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Annika Felton

Annika Felton
My research into forest ecology has involved a diverse array of forest systems around the world, at a large range of spatial scales; from the inner complexities of animal digestive physiology and plant chemistry, to landscape scale ecological dynamics and their resultant implications for forest and game management. While my projects often specifically regards animal-plant interactions and herbivore nutritional ecology, the outcomes are applicable to the maintenance of biodiversity in managed forest systems, and the ecosystem services we derive from these forests.


My current research involves both ungulates and primates.

Current research

Ungulate ecology

Research into animal-plant interactions often provides insights of direct relevance to society. This is certainly the case in Sweden where large herbivores heavily influence the management and function of our forest ecosystems.  My ongoing research projects in ungulate ecology are:        

Effects of supplementary feeding on moose health and forest damage in southern Sweden

I am currently (2017-2019) running this multidisciplinary project funded by FORMAS (the Swedish Research Council for Environment, Agricultural Sciences and Spatial Planning). Also part of the core team of this project are Dr. Emma Holmström (forestry), Dr. Jonas Malmsten (veterinary science), Dr. Ruth Nichols (genetics) and Dr. Erik Löfmarck (social science).

In this project we combine field inventories of browse availability, ungulate density, browsing pressure and damage in productions stands, with telephone interviews of land owners. We also use DNA meta-barcoding to identify plants eaten by the moose, using both rumen and fecal samples from shot individuals. The aim with the project is to fill key knowledge gaps regarding supplementary feeding by determining the effects on moose condition, diet choice and damage to young and old production forest stands. This project can be seen as a continuation of a previous (2014-2017) large research effort, partly in the same study areas, that involved the help of hundreds of volunteer hunters in southern Sweden: "Understanding the nutritional drivers of moose health and impacts in the landscape".

Interactive effects of agriculture and forestry on red deer damage in the landscape

Together with Dr. Anders Jarnemo (Högskolan i Halmstad) and Dr. Johan Månsson (Dept. Ecology SLU) I conduct a research project about the interactive effects of agriculture and forestry on red deer damage in the landscape, funded by the Swedish EPA (2016-2019). In this project we aim to assist in the development of countermeasures to crop damages and bark stripping on Norway spruce by red deer in Southern Sweden. We do this by identifying various landscape factors that affect the deer's use of different crops and how their intake of crops affect their bark stripping behaviour. Specifically, we combine controlled feeding experiments with captive red deer, with field inventories and spatial analysis of deer movements in the agricultural matrix.

Cost of red deer damage in spruce stand in southern Sweden

In collaboration with the Swedish Forest Agency, Södra Skogsägarna and LRF Skåne, I am running a project (2017-2018) aiming to quantify the cost of red deer bark damage to spruce forests in Skåne, on a stand scale. The output of this project is in the form of an application to smart phones, called "Kronhjortsskador" and is available for both i-phones and andoid telephones. See a news letter (only avaliable in Swedish) about this project.

Beyond Moose

Beyond Moose - interactions among coexisting ungulate species in Swedish landscapes modified by human land use.

I am part of the core team of a large research project called "Beyond Moose", funded by the Swedish EPA, in collaboration with the Dept. of Wildlife, Fish and Environmental studies, SLU Umeå. This is a large study of the interactions among coexisting ungulate species in Swedish landscapes modified by human land use, and how these interactions affect the performance of these species and their impacts on the landscape. We also aim to use this understanding to develop and test sustainable management practices of multi-species ungulate communities. The study involves moose, red deer, roe deer, fallow deer and wild boar, and concerns study locations from the North to the South of Sweden. I am co-supervising PhD student Robert Spitzer in this project. We use eDNA methodology, remote sensing and forest inventories to answer the research questions.


Research projects recently finalized:

Understanding the nutritional drivers of moose health and impacts in the landscape

This large research effort (2014-2017) involved the help of hundreds of volunteer hunters in more than 50 moose management units in southern Sweden. This was a large interdisciplinary project funded by the Swedish Environmental Protection Agency (EPA) and the Södra Skogsägarnas Stiftelse, and carried out in collaboration with the Swedish Veterinary Institute and the Swedish Forest Agency.

