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Determination of New Silvicultural Guidelines to Balance Sustainable Production of Forest Products and Critical Forest Habitat Structures inin ManageManaged Northern Hardwood Forests

Gregory McGee SUNY College of Environmental Science and Forestry [email protected]: 1 Forestry Drive Syracuse, New York 13210

Collaborators: Ralph Nyland, SUNY College of Environmental Science and Forestry William Leak, Mariko Yamasaki, US Forest Service, Northeast Research Station

March 31, 2007

¾Selection system with large maximum diameter limits, and reserve shelterwood systems can effectively retain biodiversity of arboreal organisms and sustain large wildlife cavities in managed northern hardwood forests. ¾Communities of canopy-dwelling organisms remain unquantified in northern hardwood systems, and great potential still exists to discover unknown arboreal species.

Funding support for this project was provided by the Northeastern States Research Cooperative (NSRC), a partnership of Northern Forest states (New Hampshire, Vermont, Maine, and New York), in coordination with the USDA Forest Service. Project Summary

„ Forest structural habitat features (SHFs) such as snags, decaying logs, and live, old/large trees are known to provide important habitat for a variety of forest-dwelling organisms in many forest types, and previous research has shown that SHF abundance can be regulated through management decisions. However, in the northern forest region, foresters currently have limited knowledge of how specific silvicultural prescriptions influence SHF abundance and how SHFs affect the abundance and diversity of many forest-dwelling taxa. Many managers seek to modify prescriptions to improve SHF availability while providing for sustainable production of wood products. Our objectives were to quantify SHF abundance in stands with definable and consistently applied silvicultural treatments, assess the role of tree diameter/age on arboreal and invertebrate communities, and incorporate wood decay functions into forest growth simulators in order to predict equilibrium log and snag volumes in modeled stands of varying management prescriptions. „ At replicate sites with known and consistently-applied management histories in central and northern New York, and New Hampshire, we sampled: decaying log and snag volumes; cavity tree densities; decayed wood bulk density in relation to known year of cutting (from 3-40 years prior to sampling); and arboreal lichen and mite communities. „ Our research conclusions are summarized below. – Stands managed under single-tree selection system for 2-3 entries over ~30 years, and even-aged maturing forests that regenerated following clearcutting or fire possessed approximately 50% of the decaying wood volume of unmanaged forests. Most of the difference was due to greater accumulation of large logs (>25 cm diameter) in the old forests. – We detected no differences in wood decay rates in even-aged versus uneven-aged managed stands. Approximately 80% of wood mass decays within 20 years. We are modifying a forest growth simulator in order to develop predictive models of equilibrium coarse woody debris volumes under differing management regimes. – Large (>55 cm dbh) sugar possess nearly twice the total lichen cover of small (15-35 cm dbh) trees. Large and small sugar maples supported different lichen communities. Stand history was less important than tree size in determining lichen abundance and composition, indicating that lichen diversity and abundance can be maintained in even- and uneven-aged managed stands as long as large, residual canopy trees are retained. – Arboreal, foliose supported more oribated mites per unit area than bare bark and crustose lichens. In what was the first quantitative assessment of arboreal mites in the northern forest, we found three new, undescribed species in a bark/lichen substrate sample totaling 0.25 m2. Research Background and Justification

¾ Forest structural habitat features (SHFs) such as standing dead trees (snags), decaying logs, and live, old/large trees are known to provide important habitat for a variety of forest-dwelling organisms in many forest types.

¾ However, habitat-wildlife relationships have not been explored for many taxa in northern hardwood systems. Old, unmanaged forests generally contain greater abundances of SHFs and associated forest-dwelling organisms than forests managed for commodity production. However, the studies from northeastern US reporting SHF abundance in managed forests have only stands receiving undefined, and inconsistently applied management prescriptions (i.e., “partial” or “selective” cuts).

These panels summarize (A) log A volumes, (B) snag basal areas, and (C) 160 live tree diameter distributions in partially-cut, 100-year-old maturing, and 120

) old-growth northern hardwood forests.

