Structure and spatial patterns of trees in old-growth northern hardwood and mixed forests of northern Maine

Stand structure including spatial patterns was studied in northern hardwood and mixed forest types in the 2000-ha old-growth Big Reed Forest Reserve in northern Maine using complete stem mapping, dendrochronology, and spatial analyses on ∼0.5 plots. The inclusion of saplings, dead wood, age distributions, spatial pattern, and interactions provided some idea of underlying processes and temporal change. Structural characteristics were most comparable to spruce-northern hardwood forests of northern New England and New York, and most characteristics matched expected patterns for old-growth forests of the region.Results indicated smaller maximum-tree sizes, lower basal areas (26–34 ) and downed-wood volumes (29–64), higher densities (475–649), but similar species longevities compared to other mesic old-growth forests further south and in the Lake States. The stands were dominated by very shade-tolerant tree species, including Fagus grandifolia Ehrh., Acer saccharum Marsh.,Picea rubens Sarg. and Abies balsamea(L.) Mill, with each species found in many crown positions and age and size classes. The sapling layer was dominated by Fagus grandifolia followed by Picea rubens. Most species had reverse-J shaped diameter distributions, but age distributions were indicative of synchronous, episodic recruitment. In most plots, Acer saccharum diameter distributions were skewed towards the mid-larger size classes. Lack of young and small Acer saccharum stems suggested change in forest composition towards Fagus grandifolia dominance. Most species formed small-scale clusters (≤15 ) perhaps in response to small gap disturbances. Snags were the dominant dead wood type and were randomly to regularly distributed in most plots. Logfall directions were unrelated to hurricane paths. Recent small-scale disturbance events and topographic position appear to be important in explaining current structure and dynamics of the hardwood and mixed forests of Big Reed Forest Reserve in northern Maine. The continued effects of beech bark disease had a greater effect on hardwood plots, whereas a recent spruce budworm outbreak had a greater effect on plots with higher conifer density.The dominance of very shade tolerant tree species in small-scale clusters, and randomly distributed snags rather than clustered uproots were indicative of the prevalence of small scale gap disturbance regimes in the hardwood and mixed forests of Big Reed Forest Reserve in northern Maine. Varying topographic position may allow for slight changes in disturbance regime leading to consequent variation in structure and dynamics. H1, a more open plot on upper exposed slopes, had distinctly different characteristics such as lower live and dead tree and sapling densities than the other plots, but more uprooted trees and Acer saccharum saplings. Such small scale gap disturbance regimes operating on an episodic basis, and effects of slight variations in this regime on stand composition and structure have significant implications for silvicultural interventions and management of these forest types.

[1]  W. Reiners,et al.  Vegetation patterns in a virgin subalpine forest at Crawford Notch, White Mountains, New Hampshire , 1983 .

[2]  Thomas G. Siccama,et al.  Presettlement and Present Forest Vegetation in Northern Vermont with Special Reference to Chittenden County , 1971 .

[3]  David R. Foster,et al.  Disturbance history, community organization and vegetation dynamics of the old-growth pisgah forest, South-Western New Hampshire, U.S.A , 1988 .

[4]  G. E. Nichols The Hemlock--White Pine--Northern Hardwood Region of Eastern North America , 1935 .

[5]  Emery R. Boose,et al.  Patterns of forest damage resulting from catastrophic wind in central New England, USA , 1992 .

[6]  G. B. Williamson Pattern and Seral Composition in an Old‐growth Beech‐Maple Forest , 1975 .

[7]  J. Henry,et al.  RECONSTRUCTING FOREST HISTORY FROM LIVE AND DEAD PLANT MATERIAL AN APPROACH TO THE STUDY OF FOREST SUCCESSION IN SOUTHWEST NEW HAMPSHIRE , 1974 .

[8]  W. A. Patterson,et al.  Mass of downed wood in northern hardwood forests in New Hampshire: potential effects of forest management , 1986 .

[9]  F. Bormann,et al.  Old-Age Stand of Hemlock-Northern Hardwood Forest in Central Vermont , 1964 .

[10]  Bengt Gunnar Jonsson,et al.  Uprooting in boreal spruce forests: long-term variation in disturbance rate , 1993 .

[11]  G. Parker,et al.  Spatial Dispersion of Woody Regeneration in an Old-Growth Forest , 1989 .

[12]  D. S. Powell,et al.  Forest statistics for Maine: 1971 and 1982 , 1984 .

[13]  S. Shifley,et al.  Ground cover in old-growth forests of the central hardwood region , 1997 .

