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The management of native regrowth forests for biodiversity has become increasingly important in recent years, as most old growth forest on mainland Australia is now protected and logging operations occur primarily in regrowth. Regrowth forest typically has higher stem density than old growth; therefore, bats (Microchiroptera) adapted to foraging along edges and in open spaces are likely to be less active in regrowth forest. Thinning is an integral component of regrowth management and could reduce structural clutter to a level suitable for bats with a range of clutter tolerances; yet little is known about the effects of thinning on bat activity, especially in Australia. In a preliminary study, we compared the activity levels of all bats, one open-space and three edge-space aerial foraging bat guilds in unthinned (n = 3) and thinned (n = 3) regrowth, in forests managed for timber production on the south coast of New South Wales. We measured bat activity at three heights in the forest (understorey, subcanopy and canopy). The number of volant insects (prey items) was also measured at each site/height combination, while 20 structural vegetation variables were measured at each site. A total of 263 bat passes was recorded in 18 detector-nights. We found high variability in bat activity for all bats and guilds between regrowth treatments and among heights, and no significant difference between levels of these factors. There was no significant interaction between logging and height factors. Vegetation structure varied significantly between unthinned and thinned sites for just two variables: the shrub layer had a higher percentage cover, and the vertical gap between canopy and understorey trees was halved, in thinned regrowth. Multivariate analysis of vegetation structure suggested just a small increase in the distance between canopy and understorey tree stems and slightly less cover in the canopy in thinned sites, changes that would represent a reduction in clutter. With the highest density of stems, the understorey of both regrowth types represented the most cluttered stratum for bats and overall bat activity was lower there (though not significantly). However, total bat activity did not indicate a clear response to insect abundance in the less cluttered subcanopy and canopy. Although the variability in bat activity within our regrowth treatments was too high to unequivocally state that thinning had no effect, results obtained from older regrowth of the same forest type suggest that overall activity is low for bats in dense regrowth forest, and that the level of thinning conducted at our sites was not sufficient to increase levels of activity consistently. We review the scientific literature on thinning effects for bats and formulate more specific hypotheses for future testing. Appropriate measures of clutter need to be identified to incorporate into indices of stand structural complexity that are relevant for bats.

Adam, M.D., Lacki, M.J. and Shoemaker, L.G. 1994. Influence of environmental conditions on flight activity of Plecotus townsendii virginianus (Chiroptera: Vespertilionidae). Brimleyana 21: 77-85.
Adams, M., Reardon, T.R., Baverstock, P.R. and Watts, C.H.S. 1988. Electrophoretic resolution of species boundaries in Australian Microchiroptera. IV. The Molossidae (Chiroptera). Australian Journal of Biological Science 41: 315-326.
Adams, M.D., Law, B.S. and French, K.O. 2005. Effect of lights on activity levels of forest bats: increasing the efficiency of surveys and species identification. Wildlife Research 32: 173-182.
Adams, M.D., Law, B.S. and French, K.O. 2009. Vegetation structure influences the vertical stratification of open- and edge-space aerial foraging bats in harvested forests. Forest Ecology and Management 258: 2090-2100.
Aldridge, H.D.J.N. and Rautenbach, I.L. 1987. Morphology, echolocation and resource partitioning in insectivorous bats. Journal of Animal Ecology 56: 763-778.
Anthony, E.L.P., Stack, M.H. and Kunz, T.H. 1981. Night roosting and the nocturnal time budget of the little brown bat, Myotis lucifugus: effects of reproductive status, prey density, and environmental conditions. Oecologia 51: 151-156.
Bailey, J.D. and Tappeiner, J.C. 1998. Effects of thinning on structural development in 40- to 100-year-old Douglas-fir stands in western Oregon. Forest Ecology and Management 108: 99-113.
Bennett, F., Lumsden, L.F. and Nicholls, A.O. 1995. Tree hollows as a resource for wildlife in remnant woodlands: spatial and temporal patterns across the northern plains of Victoria, Australia. Pacific Conservation Biology 1: 222-235.
Bowden, J. 1982. An analysis of factors affecting catches of insects in light-traps. Bulletin of Entomological Research 72: 535-556.
Brigham, R.M., Francis, R.L. and Hamdorf, S. 1997a. Microhabitat use by two species of Nyctophilus bats: a test of ecomorphology theory. Australian Journal of Zoology 45: 553-560.
