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

Detection and stratification approaches for aerial surveys of deer in prairie–parklands

Thomas J. Habib A C , David A. Moore B and Evelyn H. Merrill A
+ Author Affiliations
- Author Affiliations

A Department of Biological Sciences, University of Alberta, Edmonton, AB T6G 2E9, Canada.

B Alberta Fish and Wildlife Division, 8-4701 52 Street, Vermilion, AB T9X 1J9, Canada.

C Corresponding author. Email: thabib@ualberta.ca

Wildlife Research 39(7) 593-602 https://doi.org/10.1071/WR11175
Submitted: 16 October 2011  Accepted: 24 July 2012   Published: 7 September 2012

Abstract

Context: For management decisions that require accurate and precise estimates of large mammal population numbers, aerial surveys are considered reliable despite their cost. However, aerial surveys may still suffer from underestimation because of undetected animals and low precision as a result of inefficient sampling designs.

Aims: We assess detection of two species of deer in prairie-parkland communities of western Canada and evaluate a suite of survey design features for improving the accuracy and precision of population estimates from aerial surveys.

Methods: Modelling detection of deer was based on 100 sightability trials involving 54 radio-collared white-tailed and 46 mule deer. We used empirical survey data to simulate surveys under three alternative stratification approaches, schemes for grouping strata, and allocations of survey effort and compared the precision and accuracy of the resulting population estimates.

Key results: We observed deer in 83 of the 100 trials, with detection decreasing with small group size, reduced deer activity, low snow cover, high forest cover and observer fatigue. Survey precision and accuracy were highest when stratification was based on natural breaks, calculated via Jenks optimisation, in the values of resource-selection function (RSF), although improvement was less pronounced for estimates of mule deer abundance. Optimal or equal allocation of sampling effort among strata outperformed proportional allocation of sampling effort. Use of RSF for stratification reduced the coefficient of variation (CV) in estimates of deer numbers from 38% to 23% for white-tailed deer and from 33% to 27% for mule deer compared with past approaches.

Conclusions: Stratification based on RSF values improved precision of deer surveys the most; however, using even simple measures related to habitat selection can improve population estimates. Where deer are highly aggregated, reliably recording all variables needed to implement sightability models can prove problematic; however, survey detection adjustments are nevertheless important to account for the relatively small, but still significant, proportion of missed animals in open prairie–parkland environments.

Implications: Field experiments to assess aerial survey design are impractical because of cost. We illustrate how simulated surveys using empirical data can be useful to evaluate alternative survey designs to improve population estimates in a region when high accuracy or precision are needed to address management questions or to calibrate more cost-effective approaches.


References

Agriculture and Agri-Food Canada (2009). ‘Circa 2000 Land Cover for Agricultural Regions of Canada.’ (Government of Canada: Ottawa.) Available at http://ftp://ftp.agr.gc.ca/pub/ outgoing/nlwis-lc/LCV_CA_AAFC_30M_2000_V12/Mosaic [verified 1 May 2009].

Allen, J. R. (2005). Use of sightability models and resource selection functions to enhance aerial population surveys of elk (Cervus elaphus) in Alberta. M.Sc. Thesis, University of Alberta, Edmonton, Canada.

Allen, J. R., McInenly, L. E., Merrill, E. H., and Boyce, M. S. (2008). Using resource selection functions to improve estimation of elk population numbers. The Journal of Wildlife Management 72, 1798–1804.
Using resource selection functions to improve estimation of elk population numbers.Crossref | GoogleScholarGoogle Scholar |

Anderson, C. R., and Lindzey, F. G. (1996). Moose sightability model developed from helicopter surveys. Wildlife Society Bulletin 24, 247–259.

