Forest Stand Dynamics

Defining Forest Stand Dynamics Forest dynamics describes the underlying physical and biological forces that shape and change a forest –Continuous state of change altering forest composition and structure –Basic elements of forest dynamics: Disturbance Succession

Disturbance and Succession Forest disturbance is an event that causes change in forest structure and composition, resource availability, and the physical environment Succession is the process that gradual replacement of one community of plants by another

Disturbance and Succession

Range of Forest Disturbance Forest disturbances vary in type, frequency, spatial scale, and severity Disturbance types: Fire, wind, insects, disease, flood, ice storms, grazing/herbivory, timber harvest, road construction, conversion to nonforest A continuum of disturbance from individual tree-level to landscape scale

Major (Stand-Replacing) Disturbance

Phases of Stand Development Following Major (Stand-Replacing) Disturbance 1.Stand initiation (reorganization phase) 2.Stem exclusion (aggradation phase) 3.Understory reinitiation (transition phase) 4.Old-growth (complex phase, steady-state) Each phase of stand development is accompanied by changes in stand structure and species composition.

Stand Initiation Stage Follows major disturbance (wind, fire, clearcuts) Regeneration from seed, sprouts, or advance reproduction Rapid increase in the number of stems and biomass Structure –Single cohort (“even-aged”) stand –“Brushy” stage with herbaceous, shrub, small trees –Invasion continues until all growing space is occupied Stage ends when canopy becomes continuous and trees begin to compete with each other for light and canopy space

Stem Exclusion Stage Begins at about crown closure, characterized by onset of density dependent mortality (“self-thinning”) –Canopy continues to have one cohort and canopy too density to allow new trees to grow into canopy –Crown differentiation occurs Biggest trees tend to get bigger, the smaller ones tend to die Mortality rates are high, especially in intermediate and overtopped crown classes Least competitive individuals die –Crowns are small enough so that when a tree dies, others fill the vacant growing space by expanding their crowns Phase ends when biomass peaks

Crown Classification Overtopped

Crown Classification Dominant: Crown is larger than average and typically above the general upper level of the canopy; receives full top light, considerable side light Codominant: Top of crown is at upper canopy height; receives full top light, little from sides; medium-sized crown, usually somewhat crowded on its sides. Often wide range around “average canopy” tree. Intermediate: Top of crown is below the top of the general canopy; receives some top light from directly above, none from the side; conspicuously narrower, smaller and shorter than the average crown. Overtopped: Crown entirely below some foliage of the upper canopy; receives no direct top light; small, weak crown with low vigor

Understory Reinitiation Stage Mortality of individuals cannot be closed by adjacent individuals –Crowns of trees are now large enough so that when one overstory tree dies, the surrounding trees can not fill the gap Permanent canopy gaps form Permanent understory forms –Tree reproduction becomes re-established beneath parent stand –Primary factors influencing species composition Understory light availability Species degree of shade tolerance

Old-Growth (or Complex) Stage Natural mortality of large overstory trees produces irregular canopy gaps Mortality and recruitment and are in balance and biomass is stable Can mark transition from an even-aged to an uneven-aged stand

Stand Initiation Stem Exclusion Understory Reinitiation Old growth (Complex Stage)

Johnson, P.S., S.R. Shifley, and R. Rogers The Ecology and Silviculture of Oaks. CABI Publishing, New York, NY. 503 p. Stand Development in the Central Hardwood Region Stand Initiation Phase: 10 – 20 years

Johnson, P.S., S.R. Shifley, and R. Rogers The Ecology and Silviculture of Oaks. CABI Publishing, New York, NY. 503 p. Stand Development in the Central Hardwood Region Stem Exclusion: Begins after 10 years Concludes before age 70

Johnson, P.S., S.R. Shifley, and R. Rogers The Ecology and Silviculture of Oaks. CABI Publishing, New York, NY. 503 p. Stand Development in the Central Hardwood Region Understory Reinitation: Tree reproduction begins to develop under maturing overstory Begins after age 50, concludes before age 120

Johnson, P.S., S.R. Shifley, and R. Rogers The Ecology and Silviculture of Oaks. CABI Publishing, New York, NY. 503 p. Stand Development in the Central Hardwood Region Complex Stage (old-growth): Oak forest typically require 100 years or longer to reach this stage.

