Tanzania and Kenya. Diversity & Distributions 16 628-642.

Marshall, A.R., Jørgensbye, H.I.O., Rovero, F., Platts, P.J., White, P.C.L., Lovett, J.C. (2010) The species-area relationship and confounding variables in a threatened monkey community. American Journal of Primatology 72 325-336.

Swetnam, R.D., Marshall, A.R., Burgess, N.D. (2010) Valuing ecosystem services in the Eastern Arc Mountains of Tanzania. British Ecological Society Bulletin 41(1) 7-9.

Rovero, F., Marshall, A.R. (2009) Camera trapping photographic rate as an index of density in forest ungulates. Journal of Applied Ecology 46 1011-1017.

Menegon, M., Tolley, K., Jones, T., Rovero, F., Marshall, A.R., Tilbury, C.R. (2009) A new species of chameleon (Sauria: Chamaeleonidae: Kinyongia) from the Magombera forest and the Udzungwa Mountains National Park, Tanzania. African Journal of Herpetology 58(2) 59-70.

Rovero, F., Bowkett, A., De Luca, D., Jones, T., Marshall, A.R., Msirikale, J. and Mtui, A. (2007) Camera-trapping to study forest mammals in the Udzungwa Mountains: results from five years of research. Proceedings of the 6th TAWIRI Scientific Conference, Arusha, Tanzania 268-276.

Marshall, A.R. (2008) Assessing and Restoring Biodiversity in Tanzania’s Forests: The Case of Magombera. Proceedings of the 6th TAWIRI Scientific Conference, Arusha, Tanzania 45-84.

Burgess, N., Mwakalila, S., Madoffe, S., Ricketts, T., Olwero, N., Swetnam, R., Mbilinyi, B., Marchant, R., Mtalo, F., White, S., Munishi, P., Marshall, A., Malimbwi, R., Jambiya, G., Fisher, B., Kajembe, G., Morse-Jones, S., Kulindwa, K., Green, J., Balmford, A. (2009) Valuing the Arc – a Programme to Map and Value Ecosystem Services in Tanzania. Mountain Research Initiative Newsletter 18(3) 18-21.

Mwakalila, S., Burgess, N., Ricketts, T., Olwero, N., Swetnam, R., Mbilinyi, B., Marchant, R., Mtalo, F., White, S., Munishi, P., Marshall, A., Malimbwi, R., Smith, C., Jambiya., G., Marshall, A., Madoffe, S., Fisher, B., Kajembe., G., Morse-Jones, S., Kulindwa., K., Green, J., Balmford, A. (2009) Valuing the arc: linking science with stakeholders to sustain natural capital. The Arc Journal 23 23-28.

Topp-Jørgensen, E., Nielsen, M. R., Marshall, A.R., Pedersen, U. (2009) Mammalian density in response to different levels of bushmeat hunting in the Udzungwa Mountains, Tanzania. Tropical Conservation Science 2(1) 70-87.

Rovero, F., Marshall, A.R., Jones, T., Perkin, A. (2009) The primates of the Udzungwa Mountains: diversity, ecology and conservation. Journal of Anthropological Sciences 87(1) 93-126.

Topp-Jørgensen, J.E., Marshall, A.R., Brink, H., Pedersen, U.B. (2008) Quantifying the response of tree hyrax (Dendrohyrax validus) to human disturbance in the Udzungwa Mountains. Tropical Conservation Science 1 63-74.

Marshall, A.R., Lovett, J.C., White, P.C.L. (2008) Selection of line-transect methods for estimating the density of group-living animals: lessons from the primates. American Journal of Primatology 70, 452-462.

Bowkett, A., Rovero, F., Marshall, A.R. (2008) The use of camera-trap data to model habitat selection by antelope species in the Udzungwa Mountain forests, Tanzania. African Journal of Ecology 46(4) 479-487.

Marshall, A.R., Aloyce, Z., Mariki, S., Jones, T., Burgess, N., Kilihama, F., Massao, J., Nashanda, E., Sawe, C., Rovero, F., Watkin, J. (2007) Tanzania’s second Nature Reserve: improving the conservation status of the Udzungwa Mountains? Oryx 41(4) 429-430.