Anderson, C. R., Moody, D. S., Smith, B. L., Lindzey, F. G., and Lanka, R. P. (1998). Development and evaluation of sightability models for summer elk surveys. The Journal of Wildlife Management 62, 1055–1066.
Development and evaluation of sightability models for summer elk surveys.Crossref | GoogleScholarGoogle Scholar |

Barker, R. (2008). Theory and application of mark–recapture and related techniques to aerial surveys of wildlife. Wildlife Research 35, 268–274.
Theory and application of mark–recapture and related techniques to aerial surveys of wildlife.Crossref | GoogleScholarGoogle Scholar |

Bartmann, R. M., White, G. C., Carpenter, L. H., and Garrott, R. A. (1987). Aerial mark–recapture estimates of confined mule deer in pinyon–juniper woodland. Journal of Wildlife Management 51, 41–46.

Bird, R. D. (1961). ‘Ecology of the Aspen Parkland of Western Canada in Relation to Land Use.’ (Canada Department of Agriculture: Ottawa, Canada.)

Boyce, M. S., Vernier, P. R., Nielsen, S. E., and Schmiegelow, F. K. A. (2002). Evaluating resource selection functions. Ecological Modelling 157, 281–300.
Evaluating resource selection functions.Crossref | GoogleScholarGoogle Scholar |

Brown, S., Bart, J., Lanctot, R. B., Johnson, J. A., Kendall, S., Payer, D., and Johnson, J. (2007). Shorebird abundance and distribution on the coastal plain of the Arctic National Wildlife Refuge. The Condor 109, 1–14.
Shorebird abundance and distribution on the coastal plain of the Arctic National Wildlife Refuge.Crossref | GoogleScholarGoogle Scholar |

Burnham, K. P., and Anderson, D. R. (2002). ‘Model Selection and Inference: a Practical Information-Theoretic Approach.’ 3rd edn. (Springer Verlag: New York.)

Cairns, S. C., Lollback, G. W., and Payne, N. (2008). Design of aerial surveys for population estimation and the management of macropods in the Northern Tablelands of New South Wales, Australia. Wildlife Research 35, 331–339.
Design of aerial surveys for population estimation and the management of macropods in the Northern Tablelands of New South Wales, Australia.Crossref | GoogleScholarGoogle Scholar |

Caughley, G., Sinclair, R., and Scott-Kemmis, D. (1976). Experiments in aerial survey. The Journal of Wildlife Management 40, 290–300.
Experiments in aerial survey.Crossref | GoogleScholarGoogle Scholar |

Christman, M. C. (2000). A review of quadrat-based sampling of rare, geographically clustered populations. Journal of Agricultural Biological & Environmental Statistics 5, 168–201.
A review of quadrat-based sampling of rare, geographically clustered populations.Crossref | GoogleScholarGoogle Scholar |

Clover, M. R. (1954). A portable deer trap and catch-net. California Fish and Game 40, 367–373.

Cogan, R. D., and Diefenbach, D. R. (1998). Effect of undercounting and model selection on a sightability-adjustment estimator for elk. Journal of Wildlife Management 62, 269–279.

Diefenbach, D. R., Marshall, M. R., Mattice, J. A., and Brauning, D. W. (2007). Incorporating availability for detection in estimates of bird abundance. The Auk 124, 96–106.
Incorporating availability for detection in estimates of bird abundance.Crossref | GoogleScholarGoogle Scholar |

Dorr, B. S., Burger, L. W., and Barras, S. C. (2008). Evaluation of aerial cluster sampling of double-crested cormorants on aquaculture ponds in Mississippi. The Journal of Wildlife Management 72, 1634–1640.

Dressel, S. C., and Norcross, B. L. (2005). Using poststratification to improve abundance estimates from multispecies surveys: a study of juvenile flatfishes. Fishery Bulletin 103, 469–488.

Environment Canada (2009). ‘Canadian Climate Weather Normals, 1971–2000.’ (Government of Canada: Ottawa.) Available at http://www.climate.weatheroffice.ec.gc.ca/climate_normals/index_e.html [verified12 October 2009].

