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*Note: Some older HEE publications below have not yet been restructured into ADA compliant formats. We are actively working on this.


HEE Data Reports
  1. HEE Species List 2006-2015 (PDF: 203 KB)
  2. The Hardwood Ecosystem Experiment: Threatened and Endangered Species   (PDF: 335 KB)

Peer-Reviewed Extension Publications
  1. The Hardwood Ecosystem Experiment: 2006-2016. 2019. (PDF: 2533 KB)
  2. The Great Clearcut Controversy . 2017. (PDF: 3198 KB)
  3. The Hardwood Ecosystem Experiment: Indiana Forestry and Wildlife. 2015. (PDF: 4.9 MB)
  4. Breeding Birds and Forest Management: the Hardwood Ecosystem Experiment and the Central Hardwoods Region. 2015. (PDF: 3.2 MB)
  5. Forest Management for Reptiles and Amphibians. 2013. (PDF: 2 MB)

Published Proceedings and Symposia
  1. Swihart, R.K., M.R. Saunders, R.A. Kalb, and C.H. Michler, eds. 2013. The Hardwood Ecosystem Experiment: a framework for studying responses to forest management. Gen. Tech. Rep. NRS-P-108. Newton Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research Station. [CD ROM] 350 p.
  2. Sustainable Forestry White Paper (PDF: 17 KB)
  3. Abstracts: Managing wildlife for sustainable forests (PDF: 1.43 MB)

Peer-Reviewed Research Articles
  1. Connare, B. and K. Islam. 2023. Advancing Our Understanding of Cerulean Warbler Space Use Through Radio Telemetry. Journal of Fish and Wildlife Management; doi: https://doi.org/10.3996/JFWM-21-100
  2. Beilke, E.A., Haulton, G.S., and J.M. O’Keefe. 2023. Foliage-roosting eastern red bats select for features associated with management in a central hardwood forest, Forest Ecology, and Management, 527(1), https://doi.org/10.1016/j.foreco.2022.120604
  3. Beilke, E.A. and J.M. O’Keefe. 2022. Bats reduce insect density and defoliation in temperate forests: An exclusion experiment. Ecology. https://doi.org/10.1002/ecy.3903
  4. Divoll, T.J., Brown, V.A., McCracken, G.F., and J.M. O’Keefe. 2022. Prey size is more representative than prey taxa when measuring dietary overlap in sympatric forest bats. Environmental DNA, https://doi.org/10.1002/edn3.354.
  5. Ochs, A. E., Saunders, M. R., & R.K. Swihart. 2022. Response of terrestrial salamanders to the decade following timber harvest in hardwood forests. Forest Ecology and Management511, 120159.
  6. Connare, B. and K. Islam. 2022. Failure to advance migratory phenology in response to climate change may pose a significant threat to a declining Nearctic-Neotropical songbird. International Journal of Biometeorology, https://doi.org/10.1007/s00484-022-02239-9.
  7. Sharp, A. R., Connare, B. M., and Islam, K. 2021. Novel nest tree selection by a deciduous forest New World warbler. The Wilson Journal of Ornithology 133(4):640-645. doi.org/10.1676/20-0016
  8. Divoll, T.J., Aldrich, S.P., Haulton, G.S., and J.M. O’Keefe. 2022. Endangered Myotis bats forage in regeneration openings in a management forest. Forest Ecology and Management, 503 (1), Endangered Myotis bats forage in regeneration openings in a managed forest – ScienceDirect
  9. Milne, M.A., Gonsiorowski, J., Tuft, N., Deno, B., Ploss, T., Acosta, J., Frandsen, L. and C. Venable. 2021. Effects of fire on ground-dwelling spider (Araneae) assemblages in Central Indiana Forests. Environmental Entomology, 50(4); 781-789. https://doi.org/10.1093/ee/nvab051
  10. Sharp, A. and K. Islam. 2021. Does the intrusion of an avian nest predator elicit a change in the behaviour of a canopy-nesting passerine? Behaviour. 1-26. 10.1163/1568539X-bja10078
  11. Bergeson, S.M., Confortin, K.A., Carter, T.C., Karsk, J.R., Haulton, S., Burnett, H. 2021. Northern long-eared bats roosting in a managed forest in south-central Indiana, Forest Ecology and Management, Volume 483,118928, https://doi.org/10.1016/j.foreco.2021.118928.