Johnson, P.S., S.R. Shifley, and R. Rogers The Ecology and Silviculture of Oaks. CABI Publishing, New York, NY. 503 p. *Assume no significant stand-scale disturbances occur *Actual durations of stages of development vary with species composition, site productivity, and other factors Stand Development in the Central Hardwood Region

Tree Growth and the Environment

Photosynthesis Photosynthesis: Conversion of light energy to chemical energy –Production of carbohydrates from CO 2 and H 2 O in the presence of chlorophyll using light energy. 6CO H 2 O C 6 H 12 O 6 + 6O 2 + 6H 2 O –Photosynthetic activity is a major factor in the production of biomass –Rates of photosynthesis are influenced by both plant and environmental factors chlorophyll light

Respiration Respiration is the process by which energy fixed by photosynthesis is made available for metabolic processes

Environmental Factors Influencing Photosynthesis Light Temperature CO 2 concentration Water availability Nutrient availablity

Environmental Factors Influencing Photosynthesis Light –Photosynthesis uses solar radiation in the visible spectrum ( nm wavelengths) Wavelength range known as photosynthetically active radiation (PAR) –Light directly affects tree growth by its intensity, quality, and duration

Environmental Factors Influencing Photosynthesis Light –Light environment in a stand is influence by the vertical and horizontal forest structure Density and vertical distribution of foliage alters light transmittance from sky to forest floor Silviculturists manipulate light environment in stands by altering forest structure

Inverse Relationship between Canopy Openness and Light Availability

Light and Photosynthesis Below some light, carbon uptake is negative, as respiration exceeds photosynthesis

Light and Photosynthesis As light increases, a light compensation point is eventually reached where CO 2 through photosynthesis is exactly balanced by losses through respiration

Above the light compensation point, photosynthesis increases until the amount of carboxylation enzyme or available CO 2 limits photosynthesis. Plateau in the rate of photosynthesis is know as light saturation point

Light and Photosynthesis Light compensation points and light saturation levels vary: –Among species –Among individuals of the same species –Among leaves on the same tree –With changing environmental conditions

Environmental Factors Influencing Photosynthesis Temperature –Temperature is a very important factor in photosynthesis but unlikely to become a limiting factor in forests of temperate regions except during the winter

Environmental Factors Influencing Photosynthesis CO 2 concentration –CO 2 concentration in the atmosphere is low, about 0.03% by volume –Concentrations in the forest are often higher but show vertical gradients which change diurnally and seasonally –Stands whose structure permits continued circulation of air provide more favorable conditions from the standpoint of CO 2 supply than those with a tight canopy or those which are multi- storied. –CO 2 enrichment (i.e. atmospheric rise due to fossil fuel burning) has been shown to increase growth rates

Environmental Factors Influencing Photosynthesis Water availability –Only minute quantities of water are consumed in the process of photosynthesis –The main effect of water on photosynthesis is indirect through hydration of protoplasma and stomatal closure

Environmental Factors Influencing Photosynthesis Water availability –Issues with moisture availability for photosynthesis and hydration: Water stress in temperate regions occurs on xeric to intermediate quality sites and due to seasonal drought Water is a primary growth limiter in arid and semi-arid regions –Wider spacing is one way for the silviculturists to reduce the impact of low moisture supply in such regions arid and semi- arid regions (U.S. southwest and mountain west)

Environmental Factors Influencing Photosynthesis Water availability –Moisture availability is dictated by Soil properties –Soil texture –Soil profile and depth Topography –Aspect –Slope position –Slope shape –Elevation

Environmental Factors Influencing Photosynthesis Water availability and topography –Aspect Solar radiation exposure strongly effects evapotranspiration –North, northeast, and east slopes more cool and moist –West, southwest, and south slopes have highest transpiration loss due to perpendicular orientation to incoming solar radiation –Slope position Upper slopes drier Middle to lower slope positions more moist and productive –Slope shape Convex landforms shed water Concave accumulate water