Fielding, A. H., and Bell, J. F. (1997). A review of methods for the assessment of prediction errors in conservation presence/absence models. Environmental Conservation 24, 38–49.
A review of methods for the assessment of prediction errors in conservation presence/absence models.Crossref | GoogleScholarGoogle Scholar |

Freddy, D. J., White, G. C., Kneeland, M. C., Kahn, R. H., Unsworth, J. W., deVergie, W. J., Graham, V. K., Ellenberger, J. R., and Wagner, C. H. (2004). How many mule deer are there? Challenges of credibility in Colorado. Wildlife Society Bulletin 32, 916–927.
How many mule deer are there? Challenges of credibility in Colorado.Crossref | GoogleScholarGoogle Scholar |

Gasaway, W. C., DuBois, S. D., Reed, D. J., and Harbo, S. J. (1986). ‘Estimating Moose Population Parameters from Aerial Surveys.’ (Institute of Arctic Biology, University of Alaska: Fairbanks, AK.)

Gillies, C. S., Hebblewhite, M., Nielsen, S. E., Krawchuk, M. A., Aldridge, C. L., Frair, J. L., Saher, D. J., Stevens, C. E., and Jerde, C. L. (2006). Application of random effects to the study of resource selection by animals. Journal of Animal Ecology 75, 887–898.
Application of random effects to the study of resource selection by animals.Crossref | GoogleScholarGoogle Scholar |

Glasgow, W. M. (2000). White area ungulate management plan, 1997–98 to 1999–2000 project completion report. Alberta Fish and Wildlife, Edmonton, Canada.

Guschanski, K., Vigilant, L., McNeilage, A., Gray, M., Kagoda, E., and Robbins, M. M. (2009). Counting elusive animals: comparing field and genetic census of the entire mountain gorilla population of Bwindi Impenetrable National Park, Uganda. Biological Conservation 142, 290–300.
Counting elusive animals: comparing field and genetic census of the entire mountain gorilla population of Bwindi Impenetrable National Park, Uganda.Crossref | GoogleScholarGoogle Scholar |

Habib, T. J. (2010). Ecology and management of white-tailed deer (Odocoileus virginianus) and mule deer (O. hemionus) of East-Central Alberta in relation to chronic wasting disease. M.Sc Thesis, University of Alberta, Edmonton, Canada.

Habib, T. J., Merrill, E. H., Pybus, M. J., and Coltman, D. W. (2011). Modelling landscape effects on density–contact rate relationships of deer in eastern Alberta: implications for chronic wasting disease. Ecological Modelling 222, 2722–2732.
Modelling landscape effects on density–contact rate relationships of deer in eastern Alberta: implications for chronic wasting disease.Crossref | GoogleScholarGoogle Scholar |

Harbitz, A., Aschan, M., and Sunnana, K. (1998). Optimal effort allocation in stratified, large area trawl surveys, with application to shrimp surveys in the Barents Sea. Fisheries Research 37, 107–113.
Optimal effort allocation in stratified, large area trawl surveys, with application to shrimp surveys in the Barents Sea.Crossref | GoogleScholarGoogle Scholar |

Hata, D., and Berkson, J. (2004). Factors affecting horseshoe crab Limulus polyphemus trawl survey design. Transactions of the American Fisheries Society 133, 292–299.
Factors affecting horseshoe crab Limulus polyphemus trawl survey design.Crossref | GoogleScholarGoogle Scholar |

Highfield, L. D., Ward, M. P., Laffan, S. W., Norby, B., and Wagner, G. (2009). The impact of seasonal variability in wildlife populations on the predicted spread of foot and mouth disease. Veterinary Research 40, 18.
The impact of seasonal variability in wildlife populations on the predicted spread of foot and mouth disease.Crossref | GoogleScholarGoogle Scholar |

Hone, J. (2008). On bias, precision and accuracy in wildlife aerial surveys. Wildlife Research 35, 253–257.
On bias, precision and accuracy in wildlife aerial surveys.Crossref | GoogleScholarGoogle Scholar |

Jachmann, H. (2002). Comparison of aerial counts with ground counts for large African herbivores. Journal of Applied Ecology 39, 841–852.
Comparison of aerial counts with ground counts for large African herbivores.Crossref | GoogleScholarGoogle Scholar |

Jenks, G. F. (1967). The data model concept in statistical mapping. International Yearbook of Cartography 7, 186–190.