Brigham, R.M., Grindal, S.D., Firman, M.C. and Morissette, J.L. 1997b. The influence of structural clutter on activity patterns of insectivorous bats. Canadian Journal of Zoology 75: 131-136.
Brower, J.E., Zar, J.H. and von Ende, C.N. 1990 Field and Laboratory Methods for General Ecology 3rd Edition, W.C. Brown Publishers, Dubuque, IA.
Brown, G.W., Nelson, J.L. and Cherry, K.A. 1997. The influence of habitat structure on insectivorous bat activity in montane ash forests of the Central Highlands, Victoria. Australian Forestry 60: 138-146.
Bullen, R. and McKenzie, N.L. 2001. Bat airframe design: flight performance, stability and control in relation to foraging ecology. Australian Journal of Zoology 49: 235-261.
Crampton, L.H. and Barclay, R.M.R. 1998. Selection of roosting and foraging habitat by bats in different-aged aspen mixedwood stands. Conservation Biology 12: 1347-1358.
Erickson, J.L. and West, S.D. 2003. Associations of bats with local structure and landscape features of forested stands in western Oregon and Washington. Biological Conservation 109: 95-102.
Fenton, M.B. 1990. The foraging behaviour and ecology of animal-eating bats. Canadian Journal of Zoology 68: 411-422.
Fenton, M.B., Boyle, N.G.H., Harrison, T.M. and Oxley, D.J. 1977. Activity patterns, habitat use, and prey selection by some African insectivorous bats. Biotropica 9: 73-85.
Fenton, M.B., Cumming, D.H.M., Rautenbach, I.L., Cumming, G.S., Cumming, M.S., Ford, G., Taylor, R.D., Dunlop, J., Hovorka, M.D., Johnston, D.S., Portfors, C.V., Kalcounis, M.C. and Mahlanga, Z. 1998. Bats and the loss of tree canopy in African woodlands. Conservation Biology 12: 399-407.
Fenton, M.B. and Morris, G.K. 1976. Opportunistic feeding by desert bats ( Myotis spp.). Canadian Journal of Zoology 54: 526-530.
Florence, R.G. 1996 Ecology and Silviculture of Eucalypt Forests. CSIRO Publishing, Collingwood.
Forestry Commission of NSW. 1983. Management Plan for Batemans Bay Management Area.
Forests NSW. 2008. State Forests of New South Wales, South Coast Region, silvicultural plan - spacing. 4 September 2008.
Fullard, J.H., Koehler, C., Surlykke, A. and McKenzie, N.L. 1991. Echolocation ecology and flight morphology of insectivorous bats (Chiroptera) in south-western Australia. Australian Journal of Zoology 39: 427-438.
Ghose, K. and Moss, C.F. 2003. The sonar beam pattern of a flying bat as it tracks tethered insects. Journal of the Acoustical Society of America 114: 1120-1131.
Gibbons, P., Lindenmayer, D.B., Barry, S.C. and Tanton, M.T. 2000. Hollow formation in eucalypts from temperate forests in southeastern Australia. Pacific Conservation Biology 6: 218-228.
Glass P.H. 1993. The response of insectivorous bat activity to disturbance from logging and habitat differences in the Kioloa State Forest. MSc thesis, University of Wollongong.
Grindal, S.D. and Brigham, R.M. 1998. Short-term effects of small-scale habitat disturbance on activity by insectivorous bats. Journal of Wildlife Management 62: 996-1003.
Grindal, S.D., Morissette, J.L. and Brigham, R.M.R. 1999. Concentration of bat activity in riparian habitats over an elevational gradient. Canadian Journal of Zoology 77: 972-977.
Hayes, J.P. 1997. Temporal variation in activity of bats and the design of echolocation-monitoring studies. Journal of Mammalogy 78: 514-524.
Hecker, K.R. and Brigham, R.M. 1999. Does moonlight change vertical stratification of activity by forest-dwelling insectivorous bats? Journal of Mammalogy 80: 1196-1201.
Hickey, M.B.C. and Fenton, M.B. 1996. Behavioural and thermoregulatory responses of female hoary bats, Lasiurus cinereus (Chiroptera: Vespertilionidae), to variations in prey availability. Ecoscience 3: 414-422.
Humes, M.L., Hayes, J.P. and Collopy, M.W. 1999. Bat activity in thinned, unthinned, and old-growth forests in western Oregon. Journal of Wildlife Management 63: 553-561.
