Register      Login
Wildlife Research Wildlife Research Society
Ecology, management and conservation in natural and modified habitats
RESEARCH ARTICLE

A management experiment reveals the difficulty of altering seedling growth and palatable plant biomass by culling invasive deer

David S. L. Ramsey A , David M. Forsyth B K , Clare J. Veltman C , Sarah J. Richardson D , Robert B. Allen D E , Will J. Allen F , Richard J. Barker G , Peter J. Bellingham D , Chris L. Jacobson H , Simon J. Nicol I , Alastair W. Robertson J and Charles R. Todd A
+ Author Affiliations
- Author Affiliations

A Arthur Rylah Institute for Environmental Research, Department of Environment, Land, Water and Planning, 123 Brown Street, Heidelberg, Vic. 3084, Australia.

B Vertebrate Pest Research Unit, New South Wales Department of Primary Industries, 1447 Forest Road, Orange, NSW 2800, Australia.

C Science and Policy, Department of Conservation, c/- Private Bag 11052, Palmerston North 4442, New Zealand.

D Landcare Research, PO Box 69040, Lincoln 7640, New Zealand.

E Current address: 8 Roblyn Place, Lincoln 7608, New Zealand.

F Learning for Sustainability, PO Box 30108, Christchurch 8244, New Zealand.

G Department of Mathematics and Statistics, University of Otago, PO Box 56, Dunedin 9054, New Zealand.

H Sustainability Research Centre, University of the Sunshine Coast, Locked Bag 4, Maroochydore DC, Qld 4551, Australia.

I Institute for Applied Ecology, University of Canberra, Bruce 2617, ACT, Australia.

J Institute of Agriculture and Environment, Massey University, Private Bag 11222, Palmerston North 4442, New Zealand.

K Corresponding author. Email: dave.forsyth@dpi.nsw.gov.au

Wildlife Research 44(8) 623-636 https://doi.org/10.1071/WR16206
Submitted: 7 November 2016  Accepted: 23 July 2017   Published: 26 February 2018

Abstract

Context: There is concern that deer are shifting forests towards undesirable trajectories, and culling of deer is often advocated to mitigate these impacts. However, culling deer is expensive and sometimes controversial. To reliably ascertain whether such action is beneficial, management-scale experiments are needed. We conducted a management experiment to evaluate the benefits of culling deer in four New Zealand forests.

Aims: Our experiment tested the predictions that culling deer should increase (1) canopy tree seedling height relative growth rate (SHRGR), and (2) the foliar biomass of understorey species palatable to deer (FBP).

Methods: Each forest was divided into two 3600-ha areas, with deer culling randomly assigned to one area. Deer abundances were indexed using faecal pellet counts, and forest variables were measured at the start and end of the 8-year experiment. Deer were already at low abundance in one forest and were not culled there. We used structural equation modelling (SEM) with Bayesian variable selection to update our a priori graphical forest–deer model with data from all four forests.

Key results: Deer abundances were significantly reduced in one forest but increased or did not change in the other two forests in which deer culling occurred. Culling deer did not increase seedling height relative growth rate (SHRGR) or the foliar biomass of understorey species palatable to deer (FBP) in the three areas subject to deer culling compared with the three areas not subject to deer culling. SEM revealed no significant relationships between local-scale deer abundance and either SHRGR or FBP. Rather, tree basal area and the foliar biomass of unpalatable understorey species were important predictors of FBP and SHRGR, respectively, in some forests.

Conclusions: Our study revealed that culling deer, as currently practiced by New Zealand land managers, did not generate the desired responses in New Zealand forests, possibly due to deer not being culled to sufficiently low densities and/or because forest dynamics and abiotic drivers determined plant growth more than deer.

Implications: Managers should consider actions other than ineffective deer culling (e.g. creating canopy gaps) to alter the dynamics of New Zealand forests. Alternatively, managers will need to substantially increase culling effort above what is currently practised for this activity to substantially reduce deer populations and thus potentially alter forest dynamics.

Additional keywords: adaptive management, biological invasions, foliar biomass, herbivory, New Zealand, overabundance, seedling growth rate, ungulate impacts.