[14]  F. Serre-Bachet,et al.  Identification, presentation and interpretation of event years and pointer years in dendrochronology. , 1990 .

[15]  H. F. Morey A Comparison of Two Virgin Forests in Northwestern Pennsylvania , 1936 .

[16]  Mike Rees,et al.  5. Statistics for Spatial Data , 1993 .

[17]  W. Mccomb,et al.  Snag densities in old-growth and second-growth Appalachian forests , 1983 .

[18]  Craig G. Lorimer,et al.  AGE STRUCTURE AND DISTURBANCE HISTORY OF A SOUTHERN APPALACHIAN VIRGIN FOREST , 1980 .

[19]  B. H. Honkala,et al.  Silvics of North America , 1990 .

[20]  S. R. Gemborys Structure and dynamics in a virgin northern hardwood-spruce-fir forest--the Bowl, New Hampshire , 1996 .

[21]  S. Payette,et al.  SEED DYNAMICS OF ABIES BALSAMEA AND ACER SACCHARUM IN A DECIDUOUS FOREST OF NORTHEASTERN NORTH AMERICA , 1991 .

[22]  R. Whittaker Forest Dimensions and Production in the Great Smoky Mountains , 1966 .

[23]  O. Loucks,et al.  AGE STRUCTURE MODELS OF BALSAM FIR AND EASTERN HEMLOCK , 1976 .

[24]  Richard A. Hardt A Comparison of Structural and Compositional Characteristics of Southern Appalachian Young Second- Growth, Maturing Second-Growth, and Old-Growth Stands , 2003 .

[25]  Thomas G. Siccama,et al.  What proportion of standing trees in forests of the Northeast are dead , 1990 .

[26]  G. Mroz,et al.  Composition, structure, and aboveground biomass of two old-growth northern hardwood stands in Upper Michigan. , 1985 .

[27]  W. D. Billings,et al.  A comparison of virgin Spruce/Fir forest in the northern and southern appalachian system. , 1951 .

[28]  T. Crow,et al.  Dynamics of dead wood in old-growth hemlock-hardwood forests of northern Wisconsin and northern Michigan , 1994 .

[29]  F. H. Bormann,et al.  Catastrophic disturbance and the steady state in northern hardwood forests , 1979 .

[30]  Peter J. Diggle,et al.  Statistical analysis of spatial point patterns , 1983 .

[31]  Charles D. Canham,et al.  Causes and consequences of resource heterogeneity in forests : interspecific variation in light transmission by canopy trees , 1994 .

[32]  J. Battles,et al.  Spruce decline as a disturbance event in the subalpine forests of the northeastern United States , 1996 .

[33]  A. Bouchard,et al.  Beech-maple dynamics in an old-growth forest in southern Québec, Canada , 1994 .

[34]  Thomas R. Crow,et al.  Structural Characteristics of Old‐Growth Hemlock‐Hardwood Forests in Relation to Age , 1994 .

[35]  Yan Liu,et al.  Coarse woody debris in an old-growth deciduous forest on the Cumberland Plateau, southeastern Kentucky , 1991 .

[36]  W. Romme,et al.  FIRE AND LANDSCAPE DIVERSITY IN SUBALPINE FORESTS OF YELLOWSTONE NATIONAL PARK , 1982 .

[37]  W. Leak Characteristics of Five Climax Stands in New Hampshire , 1987 .

[38]  R. B. Davis SPRUCE-FIR FORESTS OF THE COAST OF MAINE' , 1966 .

[39]  W. Leak Age Distribution in Virgin Red Spruce and Northern Hardwoods , 1975 .

[40]  C. Lorimer THE PRESETTLEMENT FOREST AND NATURAL DISTURBANCE CYCLE OF NORTHEASTERN MAINE , 1977 .

[41]  J. H. Zar,et al.  Biostatistical Analysis, 3rd edn. , 1996 .

[42]  M. Harmon,et al.  Ecology of Coarse Woody Debris in Temperate Ecosystems , 1986 .

[43]  D. Peart,et al.  Effects of beech bark disease on the growth of American beech (Fagus grandifolia). , 1993 .

[44]  R. Holmes Computer-Assisted Quality Control in Tree-Ring Dating and Measurement , 1983 .

[45]  Marianne E. Krasny,et al.  Gradual and sudden forest canopy gaps in Allegheny northern hardwood forests , 1992 .

[46]  S. Spurr,et al.  The virgin upland forest of central New England , 1942 .

[47]  R. Seymour The red spruce-balsam fir forest of Maine: Evolution of silvicultural practice in response to stand development patterns and disturbances , 1992 .