  12. Connare, B., Delancey, C., Sharp, A., MacDonald, G., and Islam, K. 2020. “Low Breeding-Site Fidelity Suggests Dispersal of Cerulean Warblers,” Northeastern Naturalist 27(4), 669-680. https://doi.org/10.1656/045.027.0408
  13. Delancey, C. and K. Islam. 2020. Analysis of microhabitat characteristics at roost sites of Cerulean Warblers. PLOS One. https://doi.org/10.1371/journal.pone.0241501
  14. Ruhl, P.J., Kellner, K. F., and J.B. Dunning. 2020. Ecological factors explain habitat associations of mature-forest songbirds in regenerating forest clearcuts. The Wilson Journal of Ornithology doi: https://doi.org/10.1676/19-49
  15. Delancey, C., Islam, K., Kramer, G., MacDonald, G., Sharp, A., and B. Connare. 2020. Geolocators reveal migration routes, stopover sites, and nonbreeding dispersion in a population of Cerulean Warblers, Animal Migration7(1):19-26. doi: https://doi.org/10.1515/ami-2020-0003
  16. Ruhl, P.J., Flaherty, E. and J.B. Dunning. 2020. Using stable isotopes of plasma, red blood cells, feces, and feathers to assess mature-forest bird diet during the postfledging period. Canadian Journal of Zoology, 98: 39-46. https://doi.org/10.1139/cjz-2019-0109.
  17. Martin, M.K., Sheets, J.J., Sparks, D., and J.O. Whitaker. 2020. Diets of bats before and after forest management. Proceedings of the Indiana Academy of Science 129(1):56-64.
  18. Milne, M.A., Louderman, J., Foster, B., Grijalva, I., Lewis, J.J., Bishop, L., Deno, B.L., Acosta, J., Frandsen, L., and E. Stern. 2019. More spiders in Indiana: 100 new and updated distribution records. Proceedings of the Indiana Academy of Science 128(1):87-105.
  19. MacDonald, G.J. and K. Islam. 2019. Do social factors explain seasonal variation in dawn song characteristics of paired male Cerulean Warblers (Setophaga cerulea)? Bioacoustics, https://doi.org/10.1080/09524622.2019.1682671
  20. Bergeson, S.M., Holmes, J.B. and J.M. O’Keefe. 2019. Indiana bat roosting behavior differs between urban and rural landscapes, Urban Ecosystems, https://doi.org/10.1007/s11252-019-00903-4
  21. Delancey, C.D., MacDonald, G.J., and K. Islam. 2019. First confirmed hybrid pairing between a Cerulean Warbler (Setophaga cerulea) and a Black-throated Blue Warbler (Setophaga caerulescens). Wilson Journal of Ornithology, 131(1):161-165.
  22. Delancey, C.D. and Islam K. 2019. Post-fledging habitat use in a declining songbird. PeerJ 7:e7358 https://doi.org/10.7717/peerj.7358
  23. Nelson, D.L., Kellner, K.F., and Swihart, R.K. 2019. Rodent population density and survival respond to disturbance induced by timber harvest, Journal of Mammalogy 100(4):1253-1262.
  24. MacDonald, G.J., Delancey, C. D. and K. Islam. 2019. Novel vocalizations, including song, from 2 female Cerulean Warblers (Setophaga cerulea). The Wilson Journal of Ornithology 131(2): 366-373. https://doi.org/10.1676/18-104
  25. Caldwell, K.L., Carter, T.C. and J.C. Doll. 2019. A comparison of bat activity in a managed central hardwood forest. The American Midland Naturalist 181(2):225-244.
  26. Hanna, L., Kissick, A. L., McCroskey, E., and J.D. Holland. 2019. Resilience to disturbance is a cross‐scale phenomenon offering a solution to the disturbance paradox. Ecosphere 10( 4):e02682. 10.1002/ecs2.2682
  27. Stanis, S. and M.R. Saunders. 2018. Long-term overstory tree quality monitoring through multiple prescribed fires in eastern deciduous forests. In: Kirschman, Julia E., comp. 2018. Proceedings of the 19th biennial southern silvicultural research conference. e-Gen. Tech. Rep. SRS-234. Asheville, NC: U.S. Department of Agriculture Forest Service, Southern Research Station:355-362.
  28. Kellner, K.F., Ruhl, P.J., Dunning, J.B., Barnes, K.W., Saunders, M.R., and R.K. Swihart. 2018. Local-scale habitat components driving bird abundance in eastern deciduous forests. The American Midland Naturalist, 180(1):52-65.