References

Allen, R. B., Clinton, P. W., and Davis, M. R. (1997). Cation storage and availability along a Nothofagus forest development sequence in New Zealand. Canadian Journal of Forest Research 27, 323–330.
Cation storage and availability along a Nothofagus forest development sequence in New Zealand.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DyaK2sXjsFOisLo%3D&md5=4ae15f08696fbc6a9c6535ee199e74e7CAS |

Augustine, D. J., and McNaughton, S. J. (1998). Ungulate effects on the functional species composition of plant communities: herbivore selectivity and plant tolerance. The Journal of Wildlife Management 62, 1165–1183.
Ungulate effects on the functional species composition of plant communities: herbivore selectivity and plant tolerance.Crossref | GoogleScholarGoogle Scholar |

Batcheler, C. L., and Logan, P. C. (1963). Assessment of an animal-campaign in the Harper–Avoca catchment. New Zealand Forestry Research Notes 27, 1–27.

Bee, J. N., Tanentzap, A. J., Lee, W. G., Lavers, R. B., Mark, A. F., Mills, J. A., and Coomes, D. A. (2009). The benefits of being in a bad neighbourhood: plant community composition influences red deer foraging decisions. Oikos 118, 18–24.
The benefits of being in a bad neighbourhood: plant community composition influences red deer foraging decisions.Crossref | GoogleScholarGoogle Scholar |

Bellingham, P. J., and Richardson, S. J. (2006). Tree seedling growth and survival over 6 years across different microsites in a temperate rain forest. Canadian Journal of Forest Research 36, 910–918.
Tree seedling growth and survival over 6 years across different microsites in a temperate rain forest.Crossref | GoogleScholarGoogle Scholar |

Bellingham, P. J., Richardson, S. J., Mason, N. W., Veltman, C. J., Allen, R. B., Allen, W. J., Barker, R. J., Forsyth, D. M., Nicol, S. J., and Ramsey, D. S. L. (2016). Introduced deer at low densities do not inhibit the regeneration of a dominant tree. Forest Ecology and Management 364, 70–76.
Introduced deer at low densities do not inhibit the regeneration of a dominant tree.Crossref | GoogleScholarGoogle Scholar |

Brooks, S., and Gelman, A. (1998). General methods for monitoring convergence of iterative simulations. Journal of Computational and Graphical Statistics 7, 434–455.

Canham, C. D. (1988). An index for understory light levels in and around canopy gaps. Ecology 69, 1634–1638.
An index for understory light levels in and around canopy gaps.Crossref | GoogleScholarGoogle Scholar |

Cardinal, E., Martin, J.-L., Tremblay, J.-P., and Côté, S. D. (2012). An experimental study of how variation in deer density affects vegetation and songbird assemblages of recently harvested boreal forests. Canadian Journal of Zoology 90, 704–713.
An experimental study of how variation in deer density affects vegetation and songbird assemblages of recently harvested boreal forests.Crossref | GoogleScholarGoogle Scholar |

Carswell, F. E., Doherty, J. E., Allen, R. B., Brignall-Theyer, M. E., Richardson, S. J., and Wiser, S. K. (2012). Quantification of the effects of aboveground and belowground competition on growth of seedlings in a conifer–angiosperm forest. Forest Ecology and Management 269, 188–196.
Quantification of the effects of aboveground and belowground competition on growth of seedlings in a conifer–angiosperm forest.Crossref | GoogleScholarGoogle Scholar |

Caughley, G. (1983). ‘The Deer Wars. The Story of Deer in New Zealand.’ (Heinemann: Auckland.)

Clinton, P. W., Allen, R. B., and Davis, M. R. (2002). Nitrogen storage and availability during stand development in a New Zealand Nothofagus forest. Canadian Journal of Forest Research 32, 344–352.
Nitrogen storage and availability during stand development in a New Zealand Nothofagus forest.Crossref | GoogleScholarGoogle Scholar |

Coomes, D. A., and Allen, R. B. (2007). Effects of size, competition and altitude on tree growth. Journal of Ecology 95, 1084–1097.
Effects of size, competition and altitude on tree growth.Crossref | GoogleScholarGoogle Scholar |

Coomes, D. A., Allen, R. B., Forsyth, D. M., and Lee, W. G. (2003a). How reversible are the impacts of introduced deer in New Zealand forests? Conservation Biology 17, 450–459.
How reversible are the impacts of introduced deer in New Zealand forests?Crossref | GoogleScholarGoogle Scholar |

Coomes, D. A., Duncan, R. P., Allen, R. B., and Truscott, J. (2003b). Disturbances prevent stem size–density distributions in natural forests from following scaling relationships. Ecology Letters 6, 980–989.
Disturbances prevent stem size–density distributions in natural forests from following scaling relationships.Crossref | GoogleScholarGoogle Scholar |