JANIS. 1997. Nationally Agreed Criteria for the Establishment of a Comprehensive, Adequate and Representative Reserve System for Forests in Australia. A report by the Joint ANZECC/MCFFA National Forest Policy Statement Implementation Subcommittee.
Kalcounis-Ruppell, M.C., Psyllakis, J.M., Brigham, R.M. 2005. Tree roost selection by bats: an empirical synthesis using meta-analysis. Wildlife Society Bulletin 33: 1123-1132.
Kalko, E.K.V. and Schnitzler, H-U. 1993. Plasticity in echolocation signals of European pipistrelle bats in search flight: implications for habitat use and prey detection. Behavioral Ecology and Sociobiology 33: 415-428.
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Krusic, R.A., Yamasaki, M., Neefus, C.D. and Pekins, P.J. 1996. Bat habitat use in White Mountain National Forest. Journal of Wildlife Management 60: 625-631.
Kutt, A.S. 1995. Activity and stratification of microchiropteran bat communities in thinned, unthinned and old lowland regrowth forest, East Gippsland. The Victorian Naturalist 112: 86-92.
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Law, B.S. 2004. Challenges for managing bats in the State Forests of NSW. In Conservation of Australia's Forest Fauna 2nd Edition (Ed. Lunney, D.) pp. 748-760, Royal Zoological Society of New South Wales, Mosman.
Law, B.S. and Chidel, M. 2001. Bat activity 22 years after first-round intensive logging of alternate coupes near Eden, New South Wales. Australia Forestry 64: 242-247.
Law, B. and Chidel, M. 2002. Tracks and riparian zones facilitate the use of Australian regrowth forest by insectivorous bats. Journal of Applied Ecology 39: 605-617.
Law, B.S. and Chidel, M. 2004. Roosting and foraging ecology of the golden-tipped bat ( Kerivoula papuensis) on the south coast of New South Wales. Wildlife Research 31: 73-82.
Lee, Y.F. and McCracken, G.F. 2005. Dietary variation of Brazilian free-tailed bats links to migratory populations of pest insects. Journal of Mammalogy 86: 67-76.
Lloyd, A., Law, B. and Goldingay, R. 2006. Bat activity on riparian zones and upper slopes in Australian timber production forests and the effectiveness of riparian buffers. Biological Conservation 129: 207-220.
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References

Adam, M.D., Lacki, M.J. and Shoemaker, L.G. 1994. Influence of environmental conditions on flight activity of Plecotus townsendii virginianus (Chiroptera: Vespertilionidae). Brimleyana 21: 77-85.
Adams, M., Reardon, T.R., Baverstock, P.R. and Watts, C.H.S. 1988. Electrophoretic resolution of species boundaries in Australian Microchiroptera. IV. The Molossidae (Chiroptera). Australian Journal of Biological Science 41: 315-326.
Adams, M.D., Law, B.S. and French, K.O. 2005. Effect of lights on activity levels of forest bats: increasing the efficiency of surveys and species identification. Wildlife Research 32: 173-182.
Adams, M.D., Law, B.S. and French, K.O. 2009. Vegetation structure influences the vertical stratification of open- and edge-space aerial foraging bats in harvested forests. Forest Ecology and Management 258: 2090-2100.
Aldridge, H.D.J.N. and Rautenbach, I.L. 1987. Morphology, echolocation and resource partitioning in insectivorous bats. Journal of Animal Ecology 56: 763-778.
Anthony, E.L.P., Stack, M.H. and Kunz, T.H. 1981. Night roosting and the nocturnal time budget of the little brown bat, Myotis lucifugus: effects of reproductive status, prey density, and environmental conditions. Oecologia 51: 151-156.
Bailey, J.D. and Tappeiner, J.C. 1998. Effects of thinning on structural development in 40- to 100-year-old Douglas-fir stands in western Oregon. Forest Ecology and Management 108: 99-113.
Bennett, F., Lumsden, L.F. and Nicholls, A.O. 1995. Tree hollows as a resource for wildlife in remnant woodlands: spatial and temporal patterns across the northern plains of Victoria, Australia. Pacific Conservation Biology 1: 222-235.
Bowden, J. 1982. An analysis of factors affecting catches of insects in light-traps. Bulletin of Entomological Research 72: 535-556.