-1 high

ha 80 low 3 average B (m 40 std. err

) 12 -1 decaying log volume 0 ha 10 old growth maturing partially-cut 2 (n=8) (n=3) (n=3) 8 high 6 low 4 average std. err 2 C 0 snag basal area (m old growth maturing partially-cut (n=9) (n=2) (n=2) 250 200 -1 150 100 old-growth no. ha 50 maturing 0 partially-cut 15 25 35 45 55 65 75 85 95 Data are compiled from: Roskoski 1977; Roman 1980; Tritton 1980; Carbonneau 1986; Gore and Patterson dbh (cm) 1986; Dunwiddie 1993; Tyrrell and Crow 1994; Goodburn and Lorimer 1998; McGee et al. 1999 In order to provide useful information to forest owners and managers who seek to retain ecologically important structural habitat features we must:

¾ quantify the structural characteristics of stands that received well defined prescriptions, ideally over repeated entries; and ¾ more fully investigate the relationships between SHF and populations of forest-dwelling organisms that utilize them. Methodology

Experimental forest stands established by Ralph Nyland at the SUNY-ESF Huntington Wildlife Station and on NY State Forest lands, and by William Leak at the USFS Bartlett Experimental Forest, were used to quantify the structural features of even-aged and uneven-aged managed northern hardwood stands following multiple entries using precisely defined silvicultural prescriptions.

Bartlett Forest

Huntington Forest

NY State Forests Methods: forest structure study

We quantified log, snag and stump volume, and decay distributions in the following stand types: ¾ single-tree selection ¾ shelterwood ¾ clearcut w/thinning ¾ clearcut w/out thinning ¾ Late-successional / old- growth Methods: canopy lichen study Heather Root, an MS student at SUNY-ESF climbed into the canopies of 81 sugar trees in old-growth, selection system and reserve shelterwood stands to quantify arboreal lichens. She tested the following hypotheses:

A bird’s eye view of a reserve shelterwood – Tree age / size influence system at SUNY-ESF’s Huntington Wildlife epiphytic lichen abundance Research Station, Newcomb, NY. and community composition.

– Stand level management activities influence epiphytic lichen abundance and community composition. Methods: arboreal mite study

Heather Root also conducted the first description of arboreal mite communities associated with sugar maple trees in Adirondack northern hardwood forests.

Her objectives were to determine whether: (1)foliose lichen mats provide better habitat than crustose lichens and bare bark; and (2)mite abundance and community composition differ between Crustose and large sugar maple trees in foliose lichens reserve shelterwood versus old- Bare bark and foliose growth forests. lichens Methods: wildlife Laura Hartley discusses sampling cavity study protocols with Greg McGee Laura Hartley, an undergraduate student at SUNY-ESF, and Joshua Schneider an MS student at SUNY-ESF collaborated to quantify the relationship between tree size and stand-level management activities on the abundance and quality of wildlife cavities at our study sites. Laura Hartley discusses sampling protocols with Greg McGee Methods: wood decay study

We located and removed wood samples of logging debris left during operations that occurred between 2002 and 1958.

Wood bulk densities and volumes were determined in order to estimate rates of coarse woody debris mass loss through time and under differing disturbance regimes. Results: canopy lichen study

60 old growth selection shelterwood 50

40

30 ¾ Sugar maple crowns 20 had greater percent

Average percent cover percent Average 10 cover of lichens than the boles. 0 ¾ Large Medium Small Large Medium Small Crowns of large (>55 BOLE CROWN cm dbh) trees possessed nearly 2- fold greater lichen cover than those of small (15-35 cm dbh) trees. Results: canopy lichen study aS

dS aM

jM gS 2005ba sS cS OG sM oL cM aL jL oM gM mS cL

Axis 2 bL SS dL size stand type RSW gL jS mM old growth reserve shelterwood oS selection system bM sL mL

bS dM

Axis 1

¾ This ordination illustrates differences in lichen community composition on large (L), medium (M) and small (S) sugar maple trees in old-growth, reserve shelterwood and selection stands (small letters are site codes). Lichen communities on large trees were dominated by different species than communities on small trees. ¾ In old-growth and selection stands, lichen communities on small trees contained many component species of the large trees; however lichen communities on small trees in reserve shelterwoods were substantially different from those on large trees. Results: canopy lichen study

¾ The lichen species Ramalina intermedia, quercizans, and Lobaria pulmonaria which have been described elsewhere as “old growth indicators,” may be more appropriately referred to as “large tree indicators” since they occurred in all stand types, as long as larger trees were present.