Coomes, D. A., Allen, R. B., Bentley, W. A., Burrows, L. E., Canham, C. D., Fagan, L., Forsyth, D. M., Gaxiola-Alcantar, A., Parfitt, R. L., Ruscoe, W. A., Wardle, D. A., Wilson, D. J., and Wright, E. F. (2005). The hare, the tortoise and the crocodile: the ecology of angiosperm dominance, conifer persistence and fern filtering. Journal of Ecology 93, 918–935.
The hare, the tortoise and the crocodile: the ecology of angiosperm dominance, conifer persistence and fern filtering.Crossref | GoogleScholarGoogle Scholar |

Coomes, D. A., Bentley, W. A., Tanentzap, A. J., and Burrows, L. E. (2013). Soil drainage and phosphorus depletion contribute to retrogressive succession along a New Zealand chronosequence. Plant and Soil 367, 77–91.
Soil drainage and phosphorus depletion contribute to retrogressive succession along a New Zealand chronosequence.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC3sXktlKgurg%3D&md5=87c5537a8137164d82abe9b9b0deccfaCAS |

Côté, S. D., Rooney, T. P., Tremblay, J. P., Dussault, C., and Waller, D. M. (2004). Ecological impacts of deer overabundance. Annual Review of Ecology Evolution and Systematics 35, 113–147.
Ecological impacts of deer overabundance.Crossref | GoogleScholarGoogle Scholar |

Cowan, P. E. (2005). Brushtail possum. In ‘The Handbook of New Zealand Mammals’. 2nd edn. (Ed. C. M. King.) pp. 56–80. (Oxford University Press: Melbourne.)

Department of Conservation (2001). Department of Conservation’s policy statement on deer control. http://doc.govt.nz/Documents/about-doc/policies-and-plans/deer-control-policy.pdf [verified 14 October 2016].

Department of Conservation (2014). Department of Conservation Annual Report for the year ended 30 June 2014. (Department of Conservation: Wellington.)

Eggleston, J. E. (2002). Conflicting values regarding the management of deer and other wild ungulates in New Zealand: a case study in environmental ethics. M.Sc. thesis, Lincoln University, Lincoln, New Zealand.

Ferry, B., Morneau, F., Bontemps, J. D., Blanc, L., and Freycon, V. (2010). Higher treefall rates on slopes and waterlogged soils result in lower stand biomass and productivity in a tropical rain forest. Journal of Ecology 98, 106–116.
Higher treefall rates on slopes and waterlogged soils result in lower stand biomass and productivity in a tropical rain forest.Crossref | GoogleScholarGoogle Scholar |

Forsyth, D. M., Coomes, D. A., Nugent, G., and Hall, G. M. J. (2002). The diet and diet preferences of ungulates (order: Artiodactyla) in New Zealand. New Zealand Journal of Zoology 29, 323–343.
The diet and diet preferences of ungulates (order: Artiodactyla) in New Zealand.Crossref | GoogleScholarGoogle Scholar |

Forsyth, D. M., Barker, R. J., Morriss, G., and Scroggie, M. P. (2007). Modeling the relationship between fecal pellet indices and deer density. The Journal of Wildlife Management 71, 964–970.
Modeling the relationship between fecal pellet indices and deer density.Crossref | GoogleScholarGoogle Scholar |

Forsyth, D. M., Wilmshurst, J. M., Allen, R. B., and Coomes, D. A. (2010). Impacts of introduced deer and extinct moa on New Zealand ecosystems. New Zealand Journal of Ecology 34, 48–65.

Forsyth, D. M., Thomson, C., Hartley, L., MacKenzie, D. I., Price, R., Wright, E. F., Mortimer, J., Nugent, G., Wilson, L., and Livingstone, P. (2011). Long-term changes in the relative abundances of introduced deer in New Zealand estimated from faecal pellet frequencies. New Zealand Journal of Zoology 38, 237–249.
Long-term changes in the relative abundances of introduced deer in New Zealand estimated from faecal pellet frequencies.Crossref | GoogleScholarGoogle Scholar |