We produced new knowledge regarding how the availability of natural browse and supplementary food directly and indirectly influences moose winter diets and their nutritional state, body mass, health and reproduction. The final report to the EPA can be found here. This project is dependent on the voluntary involvement of hundreds of hunters in southern Sweden who deliver samples from shot moose (e.g. rumen content, organs) that we analysed using Near-Infrared Spectroscopy (NIRS), various chemical assays, veterinary assessments of organs, and macrohistology of rumen contents. The Geometric Framework for Nutrition is a central analytical tool in this and earlier projects, as well as using spatial analysis of landscape scale factors (GIS).

To inform hunters how to collect samples from the moose, we distributed instruction videos. We obtained samples from >500 individuals and produced a vast array of new knowledge, much of which has already been communicated to the stakeholders via numerous seminars and media outlets (see for example this news letter, only avaliable in Swedish). Most scientific articles from the project are currently (2018) being produced.

The effect of different types of spruce stand cleaning measures on the creation of ungulate browse

Together with Prof. Urban Nilsson at our department I studied the effect of different types of spruce stand cleaning measures (Sw = röjning) on the creation of birch browse for ungulates. We also identify how the animals utilize the browse produced and whether their browse utilization differs between re-sprouting birch stumps and standing birches. The final report of that project can be found here.


I am also involved in related projects run by colleagues:

Disturbance factors that affect the natural regeneration of oak in forests valued for conservation in Southern Sweden; PhD student Linda Pettersson.

Other examples of my work in ungulate ecology includes feeding experiments with captive moose to identify their nutritional goal and priorities; field experiments that assess how different types of supplementary feeds influence the movements and subsequent browsing behaviour by wild ungulates; laboratory analyses of the nutritional contents of moose’ natural food items, as well as agricultural crops used in fodder fields and supplementary feeding of wild game; digestion trials using moose rumen liquid to better understand the nutritional value of different foods to moose.

Primate ecology

My PhD work on spider monkeys in Bolivia (see Background) has led to continued involvement in the field of primate ecology and conservation. I am involved in a book project as a co-author. The book is called "Primate Diet and Nutrition: Needing, Finding and Using Food", edited by Joanna Lambert and Jessica Rothman. I have written review articles about primate nutritional ecology and associated scientific approaches. I am frequently reviewing articles and grant applications about primate ecology and nutrition, as well as tropical forest ecology at large.


I regularly provide lectures in three master-level courses given at the Southern Swedish Forest Research Centre.

Together with my colleague Martin Ahlström I have created and held a one-day course called "Forestry and game in balance", for employees in Swedish forestry industry; five course events Feb – Mar 2018 in southern Sweden (130 participants in total). Funded by Landsbygdsprogrammet (EU-funding).


Resources at our lab

  • Access to laboratory at SLU where we conduct a diverse array of analyses of nutritional composition of plant and other biological samples.
  • In-house access to near-infra red spectroscopy and expertise in related modelling.
  • In-house lab space for sample preparation and drying.
  • Access to laboratory at SLU and elsewhere where we conduct DNA extraction, PCR and DNA metabarcoding of biological samples.
  • At our department you find expertise in multiple disciplines: ecology, forestry, management, pathology and forest planning. I further collaborate closely with veterinarians and social scientists that are based at other departments at SLU or other universities.
  • An extensive network of hundreds of hunters in >50 moose management units (älgskötselområden) in southern Sweden, with experience in collecting samples for research.
  • Established channels for outreach of research results to stakeholders, which include forest owners, hunters, management groups and government agencies.