Environmental Factors Influencing Photosynthesis Nutrients –Photosynthetic efficiency of foliage depends decisively on soil nutrient supplies –With improving nutrient status among sites photosynthetic capacity of trees also improve –The effect is both direct (i.e., quantity of CO 2 fixed by gram of foliage) and indirect by increasing size of individual leaves, total size of crown and root system –Nutrient availability is dictated by a site’s soil properties Exception is the use of fertilization

Plant Factors Influencing Photosynthesis Leaf age Position within crown Crown class and species Sun and shade adaptations

Plant Factors Influencing Photosynthesis Leaf age –In conifers fully expanded one-year-old foliage is the most efficient of all age classes –Difference between age classes is mainly a consequence of varying rates of respiration, and by insect or disease damage

Plant Factors Influencing Photosynthesis Position within crown –The most productive leaves are in the upper crown. The lowest whorls contribute little to net photosynthesis.

Plant Factors Influencing Photosynthesis Crown class and species –Differences in photosynthetic efficiency between dominant, co- dominant, intermediate, and overtopped trees are relatively minor when one compares similarly exposed foliage and expresses efficiency per unit of leaf surface –The major factor causing differences in photosynthetic capacity of trees of different crown classes and of different species is the enormous difference found in leaf area.

Plant Factors Influencing Photosynthesis Sun and shade adaptations –Not all tree species possess the same photosynthetic efficiency –Photosynthetic rates and efficiency also varies with species shade tolerance Shade tolerant, intermediate tolerance, and intolerant –Photosynthetic efficiency varies between shade and sun leaves on the same tree Shade leaves and shade-tolerant species have higher photosynthetic efficiency per unit of leaf area under low light conditions than sun leaves Under high light conditions the reverse is true

The Carbon Budget of Trees Carbon budget of a tree (or any plant) can be expressed like a bank balance: Income = carbohydrates manufactured in photosynthesis Expenditures = carbohydrates used in growth and maintenance (construction and maintenance respiration) Balance = carbohydrates stored (so-called nonstructural carbohydrates and other compounds)

Individual Tree Growth Amount of carbohydrates produced through photosynthesis by a given tree is influence by: –Resources available to the tree (i.e., growing space) –Ability to harvest light and the roots ability to supply the foliage Extent to which a tree increases mechanical support (i.e., stem diameter) depends upon: –Carbohydrates remaining after supplying essential functions

Shade Tolerance Shade tolerance –Definition: Having the capacity to compete for survival under shaded conditions Understanding of shade tolerance is a cornerstone of silviculture Critical to silviculture in the following ways: –Regeneration of desired tree species –Regulating forest growth and understanding plant succession –Influencing species composition

Shade Tolerance and Photosynthesis Shade tolerant species –Species adapted to growing at reduced light intensities –Generally have lower compensation points and levels of light saturation than shade intolerant species Shade intolerant species saturate at relatively high light levels –Yield increased carbon gain in high light environments

Shade Tolerant vs. Intolerant Trees Regeneration –Tolerant : Regenerate and form understories beneath canopies of less tolerant trees or even beneath their own shade. Example: Red maple under oak dominated overstory –Intolerant: Regenerate most successfully in the open or in canopy gaps

Shade Tolerant vs. Intolerant Trees Ability to Persist in the Understory –Tolerant: Able to establish and persist in shaded understory –Intolerant: Sometimes establish in shaded understory, but they cannot survive for extended periods without increased understory light availability Remember: All this is relative! It is a manner of degree

Shade Tolerant vs. Intolerant Trees Response to Release –Tolerant trees: When released by canopy opening, they respond rapidly and maintain good growth Examples: Red maple, eastern hemlock –Intolerant trees: Normally die (or are significantly suppressed) following long- periods in dense shade If they are released after a long-period in dense shade, they respond with sluggish growth –Even tolerant trees may have trouble following release if in the understory too long

Shade Tolerant vs. Intolerant Trees Crown Characteristics –Tolerant trees: Have heavy crowns of several leaf layers, the innermost remaining functional in very low levels of light –Intolerant trees: Have thin, open crowns of well-lighted leaves.