Forsyth, D. M., Ramsey, D. S. L., Veltman, C. J., Allen, R. B., Allen, W. J., Barker, R. J., Jacobson, C. L., Nicol, S. J., Richardson, S. J., and Todd, C. R. (2013). When deer must die: large uncertainty surrounds changes in deer abundance achieved by helicopter- and ground-based hunting in New Zealand forests. Wildlife Research 40, 447–458.
When deer must die: large uncertainty surrounds changes in deer abundance achieved by helicopter- and ground-based hunting in New Zealand forests.Crossref | GoogleScholarGoogle Scholar |

Forsyth, D. M., Wilson, D. J., Easdale, T., Kunstler, G., Canham, C. D., Ruscoe, W. A., Wright, E. F., Murphy, L., Gormley, A. M., Gaxiola, A., and Coomes, D. A. (2015). Century-scale effects of invasive deer and rodents on the dynamics of forests growing on soils of contrasting fertility. Ecological Monographs 85, 157–180.
Century-scale effects of invasive deer and rodents on the dynamics of forests growing on soils of contrasting fertility.Crossref | GoogleScholarGoogle Scholar |

Frazer, G. W., Canham, C. D., and Lertzman, K. P. (1999). Gap Light Analyzer (GLA): imaging software to extract canopy structure and gap light transmission indices from true-colour fisheye photographs. Users’ manual and program documentation, Version 2.0. (Simon Fraser University: Burnaby, British Columbia, and the Institute of Ecosystem Studies: Millbrook, NY.) http://www.rem.sfu.ca/forestry/downloads/Files/GLAV2UsersManual.pdf [verified 14 October 2016].

Frost, H. C., Storm, G. L., Batcheller, M. J., and Lovallo, M. J. (1997). White-tailed deer management at Gettysburg National Military Park and Eisenhower National Historic Site. Wildlife Society Bulletin 25, 462–469.

Grace, J. B., Anderson, T. M., Olff, H., and Scheiner, S. M. (2010). On the specification of structural equation models for ecological systems. Ecological Monographs 80, 67–87.
On the specification of structural equation models for ecological systems.Crossref | GoogleScholarGoogle Scholar |

Hairston, N. G. (1989). ‘Ecological Experiments: Purpose, Design and Execution.’ (Cambridge University Press: Cambridge.)

Harris, L. H. (2002). ‘The Deer Menace: a History of Government Pest Control Operations 1930–1987.’ (L. H. Harris: Wellington.)

Hobbs, N. T. (1996). Modification of ecosystems by ungulates. The Journal of Wildlife Management 60, 695–713.
Modification of ecosystems by ungulates.Crossref | GoogleScholarGoogle Scholar |

Holloway, J. T. (1950). Deer and the forests of western Southland. New Zealand Journal of Forestry 6, 123–137.

Hooten, M. B., and Hobbs, N. T. (2015). A guide to Bayesian model selection for ecologists. Ecological Monographs 85, 3–28.
A guide to Bayesian model selection for ecologists.Crossref | GoogleScholarGoogle Scholar |

Husheer, S. W., and Robertson, A. W. (2005). High-intensity deer culling increases growth of mountain beech seedlings in New Zealand. Wildlife Research 32, 273–280.
High-intensity deer culling increases growth of mountain beech seedlings in New Zealand.Crossref | GoogleScholarGoogle Scholar |

Husheer, S. W., Allen, R. B., and Robertson, A. W. (2006). Suppression of regeneration in New Zealand mountain beech forests is dependent on species of introduced deer. Biological Invasions 8, 823–834.
Suppression of regeneration in New Zealand mountain beech forests is dependent on species of introduced deer.Crossref | GoogleScholarGoogle Scholar |

Jacobson, C., Allen, W., Veltman, C., Ramsey, D. S. L., Forsyth, D. M., Nicol, S., Todd, C., and Barker, R. (2009). Collaborative learning as part of adaptive management of forests affected by deer. In ‘Adaptive Environmental Management – a Practitioner’s Guide’. (Eds C. Allan, and G. H. Stankey.) pp. 275–294. (Springer: Dordrecht, the Netherlands.)

Kaji, K., Saitoh, T., Uno, H., Matsuda, H., and Yamamura, K. (2010). Adaptive management of sika deer populations in Hokkaido, Japan: theory and practice. Population Ecology 52, 373–387.
Adaptive management of sika deer populations in Hokkaido, Japan: theory and practice.Crossref | GoogleScholarGoogle Scholar |

Killmaster, C. H., Osborn, D. A., Warren, R. J., and Miller, K. V. (2007). Deer and understory plant responses to a large-scale herd reduction on a Georgia State Park. Natural Areas Journal 27, 161–168.
Deer and understory plant responses to a large-scale herd reduction on a Georgia State Park.Crossref | GoogleScholarGoogle Scholar |

King, C. M. (2005). ‘The Handbook of New Zealand Mammals.’ 2nd edn. (Oxford University Press: Melbourne.)