I finalized my PhD research in July 2008. That project was highly field intensive and involved reduced-impact logging in Bolivia, in relation to spider monkey (Ateles chamek) nutritional ecology and conservation. The candidacy was based at the Fenner School of Environment and Society, Australian National University, Canberra. The project, for which the field work took place in a certified forestry concession, assessed the importance of timber tree species to spider monkeys in terms of diet and ecology, and the capacity of RIL to maintain a suitable habitat for seed dispersers such as spider monkeys. During this period I developed a special interest in the analysis of complex multidimensional nutritional data, which I have great use of in my current work on moose in Sweden. After the thesis was finalized, I worked as a post-doc at the Fenner School for three months until the birth of my second child. This post-doc involved writing publications about sustainable resource use in managed forests.

I conducted my BSc in Ecology and Conservation Biology through Uppsala University, Sweden (1994-2000). My MSc project assessed orangutan population density, forest structure and fruit availability in hand-logged and unlogged peat swamp forests in West Kalimantan, Indonesia. I have also gained research experience participating in projects in different parts of the world, e.g: population biology of sea-turtles, Great Barrier Reef, Australia; behavioral ecology of barnacle geese on Gotland, Sweden; river otter population survey, Sweden; botanical research in Brunei Darussalaam; landscape ecology research on peccaries and jaguars in Madidi National Park, Bolivia. The latter project was organized by Wildlife Conservation Society, and during this time I co-discovered a species of titi-monkey previously unknown to science. I was also part of the WCS team that raised US$650 000 for Madidi National Park by auctioning the naming rights of this new species (Callicebus aureipalatii).


I co-supervise two PhD students (see Presentation) and supervise multiple MSc students at the Southern Swedish Forest Research Centre.