Rovero, F., Struhsaker, T.T., Marshall, A.R., Rynne, T.A., Pedersen, U.B., Butynski, T.M., Ehardt, C.L., Mtui, A.S. (2006) Abundance of diurnal primates in Mwanihana forest, Udzungwa Mountains, Tanzania: a multi-observer comparison of line-transect data. International Journal of Primatology 27(4), 675-698.

Lovett, J.C., Marshall, A.R., Carr, J. (2006) Changes in tropical forest vegetation along an altitudinal gradient in the Udzungwa Mountains National Park, Tanzania. African Journal of Ecology 44, 478-490.

Lovett, J.C., Marshall, A.R. (2006) Why conserve primates? African Journal of Ecology 44(2), 113-115.

Rovero, F., Marshall, A.R. (2005) Diversity and abundance of diurnal primates and forest antelopes in relation to habitat quality: a case study from the Udzungwa Mountains of Tanzania. In: B.A. Huber, B.J. Sinclair, K.-H. Lampe (eds.) African Biodiversity: Molecules, Organisms, Ecosystems. Proceedings of 5th International Symposium on Tropical Biology, Koenig Museum, Bonn. Springer Verlag, 297-304.

Marshall, A.R., Topp-Jørgensen, J.E., Brink, H., Fanning, E. (2005) Monkey abundance and social structure in two high elevation forest reserves in the Udzungwa Mountains of Tanzania. International Journal of Primatology 26(1), 127-145.

Rovero, F., Marshall, A.R. (2004) Estimating the abundance of forest antelopes by using line transect techniques: a case from the Udzungwa Mountains of Tanzania. Tropical Zoology 17, 267-277.

Struhsaker, T.T., Marshall, A.R., Detwiler, K., Siex, K., Ehardt, C.L., Lisbjerg, D.D., Butynski, T.M. (2004) Demographic variation among the Udzungwa red colobus (Procolobus gordonorum) in relation to gross ecological and sociological parameters. International Journal of Primatology 25(3), 615-658.

Marshall, A.R. (2004) Ecology and social demography of Udzungwa colobines: a summary of current knowledge. Folia Primatologica 75(suppl 1; Abstracts of the 20th Congress of the International Primatological Society), 23-24.

Brink, H., Topp-Jørgensen, J.E., Marshall, A.R., Fanning, E. (2002) First record in sixty-eight years of Lowe’s servaline genet (Genetta servalina lowei). Oryx 36(4), 324.

Bowkett, A., Marshall, A.R., Plowman, A. (1999) Developing and evaluating an enrichment programme in an ageing monkey house. Proceedings of the 4th International Conference on Environmental Enrichment, Edinburgh, 304-310.[/vc_column_text][/vc_tab][vc_tab title=”BOOKS AND BOOKS CHAPTERS” tab_id=”1386933536-2-74″][vc_column_text]

Boswell&Marshall2011_SavingOurForests_frontCover (1)Saving Our Forests: Why Are Forests Important and What Can We Do to Save Them?


Removal of forest trees for building and fuel is threatening forests across Africa, and this book is designed to help. The book includes a summary of the importance of forests, with a number of tips to help African villagers live sustainably, and hence help to conserve these threatened habitats.

The book is the first in a series produced to support the work of the Udzungwa Forest Project. Each volume will have a regional focus; this one relates to the Kilombero-Udzungwa area of southern Tanzania, and includes a guide to the trees and animals of the region.

Available in English or Swahili, in return for a voluntary donation to UFP (contact


Boswell, A.K., Marshall, A.R., Njilima, F.M. (in press) Tuitunze Misitu Yetu: Kwa nini Misitu ni Muhimu na Tufanye nini Kuitunza?. CIRCLE, York, ISBN TBC.

Marshall, A.R., Rovero, F., Struhsaker, T.T. (in press) Procolobus gordonorum. In: N. Rowe (ed.) All the World’s Primates. Pogonias Press, Charlestown, Rhode Island, USA.