Kosko, B. (1986). Fuzzy cognitive maps. International Journal of Man-Machine Studies 24, 65–75.
Fuzzy cognitive maps.Crossref | GoogleScholarGoogle Scholar |

Kosko, B. (1992). ‘Neural Networks and Fuzzy Systems: a Dynamical Systems Approach to Machine Intelligence.’ (Prentice-Hall Inc.: Upper Saddle River, NJ.)

Kuo, L., and Mallick, B. (1998). Variable selection for regression models. Sankhya Series B 60, 65–81.

Lusk, C. H., and Jorgensen, M. A. (2013). The whole-plant compensation point as a measure of juvenile tree light requirements. Functional Ecology 27, 1286–1294.
The whole-plant compensation point as a measure of juvenile tree light requirements.Crossref | GoogleScholarGoogle Scholar |

Lusk, C. H., Kaneko, T., Grierson, E., and Clearwater, M. (2013). Correlates of tree species sorting along a temperature gradient in New Zealand rain forests: seedling functional traits, growth and shade tolerance. Journal of Ecology 101, 1531–1541.
Correlates of tree species sorting along a temperature gradient in New Zealand rain forests: seedling functional traits, growth and shade tolerance.Crossref | GoogleScholarGoogle Scholar |

Mason, N. W. H., Peltzer, D. A., Richardson, S. J., Bellingham, P. J., and Allen, R. B. (2010). Stand development moderates effects of ungulate exclusion on foliar traits in the forests of New Zealand. Journal of Ecology 98, 1422–1433.
Stand development moderates effects of ungulate exclusion on foliar traits in the forests of New Zealand.Crossref | GoogleScholarGoogle Scholar |

McNab, W. H. (1993). A topographic index to quantify the effect of mesoscale landform on site productivity. Canadian Journal of Forest Research 23, 1100–1107.
A topographic index to quantify the effect of mesoscale landform on site productivity.Crossref | GoogleScholarGoogle Scholar |

McNulty, S. A., Porter, W. F., Mathews, N. E., and Hill, J. A. (1997). Localized management for reducing white-tailed deer populations. Wildlife Society Bulletin 25, 265–271.

Miller, B. F., Campbell, T. A., Laseter, B. R., Ford, W. M., and Miller, K. V. (2010). Test of localized management for reducing deer browsing in forest regeneration areas. The Journal of Wildlife Management 74, 370–378.
Test of localized management for reducing deer browsing in forest regeneration areas.Crossref | GoogleScholarGoogle Scholar |

Niwa, S., Mariani, L., Kaneko, N., Okada, H., and Sakamoto, K. (2011). Early-stage impacts of sika deer on structure and function of the soil microbial food webs in a temperate forest: a large-scale experiment. Forest Ecology and Management 261, 391–399.
Early-stage impacts of sika deer on structure and function of the soil microbial food webs in a temperate forest: a large-scale experiment.Crossref | GoogleScholarGoogle Scholar |

Ntzoufras, I. (2009). ‘Bayesian Modeling using WinBUGS.’ (Wiley: New York.)

Nugent, G., and Choquenot, D. (2004). Comparing cost-effectiveness of commercial harvesting, state-funded culling, and recreational deer hunting in New Zealand. Wildlife Society Bulletin 32, 481–492.
Comparing cost-effectiveness of commercial harvesting, state-funded culling, and recreational deer hunting in New Zealand.Crossref | GoogleScholarGoogle Scholar |

Nugent, G., Fraser, W., and Sweetapple, P. (2001). Top down or bottom up? Comparing the impacts of introduced arboreal possums and ‘terrestrial’ ruminants on native forests in New Zealand. Biological Conservation 99, 65–79.
Top down or bottom up? Comparing the impacts of introduced arboreal possums and ‘terrestrial’ ruminants on native forests in New Zealand.Crossref | GoogleScholarGoogle Scholar |