Selected publications

Peer-reviewed scientific publications in international journals

  1. Forbey, J.S. , R. Liu, T.T. Caughlin, M.D. Matocq, J.A. Vucetich, K.D. Kohl, M.D. Dearing, A.M. Felton. Using physiologically-based models to predict population responses to phytochemicals by wild vertebrate herbivores. Accepted in Animal on 02 August 2018.
  2. Malmsten, J. Dalin, A-M., Moutailler, S., Devillers, E., Gondard, M., Felton. A.M. Vector-borne zoonotic pathogens in Eurasian moose (Alces alces alces). Accepted in Vector-Borne and Zoonotic Diseases on 03 September 2018.
  3. Felton, A. M., H. K. Wam, C. Stolter, K. M. Mathisen, and M. Wallgren. 2018. The complexity of interacting nutritional drivers behind food selection, a review of northern cervids. Ecosphere 9:e02230.
  4. Tigabu, M., and A. M. Felton. 2018. Multivariate calibration of near infrared spectra for predicting nutrient concentrations of solid moose rumen contents. Silva Fennica 52:1. doi:10.14214/sf.7822
  5. Krizsan, S. J., A. Mateos-Rivera, S. Bertilsson, A. M. Felton, A. Anttila, M. Ramin, M. Vaga, H. Gidlund, and P. Huhtanen. 2018. An in vitro evaluation of browser and grazer fermentation efficiency and microbiota using European moose spring and summer foods. Accepted by Ecology and Evolution, 8:4183–4196. DOI:   10.1002/ece3.3  
  6. Wam, H. K., A. M. Felton, C. Stolter, L. Nybakken, and O. Hjeljord. 2018. Moose selecting for specific nutritional composition of birch places limits on food acceptability. Ecology and Evolution 8:1117-1130. DOI: 10.1002/ece3.3715.
  7. Hudson, L.N. et al. (multiple co-authors in alphabetical order). 2017. The database of the PREDICTS (Projecting Responses of Ecological Diversity In Changing Terrestrial Systems) project. Ecology and Evolution 7 (1): 145-188. DOI: 10.1002/ece3.2579
  8. Felton, A. M., A. Felton, L. Edenius, J. Cromsigt, J. Malmsten, and H. K. Wam. 2017 Interactions between ungulates, forests and supplementary feeding: the role of nutritional balancing in determining outcomes. Mammal Research 62(1):1-7. doi:10.1007/s13364-016-0301-1
  9. Felton A.M., Felton A, Raubenheimer D, Simpson SJ, Krizsan SJ, Hedwall P-O, et al. 2016. The Nutritional Balancing Act of a Large Herbivore: An Experiment with Captive Moose (Alces alces L). PLoS ONE 11(3): e0150870. doi:10.1371/journal.pone.0150870
  10. Felton, A., L. Gustafsson, J. M. Roberge, T. Ranius, J. Hjältén, J. Rudolphi, M. Lindbladh, J. Weslien, L. Rist, J. Brunet, and A. M. Felton. 2016. How climate change adaptation and mitigation strategies can threaten or enhance the biodiversity of production forests: Insights from Sweden. Biological Conservation 194:11-20. DOI 10.1016/j.biocon.2015.11.030
  11. Felton, A., P. O. Hedwall, M. Lindbladh, T. Nyberg, A. M. Felton, E. Holmström, I. Wallin, M. Löf, and J. Brunet. 2016. The biodiversity contribution of wood plantations: Contrasting the bird communities of Sweden’s protected and production oak forests. Forest Ecology and Management 365:51-60. DOI 10.1016/j.foreco.2016.01.030
  12. Felton A, Nilsson U, Sonesson J, Felton AM, Roberge J-M, Ranius T, et al. 2016. Replacing monocultures with mixed-species stands: Ecosystem service implications of two production forest alternatives in Sweden. Ambio;45(2):124-39. DOI 10.1007/s13280-015-0749-2
  13. Gustafsson, L., Felton, A., Felton, A. M., Brunet, J., Caruso, A., Hjältén, J., Lindbladh, M., Ranius, T., Roberge, J-M., Weslien, J-O.  2015. Natural versus national boundaries: The importance of considering biogeographical patterns in forest conservation policy. Conservation Letters. DOI:10.1111/conl.12087.
  14. Lindbladh, M., Hedwall, P-O., Wallin, I., Felton, A.M, Böhlenius, H. & Felton, A. 2014. Short-rotation bioenergy stands as an alternative to spruce monocultures: Implications for bird biodiversity. Silva Fennica 48 (5), DOI 10.14214/sf.1135.
  15. Hudson, L.N. et al. (multiple co-authors in alphabetical order). 2014. The PREDICTS database: a global database of how local terrestrial bioviersity responds to human impacts. Ecology and Evolution 4(24):4701-35. doi: 10.1002/ece3.1303.
  16. Raubenheimer, D., Machovsky-Capuska. G., Felton, A. M., and Simpson, S. J. 2014. Nutritional geometry: from insects to ruminants. Proc. Austr. Soc. Anim. Prod. 30:32-36.
  17. DeGabriel, J.L., Moore B.D., Felton, A.M., Ganzhorn, J.U., Stolter, C., Wallis, I.R., Johnson, C.N. & Foley, W.J. 2014. Translating nutritional ecology from the laboratory to the field: milestones in linking plant chemistry to population regulation in mammalian browsers. Oikos. 123: 298–308. DOI 10.1111/j.1600-0706.2013.00727.x
  18. Felton, A., Lindbladh, M., Elmberg, J., Felton, A.M., Andersson, E., Sekercioglu, C.H., Collingham, Y. & Huntley, B. 2014. Projecting impacts from anthropogenic climatic change on the bird communities of southern Sweden’s spruce monocultures: Will the species-poor get poorer? Ornis Fennica. 90:1-13.