  29. Ruhl, P.J., Delancey, C.D., and J.B. Dunning Jr. 2018. Roost preference, postfledging habitat use, and breeding phenology of adult female Worm-eating Warblers (Helmitheros vermivorum) on the breeding grounds. The Wilson Journal of Ornithology., 130(2): 397-409.
  30. Delancey, C., MacDonald, G., & Islam, K. 2018. American Redstarts ( Setophaga ruticilla ) usurp Cerulean Warbler ( Setophaga cerulea ) nest in Southern Indiana. The Wilson Journal of Ornithology., 130(2), 554-558.
  31. Ruhl, P., Kellner, K., Pierce, J., Riegel, J., Swihart, R., Saunders, M., & Dunning, Jr, J. 2018. Characterization of Worm-eating Warbler (Helmitheros vermivorum) breeding habitat at the landscape level and nest scale. Avian Conservation and Ecology, 13(1):11.
  32. Bergeson, S.M., O’Keefe, J.M., and Haulton, G.S. 2018. Managed forests provide roosting opportunities for Indiana bats in south-central Indiana. Forest Ecology and Management, 427:305-316.
  33. Divoll, T.J., Brown, V.A., Kinne, J., McCracken, G.F., and J.M. O’Keefe. 2018. Disparities in second-generation DNA metabarcoding results exposed with accessible and repeatable workflows. Molecular Ecology Resources, 18(3):590-601. https://doi.org/10.1111/1755-0998.12770
  34. Hale, V., B. MacGowan, L. Corriveau, D. Huse, A.F. Currylow, and S. Thompson. 2017. Radio transmitter implantation and movement in the wild Timber Rattlesnake (Crotalus horridus). Journal of Wildlife Diseases, https://doi.org/10.7589/2016-08-186
  35. Kellner, K.F. and R.K. Swihart. 2017. Herbivory on planted oak seedlings across a habitat edge created by timber harvest. Plant Ecology, 218:213-223.
  36. Kellner, K.F. and R.K. Swihart. 2017. Simulation of oak early life history and interactions with disturbance via an individual-based model, SOEL. PLoS ONE 12(6): e0179643. https://doi.org/10.1371/journal.pone.0179643
  37. MacGowan, B.J., A.F.T.Currylow and J.E.MacNeil. 2017. Short-term responses of Timber Rattlesnakes (Crotalus horridus) to even-aged timber harvests in Indiana. Forest Ecology and Management, 376: 30-36.
  38. Murray, B.D., J.D. Holand, K.S. Summerville, J.B. Dunning, M.R. Saunders, and M.A. Jenkins. 2017. Functional Diversity response to hardwood forest management varies across taxa and spatial scales. Ecological Applications 1002/eap.1532
  39. Nemes, C.E. and K. Islam. 2017. Breeding Season Microhabitat use by Cerulean Warbler (Setophaga cerulea) in an experimentally-managed forest. Forest Ecology and Management http://dx.doi.org/10.1016/j.foreco.2016.11.008
  40. Olson, M.G. and M.R. Saunders. 2017. Long-term research on managed hardwood forests in eastern North America. Forest Ecology and Management, 387: 1-2.
  41. Summerville, K.S. and R.J. Marquis. 2017. Comparing the responses of larval and adult lepidopteran communities to timber harvest using long-term, landscape-scale studies in oak-hickory forests. Forest Ecology and Management http://dx.doi.org/10.1016/j.foreco.2016.08.050
  42. Auer, S.A., K. Islam, J.R. Wagner, K.S. Summerville, and K.W. Barnes. 2016. The diet of Cerulean Warbler (Setophaga cerulea) nestlings and adult nest provisioning behaviors in S. Indiana. The Wilson Journal of Ornithology, 128(3):573-583.
  43. Barnes, K.W., K.Islam and S.A.Auer. 2016. Integrating LIDAR-derived canopy structure into cerulean warbler habitat models. Journal of Wildlife Management, 80(1): 101-116.
  44. Kellner, K.F. and R.K.Swihart. 2016. Timber harvest and drought interact to impact oak seedling growth and survival in the Central Hardwood Forest. Ecosphere 7(10): e01473
  45. Kellner, K.F., N.I.Lichti and R.K.Swihart. 2016. Midstory removal reduces effectiveness of oak (Quercus) acorn dispersal by small mammals in the Central Hardwood Forest region. Forest Ecology and Management, 375:182-190.