Nuttle, T., Ristau, T. E., and Royo, A. A. (2014). Long-term biological legacies of herbivore density in a landscape-scale experiment: forest understoreys reflect past deer density treatments for at least 20 years. Journal of Ecology 102, 221–228.
Long-term biological legacies of herbivore density in a landscape-scale experiment: forest understoreys reflect past deer density treatments for at least 20 years.Crossref | GoogleScholarGoogle Scholar |

O’Hara, R. B., and Sillanpää, M. J. (2009). A review of Bayesian variable selection methods: what, how and which. Bayesian Analysis 4, 85–117.
A review of Bayesian variable selection methods: what, how and which.Crossref | GoogleScholarGoogle Scholar |

Parkes, J., Robley, A., Forsyth, D. M., and Choquenot, D. (2006). Adaptive management experiments in vertebrate pest control in New Zealand and Australia. Wildlife Society Bulletin 34, 229–236.
Adaptive management experiments in vertebrate pest control in New Zealand and Australia.Crossref | GoogleScholarGoogle Scholar |

Peltzer, D. A., Allen, R. B., Bellingham, P. J., Richardson, S. J., Wright, E. F., Knightbridge, P. I., and Mason, N. W. H. (2014). Disentangling drivers of tree population size distributions. Forest Ecology and Management 331, 165–179.
Disentangling drivers of tree population size distributions.Crossref | GoogleScholarGoogle Scholar |

Platt, J. R. (1964). Strong inference: certain systematic methods of scientific thinking may produce much more rapid progress than others. Science 146, 347–353.
Strong inference: certain systematic methods of scientific thinking may produce much more rapid progress than others.Crossref | GoogleScholarGoogle Scholar | 1:STN:280:DC%2BC3czpvFCgug%3D%3D&md5=df07dc8b0d5e664877bdcd5d097ec0b1CAS |

Platt, K. H., Allen, R. B., Coomes, D. A., and Wiser, S. K. (2004). Mountain beech seedling responses to removal of below-ground competition and fertilizer addition. New Zealand Journal of Ecology 28, 289–293.

Plummer, M. (2003). JAGS: a program for analysis of Bayesian graphical models using Gibbs sampling. In ‘Proceedings of the 3rd International Workshop on Distributed Statistical Computing (DSC 2003)’. (Eds K. Hornik, F. Leisch and A. Zeileis.) pp. 1–10. (Austrian Association for Statistical Computing and the R Foundation for Statistical Computing: Vienna.) http://www.R-project.org/conferences/DSC-2003/Proceedings/ [verified 14 October 2016].

Porter, W. F., and Underwood, H. B. (1999). Of elephants and blind men: deer management in the U.S. national parks. Ecological Applications 9, 3–9.
Of elephants and blind men: deer management in the U.S. national parks.Crossref | GoogleScholarGoogle Scholar |

Porter, W. F., Mathews, N. E., Underwood, H. B., Sage, R. W., and Beherd, D. F. (1991). Social organization in deer: implications for localized management. Environmental Management 15, 809–814.
Social organization in deer: implications for localized management.Crossref | GoogleScholarGoogle Scholar |

Ramsey, D. S. L., Forsyth, D. M., Veltman, C. J., Nicol, S. J., Todd, C. R., Allen, R. B., Allen, W. J., Bellingham, P. J., Richardson, S. J., Jacobson, C. L., and Barker, R. J. (2012). Forest responses to deer control in a New Zealand adaptive management experiment: predictions from qualitative ecosystem models. Ecological Modelling 240, 93–104.
Forest responses to deer control in a New Zealand adaptive management experiment: predictions from qualitative ecosystem models.Crossref | GoogleScholarGoogle Scholar |

Richardson, S. J., Allen, R. B., and Doherty, J. E. (2008). Shifts in leaf N:P ratio during resorption reflect soil P in temperate rainforest. Functional Ecology 22, 738–745.
Shifts in leaf N:P ratio during resorption reflect soil P in temperate rainforest.Crossref | GoogleScholarGoogle Scholar |

Richardson, S. J., Bellingham, P. J., Allen, R. B., and Veltman, C. (2010). Initial vegetation conditions in study sites of the ‘Forests Affected by Deer’ project. Contract report prepared for Department of Conservation (LC0910/082). Landcare Research, Lincoln, New Zealand. Available at http://www.landcareresearch.co.nz/science/plants-animals-fungi/ecosystems/forest-ecosystems/consequences-of-multiple-herbivores [verified 4 November 2016].