  19. Felton, A.M, Felton, A., Rumiz, D.I., Villaroel, N., Chapman, C.A., Lindenmayer, D.B. 2013. Commercial harvesting of Ficus timber – an emerging threat to frugivorous wildlife and sustainable forestry. Biological Conservation 159:96-100. DOI 10.1016/j.biocon.2012.10.025
  20. Wallis, I.R., Edwards, M.J., Windley, H., Krockenberger, A.K., Felton, A.M., et al. 2012. Food for folivores – nutritional explanations linking diets to population density. Oecologia, 169:281-291. DOI 10.1007/s00442-011-2212-9
  21. Driscoll, D.A., Felton, A., Gibbons, P., Munro, N.T., Felton, A.M, Lindenmayer, D.B. 2011. Priorities in policy and management when existing biodiversity stressors interact with climate-change. Climatic Change, 111(3-4):533-557. DOI 10.1007/s10584-011-0170-1
  22. Felton, A.M, Felton, A., Foley, W.J., Lindenmayer, D.B. 2010. The role timber tree species play in the nutritional ecology of spider monkeys (Ateles chamek). Forest Ecology and Management 259(8):1642-1649. DOI: 10.1016/j.foreco.2010.01.042
  23. Felton, A., Fischer, J., Lindenmayer, D.B., Montague-Drake, R., Lowe, W.R., Saunders, D., Felton, A.M. et al. 2009. Climate change, conservation and management: An assessment of the peer-reviewed scientific journal literature. Biodiversity Conservation 18:2243-2253. DOI 10.1007/s10531-009-9652-0
  24. Felton, A.M., Felton, A., Wood, J.T., Foley, W.J., Raubenheimer, D., Wallis, I.R, Lindenmayer, D.B. 2009. Nutritional ecology of spider monkeys (Ateles chamek) in lowland Bolivia: How macro-nutrient balancing influences food choices. International Journal of Primatology 30: 675-696. DOI 10.1007/s10764-009-9367-9
  25. Felton, A.M., Felton, A., Raubenheimer, D., Simpson, S.J., Foley, W.J., Wood, J.T., Wallis, I.R., Lindenmayer, D.B. 2009. Protein content of diets dictates the daily energy intake of a free-ranging primate. Behavioural Ecology 20: 685-690. DOI 10.1093/beheco/arp021
  26. Felton, A.M. Felton, A., Lindenmayer, D.B., Foley, W.J. 2009. Nutritional goals of wild primates. Functional Ecology 23:70-78. DOI 10.1111/j.1365-2435.2008.01526.x
  27. Felton, A.M., Felton, A., Wood, J.T., Lindenmayer, D.B. 2008. Diet and feeding ecology of the Peruvian spider monkey (Ateles chamek) in a Bolivian forest: The importance of Ficus as a staple food resource. International Journal of Primatology 29:379-403. DOI 10.1007/s10764-008-9241-1
  28. Felton, A, Wood, J, Felton, A.M et al.  2008. Bird community responses to reduced-impact logging in a certified forestry concession in lowland Bolivia, Biological Conservation 141:545-555. DOI 10.1016/j.biocon.2007.11.009
  29. Felton, A, Wood, J, Felton, A.M et al. 2008. A comparison of bird communities in the anthropogenic and natural-tree fall gaps of a reduced-impact logged forest in Bolivia, Bird Conservation International 18:129-143.  DOI 10.1017/S0959270908000117
  30. Felton, A, Felton, A.M et al. 2008. The display of a reddish hermit (Phaethornis ruber) in a lowland rainforest, Bolivia. Wilson Journal of Ornithology 120(1):201-204. doi:
  31. Lindenmayer, D.B, Fischer, J, Felton, A, Montague, R, Manning, AD, Simberloff, D, Youngentob, K, Saunders, D, Wilson, D, Felton, A.M, et al. 2007. The complementarity of single-species and ecosystem-oriented research in conservation research, Oikos, 116(7):1220-1226. DOI 10.1111/j.2007.0030-1299.15683.x
  32. Felton, A, Felton, AM, Hennessey, B.A. & Lindenmayer, D.B. 2007. Birds surveyed in the harvested and unharvested areas of a reduced-impact logged forestry concession, located in the lowland forests of Bolivia, Check List 3(1):43-50. DOI:
  33. Felton, A, Felton, A.M, Woods, J. & Lindenmayer, D.B. 2006. Vegetation structure, phenology, and regeneration in the natural and anthropogenic tree-fall gaps of a reduced-impact logged Bolivian forest. Forest Ecology Management, 235: 186-193.  
  34. Felton, A, Felton, A.M, Wallace, R.B. & Gomez, H. 2006. Identification, behavioral observations, and notes on the distribution of the Titi monkeys Callicebus modestus and C. olallae. Prim. Cons., 20: 41-46. doi:
  35. Wallace, R.B, Gomez, H, Felton, A.M & Felton, A. 2006. On a new species of Titi monkey from Bolivia. Primate Conservation 20: 29-39.  doi:
  36. Felton, A, Alford, R.A, Felton, A.M & Schwarzkopf, L. 2006. Multiple mate choice criteria and the importance of age for male mating success in the microhylid frog, Cophixalus ornatus. Behavioral Ecology & Sociobiology 59(6): 786-795. DOI 10.1007/s00265-005-0124-6
  37. Felton, A.M, Engström, L.M., Felton, A. & Knott, C.D. 2003. Orangutan Pongo p. pygmaeus population density, forest structure and fruit availability in hand-logged and unlogged peat swamp forests in West Kalimantan, Indonesia. Biological Conservation 114(1): 91-101. doi:10.1016/S0006-3207(03)00013-2