  46. Kellner, K.F., P.J.Ruhl, J.B.Dunning, J.K.Riegel and R.K.Swihart. 2016. Multi-scale responses of breeding birds to experimental forest management in Indiana. Forest Ecology and Management, 382:64-75.
  47. Milne, M.A., B. Foster, J.J. Lewis, L. Bishop, A. Hoffman, T. Ploss, and B. Deno. 2016. Spiders in Indiana: seventy-one new and updated distribution records. Proceedings of the Indiana Academy of Science, 125(1): 75-85.
  48. Murray, B.D., C.R.Webster, M.A.Jenkins, M.R.Saunders and G.S.Haulton. 2016. Ungulate impacts on herbaceous-layer plant communities in even-aged and uneven-aged forests. Ecosphere, 7(6): e01378
  49. Leonard, O.D., J.W.Moore, J.K.Riegel, A.R.Meier, J.B.Dunning and K.F.Kellner, R.K.Swihart. 2015. Effect of variation in forest harvest intensity on winter occupancy of barred owls and eastern screech-owls in deciduous forests of the east-central United States. Journal of Field Ornithology, 86(2): 115-129.
  50. Nemes, C.E., K.Islam and D.M.Pirtle. 2015. First documentation of a “double-decker” Cerulean Warbler (Setophaga cerulea) nest. The Wilson Journal of Ornithology, 127(3):534-538.
  51. Olson, Z.H., B.J.MacGowan, M.T.Hamilton, A.F.T.Currylow and R.N.Williams. 2015. Survival of timber rattlesnakes: Investigating individual, environmental, and ecological effects. Herpetologica, 71:274-279.
  52. Powell, G., R.M.Brattain and J.D.Holland. 2015. New distributional records for the genus Pytho. The Coleopterists’ Bulletin, 66:761-765.
  53. Ruhl, P.J., Jeffrey K. Riegel, and John B. Dunning. 2015. Confirmation of successful chestnut-sided warbler breeding in south-central Indiana. Proceedings of the Indiana Academy of Science, 124:38-42.
  54. Ruhl, P.J. and J.B.Dunning. 2015. Morphometrics of Worm-eating Warblers in south-central Indiana: hatching year and after hatching year comparison. North American Bird Bander, 43:81-84.
  55. Summerville, K.S. 2015. Species traits fail to consistently predict moth persistence in managed forest stands. Great Lakes Entomologist, 48:114-124.
  56. Wagner, J.R., K. Islam, and K. Summerville. 2015. Cerulean Warbler territory size is influenced by prey-rich tree abundance. Proceedings of the Indiana Academy of Science,124:70-79.
  57. Currylow, A.F., A.J.Johnson and R.N.Williams. 2014. Evidence of ranavirus infections among sympatric larval amphibians and box turtles. Journal of Herpetology, 48(1): 117-121.
  58. Kellner, K.F. and R.K.Swihart. 2014. Changes in small mammal microhabitat use following silvicultural disturbance. American Midland Naturalist, 172: 349-359.
  59. Kellner, K.F., J.R. Riegel and R.K. Swihart. 2014. Effects of silvicultural disturbance on acorn infestation and removal in the central hardwood forest region. New Forests, 45(2): 265-281.
  60. Kimble, S.J.A., O.E. Rhodes, Jr. and R.N. Williams. 2014. Relatedness and other finescale genetic processes in the box turtle. Conservation Genetics, 15(4): 967-979.
  61. Kimble, S.J.A., O.E.Rhodes, Jr. and R.N.Williams. 2014. Unexpectedly low rangewide population genetic structure of the imperiled eastern box turtles. . PLoS ONE 9:e92274
  62. LaGrange, S.M., S.J.A.Kimble, B.J.MacGowan and R.N.Williams. 2014. Seasonal variance in hematology and plasma chemistry values of the timber rattlesnake (Crotalus horridus). Jounal of Wildlife Diseases DOI 10.7589/2013-10-267.
  63. MacNeil, J.E. and R.N.Williams. 2014. Effects of timber harvests and silvicultural edges on terrestrial salamanders. PLoS ONE, 9(12):e114683
  64. Owen, D.A.S., E.T.Carter, M.L.Holding, K.Islam and I.T.Moore. 2014. Roads are associated with a blunted stress response in a North American pit viper. General and Comparative Endocrinology, 202: 87-92.