Shade Tolerant vs. Intolerant Trees Natural Pruning –Tolerant trees: clean their boles of side branches relatively slowly because the leaves remain alive in low light Examples: eastern hemlock, sugar maple, American beech –Intolerant trees: Clean their trunks rapidly, "self-pruning", even if grown in the open Examples: shortleaf pine, yellow-poplar

Shade Tolerant vs. Intolerant Trees Bole Form –Tolerant trees: Because of differences in the degree of natural pruning, tolerant trees have more cone-shaped boles Examples: American beech, sugar maple, hemlock –Intolerant trees: Tend to have cylindrical-shaped boles Examples: poplars, southern pines

Shade Tolerant vs. Intolerant Trees Seed Production –Tolerant trees: Reach seed bearing age late and may produce periodic seed crops –Intolerant trees: Produce seed early in live and produce large, regular seed crops

Adaptive Strategies in Reference to Tolerance Intolerants –Capacity for rapid establishment on disturbed sites Early seed production Prolific seeders Light seeds, wide dispersal Rapid germination –Fast juvenile growth in full light –Adaptation to extreme sites (dry, wet, cold, hot) –Colonize from a refuge site

Adaptive Strategies in Reference to Tolerance Tolerant species –Typically adapted to sheltered, moist, fertile sites –Gradually replace intolerants in the absence of disturbance A special case: gap-phase species T olerant when juvenile Capable of outgrowing tolerant species when canopy gap created Example: eastern white pine

Silvics in Central Hardwood Forest Region

Central Hardwood Forest Region

25.4 million acre forestland base

Silvical Characteristics of KY Major Species SpeciesSeed Dissmemination Ecological Strategy GravityAnimalsWindExploitiveConservative Yellow-poplar X X White oakXX X Chestnut oakXX X Black oakXX X Northern red oakXX XX Scarlet oakXX XX Sugar maple X XX Red maple X X Pignut hickoryXX X American beechXX X

Silvical Characteristics of KY Major Species SpeciesShade Tolerance Growth Rate Longevity IntolerantIntermediateTolerant SlowMediumFast < >200 Yellow-poplarX X X White oak X X X Chestnut oak X X X Black oak X X X Northern red oak X X X Scarlet oakX X X Sugar maple X X X Red maple X X X Pignut hickory X X X American beech X X X

Need More Information? Silvics of North America Volume 1: ConifersVolume 2: Hardwoods

Growth and Yield of Stands

The Stand The basic unit for silvicultural practice Stands are usually classified by age, composition, and structure

Site Site is the sum of the effective environmental conditions under which a forest lives Site factors can be grouped as: –Climatic –Edaphic –Physiographic –Biotic Site quality is the capacity of a site for production –Two categories of site indicators are used Direct measurement of environment Correlates such as Site Index

Site Site Index (SI): A measure of actual or potential forest productivity expressed in terms of the average height of dominants and co- dominants in the stand at an index age (base age) for a particular species. –Base age: 50 years for hardwoods 25 years for southern pine

Example: Site Index Chart

Growth Growth is increase in size of an individual or a stand Growth is usually expressed as a change in size per unit time and area

Mean Annual Growth Mean Annual Increment (MAI): Average annual growth a stand has exhibited up to a specified age where, Y a = yield at given age a = age

MAI Calculations Example: –A stand was inventoried in 2013 –In 2013, the stand was 60 years old –Stand volume was 3200 ft 3 per acre in 2013 Calculate MAI in year 2013:

Periodic Growth Periodic Annual Increment (PAI): Average annual growth a stand exhibited during a specific time period where, Y is the yield at times 1 and 2 T 1 represents the year starting the growth period, and T 2 is the end year

PAI Calculations Example: –A stand was inventoried in 2003 and 2013 –In 2013, the stand was 60 years old –Stand volume was 2000 ft 3 per acre in 2003 and 3200 ft 3 per acre in 2013 Calculate PAI:

Relationship between PAI and MAI Net Yield Periodic annual increment (PAI) and mean annual growth (MAI) rises, peaks, and declines through time

Relationship between PAI and MAI Net Yield Stage 1: Rotation for maximum fiber production ends when PAI = MAI

Relationship between PAI and MAI Net Yield Stage 2: Rotation for sawtimber production with exact length determined by economic criteria

Relationship between PAI and MAI Net Yield Stage 3: Understory reinitiation phase where mortality exceeds production and standing volume declines progressively

Yield Yield is the quantity of harvestable material or attributes produced on a defined area of land Yield is usually expressed as a rate, quantity per unit time and area The most fundamental forest yield calculation relates solar energy input to crop output