¾ A group of nitrophilous species, which are well- dispersed, and establish quickly, characterized small trees and were more abundant on trees in the reserve shelterwoods and selection stands. Results: arboreal mite study

¾ In a sample of bark and arboreal lichen substrate totaling approximately 0.25 m2, twenty-five oribated mite species were sampled, three of which were undescribed species.

¾ Heather Root proposed naming one of the new species Anachipteria sacculifera. The species name “sacculifera” refers to the nature of the pore-like sensory organs, which take on a sack-like form in this species (images 14, 24, 25). Results: arboreal mite study

¾ Foliose lichens provided vital Aveage (1SE) mite density on bare bark, and in crustose habitat for the lichens and three foliose lichens on sugar maples in mite Adirondack northern hardwood forests. communities. There were 35 approximately 4- 30 to 7-x more mites in foliose lichens 25 than on bare 20 bark and 15 crustose lichens. 10 5 ¾ Mite abundance

no. individuals/sample did not differ 0 among common bark crustose foliose foliose foliose species of foliose Flacap Parsqu Punrud lichens.

Crustose lichen is Pertusaria velata Foliose lichens are Flavoparmelia caperata, Parmelia squarossa, Punctelia rudecta Results: arboreal mite study

Arboreal mite indicator species analysis for five substrates associated with sugar maple trees in Adirondack northern hardwood old growth and reserve shelterwood stands. ¾ Communities differed among Mite species Substrate or Stand Type Indicator value P bark, crustose lichen, and Achipteriidae imm. Foliose lichen 47.6 .004 foliose lichens. Anachipteria n. sp. Foliose lichen 59.1 .002 ¾ Trees in old Carabodes higginsi Foliose lichen 70.5 .001 growth and Ommatocepheus sp. imm. Bark 31.2 .013 reserve shelterwood stands supported different mite Achipteriidae imm. Reserve shelterwood 70.1 .102 communities. Anachipteria magnilamellata Old growth 100 .102 Anachipteria n. sp. Reserve shelterwood 75.1 .102 Caleremaeus sp. 1 Reserve shelterwood 81.8 .102 Platyliodes sp. imm. Old growth 100 .102 Results: Coarse Woody Debris Comparisons

Consistent with previous studies, old, unmanaged forests tended to have greater total coarse woody debris volume than maturing, 100-yr old stands, and stands managed under strict single-tree selection system guidelines for 2- 3 rotations over 30 years. Equal volumes of small-diameter material were produced in all stand types, reflecting the mortality of suppressed trees and loss of smaller branches. The old stand contained greater volumes of material >25 cm dia.

200 >49 cm 26-49 cm 18 0 10 - 2 5 c m 16 0 <10 cm 14 0 12 0 10 0

80 60 40 20

0

100 yr s e le c tio n o ld

stand type Results: Incorporation of Wood Decay Processes in Stand Growth Models Masters student Joshua Schneider is competing an integration of coarse woody debris decay functions into a modified Fiber 3.0 forest stand growth simulator to model long-term coarse woody debris dynamics in selection system stands. This work is being coordinated with another NSRC-funded project under the direction of Ralph Nyland, Bill Porter and Ed Bevilacqua.

Percent mass loss of coarse woody debris residue in managed northern hardwood forests as a function of cutting year

100 80 60 40

mass loss (%) 20 0 1950 1960 1970 1980 1990 2000 2010

year Results: Incorporation of Cavity Development in Stand Growth Models Joshua Schneider is also analyzing cavity occurrence as a function of stand history, tree size and tree species, and will be integration cavity development functions into the Fiber 3.0 forest stand growth simulator to model cavity occurrence in selection system stands.

Relationship between tree diameter and percent of trees containing at least one functional cavity of a given opening 20 size.