  65. Summerville, K.S. 2014. Do seasonal temperatures, species traits and nearby timber harvest predict variation in moth species richness and abundance in unlogged deciduous forests? Agricultural and Forest Entomology, 16: 80-86.
  66. Wagner, J.R. and K.Islam. 2014. Nest-site selection and breeding ecology of the Cerulean Warbler in southern Indiana. Northeastern Naturalist, 21: 515-528.
  67. Auer, S.A., K.Islam, K.W.Barnes and J.A.Brown. 2013. Documentation of Red-bellied Woodpecker predation of a Cerulean Warbler nestling. The Wilson Journal of Ornithology, 125(3) 642-646.
  68. Currylow, A.F., M.S.Tift, J.L.Meyer, D.E.Crocker and R.N.Williams. 2013. Seasonal variations in plasma vitellogenin and sex steroids in male and female eastern box turtles (Terrapene c. carolina). General and Comparative Endocrinology, 180:48-55.
  69. Kaminski, K.J. and K. Islam. 2013. Effects of forest treatments on abundance and spatial characteristics of cerulean warbler territories. American Midland Naturalist, 170:111 – 120.
  70. Kellner, K.F., N.A.Urban and R.K.Swihart. 2013. Short-term responses of small mammals to timber harvest in the Central Hardwoods. Journal of Wildlife Management, 77(8): 1650-1663.
  71. MacNeil, J.E. and R.N.Williams. 2013. Effectiveness of two artificial cover objects in sampling terrestrial salamanders. Herpetological Conservation and Biology, 8(3): 552-560.
  72. Summerville, K.S. 2013. Forest lepidopteran communities are more resilient to shelterwood harvest compared to more intensive logging regimes. Ecological Applications, 23(5): 1101-1112.
  73. Summerville, K.S., J.L. Lane and D. Courard-Hauri. 2013. Stability in forest lepidopteran communities: how sensitive are pest species to experimental forest management. Insect Conservation and Diversity, 6:(3)265-275.
  74. Currylow, A.F., B.J.MacGowan and R.N.Williams. 2012. Hibernal thermal ecology of eastern box turtles within a managed forest landscape. Journal of Wildlife Management, 77(2): 326-335.
  75. Currylow, A.F., B.J.MacGowan and R.N.Williams. 2012. Short-term forest management effects on a long-lived ecotherm. PLoS ONE, 7(7):e40473.
  76. Islam, K., J.Wagner, R.Dibala, M.M.MacNeil, K.J.Kaminski and L.P.Young. 2012. Cerulean Warbler (Setophaga cerulea) response to changes in forest structure in Indiana. Ornitologia Neotropical, 23: 335-341
  77. Kimble, S.J.A. and R.N.Williams. 2012. Temporal variance in hematologic and plasma bi0chemical reference intervals for free-ranging eastern box turtles (Terrapene carolina carolina). Journal of Wildlife Diseases, 48(3):799-802.
  78. Urban, N.A., R.K.Swihart, M.C.Malloy and J.B.Dunning. 2012. Improving selection of indicator species when detection is imperfect. Ecological Indicators, 15: 188-197.
  79. Currylow, A.F., P.A.Zollner, B.J.MacGowan and R.N.Williams. 2011. A survival estimate of Midwestern adult Eastern box turtles using radio telemetry. The American Midland Naturalist, 165(1): 143-149.
  80. Summerville, K.S. 2011. Managing the forest for more than the trees: effects of experimental timber harvest on forest Lepidoptera. Ecological Applications, 21(3): 806-816.
  81. Urban, N.A. and R.K.Swihart. 2011. Small mammal responses to forest management for oak regeneration in southern Indiana. Forest Ecology and Management, 261:353-361.
  82. Holland, J.D. 2010. Isolating spatial effects on beta diversity to inform forest landscape planning. Landscape Ecology, 25(9): 1349-1362.
  83. Summerville, K.S., D.Courard-Hauri and M.M.Dupont. 2009. The legacy of timber harvest: Do patterns of species dominance suggest recovery of Lepidopteran communities in managed hardwood stands? Forest Ecology and Management, 259(1): 8-13.
  84. Summerville, K.S., M.M.Dupont, A.V.Johnson and R.L.Krehbiel. 2008. Spatial structure of forest Lepidopteran communities in oak hickory forests of Indiana. Environmental Entomology, 37(5): 1224-1230.