Original contributions to refereed scientific conferences

  • Forbey, J.S. , R. Liu, T.T. Caughlin, M.D. Matocq, J.A. Vucetich, K.D. Kohl, M.D. Dearing, A.M. Felton. Using physiologically-based models to predict population responses to phytochemicals by wild vertebrate herbivores. Accepted in Animal on 02 August 2018.  Invited keynote lecture delivered ny Annika M. Felton and Jennifer S. Forbey at the International Symposium for the Nutrition of Herbivores (ISNH), Cleremont-Ferrand, 5th September 2018.
  • Krizsan, S. J., Felton, A.M., Ramin, M., Anttila, A., Vaga, M., Gidlund, H. and Huhtanen, P. 2013. A comparison of herbivore digestion efficiency in vitro using moose spring and summer foods. Proceedings of the 4th Nordic Feed Science Conference, Uppsala, Sweden, 12-13 June 2013. Eds Uden, P., Eriksson, T. Rustas, B. O. et al. pp 118-122. (project concept, experimental design, field & lab work, results interpretation, writing & editing).


Book chapters

  • Felton, A.M., Lambert, J. In process. The role of different nutrients in the food choice of primates. In: Primate diet and nutrition: needing, finding and using food. Eds. Lambert, J.E. and Rothman, J. University of Chicago Press.
  • Shimooka, Y., Campbell, C.J., Di Fiore, A., Felton, A.M, et al., 2008. Demography and group composition of Ateles. In: Spider monkeys: behavior, ecology and evolution of the genus Ateles. Ed. Campbell, C.J. Cambridge University Press. Pp: 329-348.

Master theses I have supervised

  • Widén, A. 2017. Effects of supplementary feeding on bark stripping by red deer (Cervus elaphus). Lunds Universitet.
  • Rautiainen, H. 2017. Nutritional ecology of roe deer (Capreolus capreolus L.) and fallow deer (Dama dama L.) – A case study of the browser – grazer dichotomy. SLU, Department of Ecology.
  • Karlström, J. 2016. Södras viltfoderhantering vid röjning – instruktioner och uppföljningsrutiner. Southern Swedish Forest Research Centre, SLU Alnarp.
  • Blomqvist, J. 2016 Supplementary feeding of ungulates in southern Sweden – relationships between supplementary feeding, browsing damages and land use. Southern Swedish Forest Research Centre, SLU Alnarp.
  • Westerström, S. 2015. Ungulate browsing pressure on Populus and feeding patterns in southern Sweden. Southern Swedish Forest Research Centre, SLU Alnarp.
  • Jönsson, A. 2015. Stand size effects on the proportion of damage by ungulates in poplar stands. Southern Swedish Forest Research Centre, SLU Alnarp.
  • Otto, P. 2013. Winter feeding site choice of ungulates in relation to food quality. Dept. Wildlife, Fish and Environmental Studies, SLU Umeå.



  • Felton, A.M. 2008. The Nutritional Ecology of Spider Monkeys (Ateles chamek) in the Context of Reduced- Impact Logging. PhD thesis, Fenner School of Environment and Society, Australian National University. (Open Access Digital Theses)