Simard, M. A., Dussault, C., Huot, J., and Côté, S. D. (2013). Is hunting an effective tool to control overabundant deer? A test using an experimental approach. The Journal of Wildlife Management 77, 254–269.
Is hunting an effective tool to control overabundant deer? A test using an experimental approach.Crossref | GoogleScholarGoogle Scholar |

Stewart, G. H., and Burrows, L. E. (1989). The impact of white-tailed deer Odocoileus virginianus on regeneration in the coastal forests of Stewart Island, New Zealand. Biological Conservation 49, 275–293.
The impact of white-tailed deer Odocoileus virginianus on regeneration in the coastal forests of Stewart Island, New Zealand.Crossref | GoogleScholarGoogle Scholar |

Stewart, G. H., Wardle, J. A., and Burrows, L. E. (1987). Forest understorey changes after reduction in deer numbers, northern Fiordland, New Zealand. New Zealand Journal of Ecology 10, 35–42.

Tanentzap, A. J., Burrows, L. E., Lee, W. G., Nugent, G., Maxwell, J. M., and Coomes, D. A. (2009). Landscape-level vegetation recovery from herbivory: progress after four decades of invasive red deer control. Journal of Applied Ecology 46, 1064–1072.
Landscape-level vegetation recovery from herbivory: progress after four decades of invasive red deer control.Crossref | GoogleScholarGoogle Scholar |

Tanentzap, A. J., Bazely, D. R., Koh, S., Timciska, M., Haggith, E. G., Carleton, T. J., and Coomes, D. A. (2011). Seeing the forest for the deer: do reductions in deer-disturbance lead to forest recovery? Biological Conservation 144, 376–382.
Seeing the forest for the deer: do reductions in deer-disturbance lead to forest recovery?Crossref | GoogleScholarGoogle Scholar |

Tremblay, J.-P., Huot, J., and Potvin, F. (2006). Divergent nonlinear responses of the boreal forest field layer along an experimental gradient of deer densities. Oecologia 150, 78–88.
Divergent nonlinear responses of the boreal forest field layer along an experimental gradient of deer densities.Crossref | GoogleScholarGoogle Scholar |

Tremblay, J.-P., Huot, J., and Potvin, F. (2007). Density-related effects of deer browsing on the regeneration dynamics of boreal forests. Journal of Applied Ecology 44, 552–562.
Density-related effects of deer browsing on the regeneration dynamics of boreal forests.Crossref | GoogleScholarGoogle Scholar |

Walters, C. (1986). ‘Adaptive Management of Renewable Resources.’ (MacMillan: New York.)

Walters, C., and Holling, C. S. (1990). Large-scale management experiments and learning by doing. Ecology 71, 2060–2068.
Large-scale management experiments and learning by doing.Crossref | GoogleScholarGoogle Scholar |

Wardle, J. A. (1984). ‘The New Zealand Beeches: Ecology, Utilisation and Management.’ (New Zealand Forest Service: Christchurch.)

Wardle, D. A., Barker, G. M., Yeates, G. W., Bonner, K. I., and Ghani, A. (2001). Introduced browsing mammals in New Zealand natural forests: aboveground and belowground consequences. Ecological Monographs 71, 587–614.
Introduced browsing mammals in New Zealand natural forests: aboveground and belowground consequences.Crossref | GoogleScholarGoogle Scholar |

Williams, B. K. (2011). Adaptive management of natural resources – framework and issues. Journal of Environmental Management 92, 1346–1353.
Adaptive management of natural resources – framework and issues.Crossref | GoogleScholarGoogle Scholar |

Wiser, S. K., Hurst, J. M., Allen, R. B., and Wright, E. F. (2011). New Zealand’s forest and shrubland communities: a quantitative classification based on a nationally representative plot network. Applied Vegetation Science 14, 506–523.
New Zealand’s forest and shrubland communities: a quantitative classification based on a nationally representative plot network.Crossref | GoogleScholarGoogle Scholar |

Wright, D. M., Tanentzap, A. J., Flores, O., Husheer, S. W., Duncan, R. P., Wiser, S. K., and Coomes, D. A. (2012). Impacts of culling and exclusion of browsers on vegetation recovery across New Zealand forests. Biological Conservation 153, 64–71.
Impacts of culling and exclusion of browsers on vegetation recovery across New Zealand forests.Crossref | GoogleScholarGoogle Scholar |