Johnson, C. J., Nielsen, S. E., Merrill, E. H., McDonald, T. L., and Boyce, M. S. (2006). Resource selection functions based on use-availability data: theoretical motivation and evaluation methods. The Journal of Wildlife Management 70, 347–357.
Resource selection functions based on use-availability data: theoretical motivation and evaluation methods.Crossref | GoogleScholarGoogle Scholar |

Khaemba, W. M., Stein, A., Rasch, D., De Leeuw, J., and Georgiadis, N. (2001). Empirically simulated study to compare and validate sampling methods used in aerial surveys of wildlife populations. African Journal of Ecology 39, 374–382.
Empirically simulated study to compare and validate sampling methods used in aerial surveys of wildlife populations.Crossref | GoogleScholarGoogle Scholar |

Lehtonen, R., and Pahkinen, E. (2004). ‘Practical Methods for Design and Analysis of Complex Surveys.’ (John Wiley and Sons: New York.)

Manning, J. A., and Garton, E. O. (2012). A sightability model for correcting visibility and availability biases in standardized surveys of breeding burrowing owls in southwest agroecosystem environments. The Journal of Wildlife Management 76, 65–74.
A sightability model for correcting visibility and availability biases in standardized surveys of breeding burrowing owls in southwest agroecosystem environments.Crossref | GoogleScholarGoogle Scholar |

Månsson, J., Hauser, C. E., Andrén, H., and Possingham, H. P. (2011). Survey method choice for wildlife management: the case of moose Alces alces in Sweden. Wildlife Biology 17, 176–190.
Survey method choice for wildlife management: the case of moose Alces alces in Sweden.Crossref | GoogleScholarGoogle Scholar |

McIntosh, T. E., Rosatte, R. C., Hamr, J., and Murray, D. L. (2009). Development of a sightability model for low-density elk populations in Ontario, Canada. The Journal of Wildlife Management 73, 580–585.
Development of a sightability model for low-density elk populations in Ontario, Canada.Crossref | GoogleScholarGoogle Scholar |

McPherson, J. M., Jetz, W., and Rogers, D. J. (2004). The effects of species’ range sizes on the accuracy of distribution models: ecological phenomenon or statistical artefact? Journal of Applied Ecology 41, 811–823.
The effects of species’ range sizes on the accuracy of distribution models: ecological phenomenon or statistical artefact?Crossref | GoogleScholarGoogle Scholar |

Mier, K. L., and Picquelle, S. J. (2008). Estimating abundance of spatially aggregated populations: comparing adaptive sampling with survey designs. Canadian Journal of Fisheries and Aquatic Sciences 65, 176–197.
Estimating abundance of spatially aggregated populations: comparing adaptive sampling with survey designs.Crossref | GoogleScholarGoogle Scholar |

Moore, D. (2003). ‘WMU 234 Ungulate Survey: January 18–20, 2003.’ (Alberta Fish and Wildlife: Vermilion, Canada.)