Marshall, A.R., Fashing, P. (in press) Colobus angolensis palliatus. In: N. Rowe (ed.) All the World’s Primates. Pogonias Press, Charlestown, Rhode Island, USA.

Boswell, A.K., Marshall, A.R. (2011) Saving Our Forests: Why Are Forests Important and What Can We Do to Save Them? CIRCLE, York, ISBN 978-0-9569862-0-7.

Marshall, A.R., Munishi, P.K.T. (2011) Valuing the Arc: measuring and monitoring forest carbon for offsetting. In: Kareiva, P., Ricketts, T., Daily, G., Tallis, H., Polasky, S. (eds.) The Theory and Practice of Ecosystem Service Valuation. Oxford University Press, Oxford.

Lovett, J.C., Marchant, R., Marshall, A.R., Barber, J. (2007) Tropical Moist Forests. In: R. Hesler and Harrison (eds.) Biodiversity Under Threat. Royal Society of Chemistry 161-192.[/vc_column_text][/vc_tab][vc_tab title=”TECHNICAL REPORTS” tab_id=”1386934099514-2-6″][vc_column_text]

Circle- Flamingo Land BAP

Flamingo Land Biodiversity Action Plan


The Flamingo Land Biodiversity Action Plan (BAP) has been developed by CIRCLE (Centre for the Integration of Research, Conservation and Learning), a joint collaboration between Flamingo Land Theme Park and Zoo and the University of York’s Environment Department. Advice was given by representatives from the Ryedale BAP, local naturalists and staff at Flamingo Land. Read more>>>

Marshall, A.R., Wilson, S., Needham, N. (2013) Top Ten Mammal Species Reliant on Zoos. BIAZA Field Programme Committee.

Marshall, A.R., Robinson, J.E., Hindle, B. (2012) Top Ten Species Reliant on Zoos. BIAZA Field Programme Committee.

Hambly, N., Marshall, A.R. (2011) Flamingo Land Biodiversity Action Plan: 2011-2016. CIRCLE, York.

Marshall, A.R., Platts, P.J., Marchant, R., Gereau, R., Kindeketa, W. (2009) Conserving biodiversity along the Kenya-Tanzania border in the face of climate change. Report to the Liz Claiborne Art Ortenburg Foundation.

Marshall, A.R. (2008) Ecological Report on Magombera Forest. Worldwide Fund for Nature – Tanzania Programme Office.

Marshall, A.R., Mtoka, S.N. (2008) Taarifa ya Kiikolojia ya Msitu wa Magombera. Report to the Critical Ecosystem Partnership Fund, WWF Tanzania Programme Office, Dar es Salaam, Tanzania.

Marshall, A.R. (2007) Disturbance in the Udzungwas: Responses of Monkeys and Trees to Forest Degradation. Ph.D. Thesis, University of York, UK.[/vc_column_text][/vc_tab][vc_tab title=”PhD THESES” tab_id=”1404145310650-4-8″][vc_column_text]

CIRCLE-Latham2013_PhDthesisCover_225pxLatham, J.E. (2013) Evaluating failures in tropical forest management: Incorporating local perspectives into global conservation strategies.


This thesis makes a rare attempt to consider the needs of local people living alongside conservation areas in Africa. The findings show how forest conservation policy has evolved from a purely wildlife-based approach, to a multi-level approach aiming to help wildlife, livelihoods, and local economy. The realities however are much more complicated, and the thesis demonstrates that people are not always aware of conservation management, and are often left with limited natural resources and income, as a result of forest protection.

Overall, the thesis shows that forest conservation and management interventions need to account for the perspectives and needs of local forest-adjacent communities. Education, governance and forest dependency were identified as the three main areas of concern among local people.


Latham, J.E. (2013) Evaluating failures in tropical forest management: Incorporating local perspectives into global conservation strategies. Ph.D. thesis, University of York.[/vc_column_text][/vc_tab][vc_tab title=”GENERAL PUBLICATIONS” tab_id=”1386934100815-3-0″][vc_column_text]World zoo publication database in preparation.[/vc_column_text][/vc_tab][/vc_tabs][/vc_column][/vc_row]

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