Brigham, R.M., Francis, R.L. and Hamdorf, S. 1997a. Microhabitat use by two species of Nyctophilus bats: a test of ecomorphology theory. Australian Journal of Zoology 45: 553-560.
Brigham, R.M., Grindal, S.D., Firman, M.C. and Morissette, J.L. 1997b. The influence of structural clutter on activity patterns of insectivorous bats. Canadian Journal of Zoology 75: 131-136.
Brower, J.E., Zar, J.H. and von Ende, C.N. 1990 Field and Laboratory Methods for General Ecology 3rd Edition, W.C. Brown Publishers, Dubuque, IA.
Brown, G.W., Nelson, J.L. and Cherry, K.A. 1997. The influence of habitat structure on insectivorous bat activity in montane ash forests of the Central Highlands, Victoria. Australian Forestry 60: 138-146.
Bullen, R. and McKenzie, N.L. 2001. Bat airframe design: flight performance, stability and control in relation to foraging ecology. Australian Journal of Zoology 49: 235-261.
Crampton, L.H. and Barclay, R.M.R. 1998. Selection of roosting and foraging habitat by bats in different-aged aspen mixedwood stands. Conservation Biology 12: 1347-1358.
Erickson, J.L. and West, S.D. 2003. Associations of bats with local structure and landscape features of forested stands in western Oregon and Washington. Biological Conservation 109: 95-102.
Fenton, M.B. 1990. The foraging behaviour and ecology of animal-eating bats. Canadian Journal of Zoology 68: 411-422.
Fenton, M.B., Boyle, N.G.H., Harrison, T.M. and Oxley, D.J. 1977. Activity patterns, habitat use, and prey selection by some African insectivorous bats. Biotropica 9: 73-85.
Fenton, M.B., Cumming, D.H.M., Rautenbach, I.L., Cumming, G.S., Cumming, M.S., Ford, G., Taylor, R.D., Dunlop, J., Hovorka, M.D., Johnston, D.S., Portfors, C.V., Kalcounis, M.C. and Mahlanga, Z. 1998. Bats and the loss of tree canopy in African woodlands. Conservation Biology 12: 399-407.
Fenton, M.B. and Morris, G.K. 1976. Opportunistic feeding by desert bats ( Myotis spp.). Canadian Journal of Zoology 54: 526-530.
Florence, R.G. 1996 Ecology and Silviculture of Eucalypt Forests. CSIRO Publishing, Collingwood.
Forestry Commission of NSW. 1983. Management Plan for Batemans Bay Management Area.
Forests NSW. 2008. State Forests of New South Wales, South Coast Region, silvicultural plan - spacing. 4 September 2008.
Fullard, J.H., Koehler, C., Surlykke, A. and McKenzie, N.L. 1991. Echolocation ecology and flight morphology of insectivorous bats (Chiroptera) in south-western Australia. Australian Journal of Zoology 39: 427-438.
Ghose, K. and Moss, C.F. 2003. The sonar beam pattern of a flying bat as it tracks tethered insects. Journal of the Acoustical Society of America 114: 1120-1131.
Gibbons, P., Lindenmayer, D.B., Barry, S.C. and Tanton, M.T. 2000. Hollow formation in eucalypts from temperate forests in southeastern Australia. Pacific Conservation Biology 6: 218-228.
Glass P.H. 1993. The response of insectivorous bat activity to disturbance from logging and habitat differences in the Kioloa State Forest. MSc thesis, University of Wollongong.
Grindal, S.D. and Brigham, R.M. 1998. Short-term effects of small-scale habitat disturbance on activity by insectivorous bats. Journal of Wildlife Management 62: 996-1003.
Grindal, S.D., Morissette, J.L. and Brigham, R.M.R. 1999. Concentration of bat activity in riparian habitats over an elevational gradient. Canadian Journal of Zoology 77: 972-977.
Hayes, J.P. 1997. Temporal variation in activity of bats and the design of echolocation-monitoring studies. Journal of Mammalogy 78: 514-524.
Hecker, K.R. and Brigham, R.M. 1999. Does moonlight change vertical stratification of activity by forest-dwelling insectivorous bats? Journal of Mammalogy 80: 1196-1201.
Hickey, M.B.C. and Fenton, M.B. 1996. Behavioural and thermoregulatory responses of female hoary bats, Lasiurus cinereus (Chiroptera: Vespertilionidae), to variations in prey availability. Ecoscience 3: 414-422.