[48]  S. Payette,et al.  Disturbance regime of a cold temperate forest as deduced from tree-ring patterns: the Tantaré Ecological Reserve, Quebec , 1990 .

[49]  J. Worrall,et al.  Etiology of canopy gaps in spruce–fir forests at Crawford Notch, New Hampshire , 1988 .

[50]  W. Leak Some effects of forest preservation , 1974 .

[51]  Richard M. Klein,et al.  Spruce–fir decline and gap dynamics on Camels Hump, Vermont , 1992 .

[52]  Melinda Moeur,et al.  Characterizing Spatial Patterns of Trees Using Stem-Mapped Data , 1993, Forest Science.

[53]  N. H. Russell The Beech Gaps of the Great Smoky Mountains , 1953 .

[54]  C. Martin Distribution of tree species in an undisturbed northern hardwood-spruce-fir forest, the Bowl, N. H. , 1977 .

[55]  Bijan Payandeh Spatial pattern of trees in the major forest types of northern Ontario , 1974 .

[56]  Dietrich Stoyan,et al.  Marked Point Processes in Forest Statistics , 1992, Forest Science.

[57]  E. Cook,et al.  On predicting the response of forests in eastern North America to future climatic change , 1991 .

[58]  C. Federer,et al.  Growth decline in red spruce and balsam fir relative to natural processes , 1986 .

[59]  R. P. McIntosh,et al.  The Spruce‐Fir Forests of the Catskill Mountains , 1964 .

[60]  Tim Parshall,et al.  Canopy mortality and stand-scale change in a northern hemlock–hardwood forest , 1995 .

[61]  J. H. Zar,et al.  Biostatistical Analysis (5th Edition) , 1984 .

[62]  C. Canham Suppression and release during canopy recruitment in Acer saccharum , 1985 .

[63]  B. C. Larson,et al.  The Ecology and Silviculture of Mixed-Species Forests , 1992, Forestry Sciences.

[64]  D. Foster From coastal wilderness to fruited plain , 1995 .

[65]  F. Stearns The Composition of the Sugar Maple‐Hemlock‐Yellow Birch Association in Northern Wisconsin , 1951 .

[66]  S. Manabe,et al.  On the Distribution of Climate Change Resulting from an Increase in CO2 Content of the Atmosphere , 1980 .

[67]  L. Conkey Decline in old-growth red spruce in Western Maine: an analysis of wood density and climate , 1988 .

[68]  John A. Helms,et al.  Principles of Silviculture , 1979 .

[69]  D. Houston,et al.  Beech bark disease , 1998 .

[70]  R. T. Ward Some Aspects of Regeneration Habits of the American Beech , 1961 .

[71]  W. Leak Species and structure of a virgin northern hardwood stand in New Hampshire , 1973 .

[72]  D. D. Reed,et al.  Spatial autocorrelation of individual tree characteristics in loblolly pine stands , 1985 .

[73]  M. Applequist A simple pith locator for use with off-center increment cores , 1958 .

[74]  Richard Brewer,et al.  Wind Throw and Tree Replacement in a Climax Beech-Maple Forest , 1978 .

[75]  C. Canham Suppression and release during canopy recruitment in Fagus grandifolia , 1990 .

[76]  W. C. Fischer,et al.  Habitat Management for Hole-Nesting Birds in Forests of Western Larch and Douglas-fir , 1979 .

[77]  L. Irland,et al.  B819: The Spruce Budworm Outbreak in Maine in the 1970's–Assessment and Directions for the Future , 1988 .

[78]  G. Whitney,et al.  FIFTY YEARS OF CHANGE IN THE ARBOREAL VEGETATION OF HEART'S CONTENT, AN OLD-GROWTH HEMLOCK-WHITE PINE-NORTHERN HARDWOOD STAND' , 1984 .

[79]  Noel A Cressie,et al.  Statistics for Spatial Data. , 1992 .

[80]  W. H. Martin Characteristics of old-growth mixed mesophytic forests , 1992 .

[81]  G. E. Nichols The Vegetation of Connecticut IV. Plant Societies in Lowlands , 1915 .

[82]  L. Frelich,et al.  Age-class distribution and spatial patterns in an old-growth hemlock–hardwood forest , 1994 .

[83]  A. F. Hough,et al.  The Ecology and Silvics of Forests in the High Plateau of Pennsylvania , 1943 .

[84]  J. Hett A Dynamic Analysis of Age in Sugar Maple Seedlings , 1971 .

[85]  W. Platt,et al.  Replacement Patterns of Beech and Sugar Maple in Warren Woods, Michigan , 1996 .