Nakagawa, S., and Cuthill, I. C. (2007). Effect size, confidence interval and statistical significance: a practical guide for biologists. Biological Reviews of the Cambridge Philosophical Society 82, 591–605.
Effect size, confidence interval and statistical significance: a practical guide for biologists.Crossref | GoogleScholarGoogle Scholar |

Pearse, A. T., Dinsmore, S. J., Kaminski, R. M., and Reinecke, K. J. (2008). Evaluation of an aerial survey to estimate abundance of wintering ducks in Mississippi. The Journal of Wildlife Management 72, 1413–1419.
Evaluation of an aerial survey to estimate abundance of wintering ducks in Mississippi.Crossref | GoogleScholarGoogle Scholar |

Pearse, A. T., Reinecke, K. J., Dinsmore, S. J., and Kaminski, R. M. (2009). Using simulation to improve wildlife surveys: wintering mallards in Mississippi, USA. Wildlife Research 36, 279–288.
Using simulation to improve wildlife surveys: wintering mallards in Mississippi, USA.Crossref | GoogleScholarGoogle Scholar |

Pettorelli, N., Cote, S. D., Gingras, A., Potvin, F., and Huot, J. (2007). Aerial surveys vs hunting statistics to monitor deer density: the example of Anticosti Island, Quebec, Canada. Wildlife Biology 13, 321–327.
Aerial surveys vs hunting statistics to monitor deer density: the example of Anticosti Island, Quebec, Canada.Crossref | GoogleScholarGoogle Scholar |

Pollock, K. H., and Kendall, W. L. (1987). Visibility bias in aerial surveys – a review of estimation procedures. The Journal of Wildlife Management 51, 502–510.
Visibility bias in aerial surveys – a review of estimation procedures.Crossref | GoogleScholarGoogle Scholar |

Pople, A. R., Cairns, S. C., Menke, N., and Payne, N. (2006). Estimating the abundance of eastern grey kangaroos (Macropus giganteus) in south-eastern New South Wales, Australia. Wildlife Research 33, 93–102.
Estimating the abundance of eastern grey kangaroos (Macropus giganteus) in south-eastern New South Wales, Australia.Crossref | GoogleScholarGoogle Scholar |

Potvin, F., and Breton, L. (2005). From the field: testing 2 aerial survey techniques on deer in fenced enclosures – visual double-counts and thermal infrared sensing. Wildlife Society Bulletin 33, 317–325.
From the field: testing 2 aerial survey techniques on deer in fenced enclosures – visual double-counts and thermal infrared sensing.Crossref | GoogleScholarGoogle Scholar |

Prenzlow, D. M., and Lovvorn, J. R. (1997). Design and results of a waterfowl breeding population survey for Wyoming. The Journal of Wildlife Management 61, 758–767.
Design and results of a waterfowl breeding population survey for Wyoming.Crossref | GoogleScholarGoogle Scholar |

Ransom, J. I. (2012). Detection probability in aerial surveys of feral horses. The Journal of Wildlife Management 76, 299–307.
Detection probability in aerial surveys of feral horses.Crossref | GoogleScholarGoogle Scholar |

Rice, C. G., Jenkins, K. J., and Chang, W. Y. (2009). A sightability model for mountain goats. The Journal of Wildlife Management 73, 468–478.
A sightability model for mountain goats.Crossref | GoogleScholarGoogle Scholar |

Riley, S. J., DeGloria, S. D., and Elliot, R. (1999). A terrain ruggedness index that quantifies topographic heterogeneity. Intermountain Journal of Sciences 5, 23–27.

Samuel, M. D., Garton, E. O., Schlegel, M. W., and Carson, R. G. (1987). Visibility bias during aerial surveys of elk in northcentral Idaho. The Journal of Wildlife Management 51, 622–630.
Visibility bias during aerial surveys of elk in northcentral Idaho.Crossref | GoogleScholarGoogle Scholar |

Sinclair, A. R. E., Fryxell, J. M., and Caughley, G. (2006). ‘Wildlife Ecology, Conservation, and Management.’ 2nd edn. (Blackwell Publishing: Malden, MA.)

Skalski, J. R., Millspaugh, J. J., and Spencer, R. D. (2005). Population estimation and biases in paintball, mark–resight surveys of elk. Journal of Wildlife Management 69, 1043–1052.