15 3-5 cm 6-10 cm 10 11-15 cm with cavity 5 percent of trees of percent

0

15-24 25-34 35-44 45-54 >54 tree diameter Implications and applications in the Northern Forest region

¾ Much of the northern forest is privately owned and managed and the regional maintenance of biodiversity in this working forest will require knowledge of the impacts of forest management decisions on a variety of forest dwelling organisms. ¾ Our data advance the collective understanding of the ecological roles of large, old residual trees. Specifically, large trees ¾ support greater amounts of arboreal lichens; ¾ provide habitat for unique assemblages of lichens not found on smaller, younger trees; ¾ support diverse and unknown invertebrate communities in their lichen mats; ¾ contain greater numbers of large wildlife cavities than smaller trees. Implications and applications in the Northern Forest region (cont’d)

¾ Our research will be used modify existing stand growth models to predict CWD supply in selection system forests of differing residual basal areas, diameter limits and entry cycles, which will allow resource managers to make educated decisions regarding the costs and benefits of retaining varying levels of structural complexity in managed woodlots. ¾ Furthermore, our research indicates that much remains unknown about communities of arboreal organisms in northern forests. Future Directions: We will continue working to elucidate the Snail and Evernia mesomorpha relationships between SHF abundance and populations of forest- dwelling organisms in northern forests. In particular, Heather Root’s study revealed that epiphytic lichens serve as important habitat for mites, but numerous other organisms clearly utilize lichen mats for food or shelter. We also plan to pursue more detailed studies of the communities and trophic interactions of arboreal organisms.

Slug on Lobaria quercizans and Frullania eboracensis List of products

Peer Reviewed Publications ¾ Root, H.T., A.Y. Kawahara and R.A. Norton. (in review). Anachipteria sacculifera n. sp. (Acari: Oribatida: Achipteriidae) from arboreal lichens in New York State. Acarologia. ¾ Root, H.T., G.G. McGee and R.D. Nyland. 2007. Effects of two silvicultural management regimes with large tree retention on epiphytic lichen communities in Adirondack northern hardwoods, New York, USA. Canadian Journal of Forest Research (in press). ¾ Root, H.T., G.G. McGee and R.A. Norton. 2007. Arboreal mite communities on epiphytic lichens of the Adirondack Mountains of New York, USA. The Northeastern Naturalist (in press).

Theses ¾ Schneider, Joshua. (in preparation for May 2007). ¾ Root, Heather, T. 2006. Arboreal lichen and mite communities in Adirondack northern hardwood forests and their responses to forest management. Masters Thesis. State University of New York College of Environmental Science and Forestry. Syracuse, New York. 78 pp.

Other Publications ¾ McGee, G.G. Forest Ecosystem Management: A New Paradigm for Preserving Forest Biodiversity. 2007. Network for Conservation Educators and Practitioners, American Museum of Natural History. [Available online at: http://cbc.amnh.org/ncep/us/module_files.php?module_instance_id=400]. List of products, continued

Posters ¾ McGee, G. H. Root and L. Hartley. The Maintenance of Biodiversity Through Green Tree Retention in Managed Northern Hardwood Forests (poster). NY SAF Annual meeting, Liverpool, NY, February, 2006. ¾ Hartley, L., G. McGee, R. Nyland, W. Leak, M. Yamasaki. Comparison of wildlife cavity abundance in northern hardwood forests receiving different silvicultural treatments (poster). NY SAF Annual Meeting, Liverpool, NY, February, 2006.

Professional Presentations ¾ Root, H., and G.G. McGee. Epiphytic Lichen Community Response to Forest Management on in the Adirondack Mountains of New York (oral presentation) Northeastern Natural History Conference, Albany, NY, April, 2006.

¾ Root., H. and G.G. McGee The effects of forest management on epiphytic lichen communities of the Adirondack Mountains of New York. American Bryological and Lichenological Society, Annual Meeting, June, 2006. List of products, continued

Other media outlets ¾ Root and McGee interviews with Michael Cannon, Associated Press, Albany, May 2006 leading to the following outlets. • http://science.monstersandcritics.com/news/article_1158101.php/Three_ new_species_of_mites_are_found • http://www.physorg.com/news65164275.html • http://www.newswise.com/articles/view/519892/ • http://news.webindia123.com/news/Articles/Science/20060425/315715.h tml • http://www.postchronicle.com/news/science/article_21215828.shtml

¾ Root interview w/ Brian Mann, North Country Public Radio, Canton, 4/27/06; aired 5/4/06. ¾ Root, McGee and Nyland on-film interviews w/ Thom Hallock, WPTZ Burlington, for “Champlain Connection” feature segment, July 2005.