Humes, M.L., Hayes, J.P. and Collopy, M.W. 1999. Bat activity in thinned, unthinned, and old-growth forests in western Oregon. Journal of Wildlife Management 63: 553-561.
JANIS. 1997. Nationally Agreed Criteria for the Establishment of a Comprehensive, Adequate and Representative Reserve System for Forests in Australia. A report by the Joint ANZECC/MCFFA National Forest Policy Statement Implementation Subcommittee.
Kalcounis-Ruppell, M.C., Psyllakis, J.M., Brigham, R.M. 2005. Tree roost selection by bats: an empirical synthesis using meta-analysis. Wildlife Society Bulletin 33: 1123-1132.
Kalko, E.K.V. and Schnitzler, H-U. 1993. Plasticity in echolocation signals of European pipistrelle bats in search flight: implications for habitat use and prey detection. Behavioral Ecology and Sociobiology 33: 415-428.
Keenan, R.J. and Ryan, M.F. 2004. Old growth forests in Australia: conservation status and significance for timber production. Revised Edition. Bureau of Rural Sciences, Australian Government.
Krusic, R.A., Yamasaki, M., Neefus, C.D. and Pekins, P.J. 1996. Bat habitat use in White Mountain National Forest. Journal of Wildlife Management 60: 625-631.
Kutt, A.S. 1995. Activity and stratification of microchiropteran bat communities in thinned, unthinned and old lowland regrowth forest, East Gippsland. The Victorian Naturalist 112: 86-92.
Law, B., Anderson, J. and Chidel, M. 1998. A bat survey in State Forests on the south-west slopes region of New South Wales with suggestions of improvements for future surveys. Australian Zoologist 30: 467-479.
Law, B.S. 2004. Challenges for managing bats in the State Forests of NSW. In Conservation of Australia's Forest Fauna 2nd Edition (Ed. Lunney, D.) pp. 748-760, Royal Zoological Society of New South Wales, Mosman.
Law, B.S. and Chidel, M. 2001. Bat activity 22 years after first-round intensive logging of alternate coupes near Eden, New South Wales. Australia Forestry 64: 242-247.
Law, B. and Chidel, M. 2002. Tracks and riparian zones facilitate the use of Australian regrowth forest by insectivorous bats. Journal of Applied Ecology 39: 605-617.
Law, B.S. and Chidel, M. 2004. Roosting and foraging ecology of the golden-tipped bat ( Kerivoula papuensis) on the south coast of New South Wales. Wildlife Research 31: 73-82.
Lee, Y.F. and McCracken, G.F. 2005. Dietary variation of Brazilian free-tailed bats links to migratory populations of pest insects. Journal of Mammalogy 86: 67-76.
Lloyd, A., Law, B. and Goldingay, R. 2006. Bat activity on riparian zones and upper slopes in Australian timber production forests and the effectiveness of riparian buffers. Biological Conservation 129: 207-220.
Loeb, S.C. and O'Keefe, J.M. 2006. Habitat use by forest bats in South Carolina in relation to local, stand, and landscape characteristics. Journal of Wildlife Management 70: 1210-1218.
Loyn, R.H., Fagg, P.C., Piggin, J.E., Morton, A.G. and Tolhurst, K.G. 1983. Changes in the composition of understorey vegetation after harvesting eucalypts for sawlogs and pulpwood in East Gippsland. Australian Journal of Ecology 8: 43-53.
Lumsden, L.F. and Bennett, A.F. 2005. Scattered trees in rural landscapes: foraging habitat for insectivorous bats in south-eastern Australia. Biological Conservation 122: 205-222.
Lunney, D., Barker, J., Priddel, D. and O'Connell, M. 1988. Roost selection by Gould's long-eared bat, Nyctophilus gouldi Tomes (Chiroptera: Vespertilionidae), in logged forest on the south coast of New South Wales. Australian Wildlife Research 15: 375-384.
McElhinny, C., Gibbons, P., Brack, C. and Bauhus, J. 2005. Forest and woodland stand structural complexity: its definition and measurement. Forest Ecology and Management 218: 1-24.
Medhurst, J.L., Beadle, C.L. and Neilsen, W.A. 2001. Early-age and later-age thinning affects growth, dominance and intraspecific competition in Eucalyptus nitens plantations. Canadian Journal of Forest Research-Revue Canadienne de Recherche Forestiere 31: 187-197.
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