Smith, D. R., Conroy, M. J., and Brakhage, D. H. (1995). Efficiency of adaptive cluster sampling for estimating density of wintering waterfowl. Biometrics 51, 777–788.
Efficiency of adaptive cluster sampling for estimating density of wintering waterfowl.Crossref | GoogleScholarGoogle Scholar |

Smith, D. R., Brown, J. A., and Lo, N. C. H. (2004). Applications of adaptive sampling to biological populations. In ‘Sampling for Rare or Elusive Species: Concepts, Designs, and Techniques for Estimating Population Parameters’. (Ed. W. L. Thompson.) pp. 77–122. (Island Press: Washington, DC.)

Smith, D. R., Gray, B. R., Newton, T. J., and Nichols, D. (2010). Effect of imperfect detectability on adaptive and conventional sampling: simulated sampling of freshwater mussels in the upper Mississippi River. Environmental Monitoring and Assessment 170, 499–507.
Effect of imperfect detectability on adaptive and conventional sampling: simulated sampling of freshwater mussels in the upper Mississippi River.Crossref | GoogleScholarGoogle Scholar |

Steinhorst, R. K., and Samuel, M. D. (1989). Sightability adjustment methods for aerial surveys of wildlife populations. Biometrics 45, 415–425.
Sightability adjustment methods for aerial surveys of wildlife populations.Crossref | GoogleScholarGoogle Scholar |

Swets, J. A. (1988). Measuring the accuracy of diagnostic systems. Science 240, 1285–1293.
Measuring the accuracy of diagnostic systems.Crossref | GoogleScholarGoogle Scholar | 1:STN:280:DyaL1c3jsF2jtQ%3D%3D&md5=2e6d591a373c6cf9cf3ad77a2d974629CAS |

Thompson, S. K. (1990). Adaptive cluster sampling. Journal of the American Statistical Association 85, 1050–1059.
Adaptive cluster sampling.Crossref | GoogleScholarGoogle Scholar |

Thompson, S. K. (2002). ‘Sampling.’ 2nd edn. (John Wiley and Sons: New York.)

Thomson, J. A., Cooper, A. B., Burkholder, D. A., Heithaus, M. R., and Dill, L. M. (2012). Heterogeneous patterns of availability for detection during visual surveys: spatiotemporal variation in sea turtle dive-surfacing behaviour on a feeding ground. Methods in Ecology and Evolution 3, 378–387.
Heterogeneous patterns of availability for detection during visual surveys: spatiotemporal variation in sea turtle dive-surfacing behaviour on a feeding ground.Crossref | GoogleScholarGoogle Scholar |

Udevitz, M. S., Shults, B. S., Adams, L. G., and Kleckner, C. (2006). Evaluation of aerial survey methods for Dall’s sheep. Wildlife Society Bulletin 34, 732–740.

Unsworth, J. W., Kuck, L., and Garton, E. O. (1990). Elk sightability model validation at the National-Bison-Range, Montana. Wildlife Society Bulletin 18, 113–115.

Unsworth, J. W., Leban, F. A., Garton, E. O., Leptich, D. J., and Zager, P. (1999). ‘Aerial Survey: User’s Manual.’ Electronic edn. (Idaho Department of Fish and Game: Boise, ID.)

Wilson, G. J., and Delahay, R. J. (2001). A review of methods to estimate the abundance of terrestrial carnivores using field signs and observation. Wildlife Research 28, 151–164.
A review of methods to estimate the abundance of terrestrial carnivores using field signs and observation.Crossref | GoogleScholarGoogle Scholar |

Wittmer, H. U., McLellan, B. N., Seip, D. R., Young, J. A., Kinley, T. A., Watts, G. S., and Hamilton, D. (2005). Population dynamics of the endangered mountain ecotype of woodland caribou (Rangifer tarandus caribou) in British Columbia, Canada. Canadian Journal of Zoology 83, 407–418.

Zar, J. H. (1999). ‘Biostatistical Analysis.’ 4th edn. (Prentice Hall: Upper Saddle River, NJ.)