Gross versus Net Yield Gross yield –Total amount produced on a given site at a given age (e.g., volume of living trees + volume of mortality) –Rises throughout stand development, but declines in complex phase Net yield –Yield (volume or biomass) available for removal at any given age –Rises throughout much of a stand’s life, but gradually falls below gross yield as mortality accumulates

Growth Patterns of Even-Aged Stands Tree Density –Decreases continuously due to mortality as the stand ages Schnur, G.L Yield, stand, and volume tables for even-aged upland oak forests. US Department of Agriculture, Technical Bulletin No p.

Average Tree Diameter –Diameter (dbh) of average tree increases throughout the life of the stand as trees grow and as smaller trees suffer a disproportionately higher mortality rate Schnur, G.L Yield, stand, and volume tables for even-aged upland oak forests. US Department of Agriculture, Technical Bulletin No p.

Basal Area –Basal area increase throughout the life of a stand. In some species, basal area rises to a plateau and then remains fairly constant (growth balanced by mortality). Schnur, G.L Yield, stand, and volume tables for even-aged upland oak forests. US Department of Agriculture, Technical Bulletin No p.

Tree Height –Height of dominant and codominant trees increases through life of stand –Normally level off or flatten as stands become decadent

Influence of Site Quality –Average height growth of canopy trees is primarily dependent on site quality except at extremely low or high densities Site Quality and Stand Growth

As site quality (SI) increases overstory trees grow in height more quickly Schnur, G.L Yield, stand, and volume tables for even-aged upland oak forests. US Department of Agriculture, Technical Bulletin No p.

Stand density is lower at a given age on high quality sites when compared to low quality sites* *Assuming stands have similar species composition, disturbance histories, and initial stand conditions.

Average tree diameter is larger at a given age on high quality sites when compared to low quality sites* *Assuming stands have similar species composition, disturbance histories, and initial stand conditions.

At a given age, taller and larger trees are present on higher quality sites when compared to lower quality sites. Hence, more volume accumulates on high quality sites.* *Assuming stands have similar species composition, disturbance histories, and initial stand conditions.

Influence of Site Quality on Stand Development As site quality (SI) increases: –Trees grow in height more quickly –Stands develop closed canopy more rapidly –Competition induced mortality begins earlier –More rapid stand development results in: Lower densities, larger average diameter, and more volume at a given age on high quality sites when compared to low quality sites –More rapid volume accumulates because of taller, larger trees present at a given age

Influence of Species on Growth Source: Assmann 1970

Influence of Species on Height Growth

Influence of Species on Diameter Growth

Influence of Stand Density on Height Growth Height growth of overstory trees is only effected by extreme stand densities Open-grown trees and overstory trees growing in extremely high densities will generally have reduced heights when compared to other trees growing on a similar quality site Height growth of intermediate and overtopped crown class trees is reduced by shading effects of the overstory

Influence of Stand Density on Diameter Growth Individual tree diameter growth increases as the amount of available growing space increases 1200 trees ac trees ac trees ac trees ac trees ac -1

Influence of Stand Density on Diameter Growth Stand density is a primary driver of tree diameter growth However, at a given stand density, diameter growth is generally higher on better quality sites

Relationship between planting spacing and stand density over time Trees per Hectare Density dependant mortality increases as the amount of available growing space decreases Influence of Stand Density on Mortality

Relationship Between Density and Tree Volume Growth CrowdedIsolated Trees per Acre Wide Spaced Well Spaced Volume per Tree Patterns in volume per tree mirrors amount of growing space available per tree. Adapted from: Daniel et al. 1979, Smith et al. 1997

Crowded Isolated Trees per Acre Wide Spaced Well Spaced Volume per Acre Total Volume Merchantable Volume or Total Volume in Species Susceptible to Stagnation at High Densities Adapted from: Daniel et al. 1979, Smith et al Relationship Between Density and Tree Volume Growth

Relationship Between Density and Tree/Stand Volume Growth Crowded Isolated Trees per Acre Wide Spaced Well Spaced Volume per Tree Volume per Acre Total Volume Merchantable Volume or Total Volume in Species Susceptible to Stagnation at High Densities Patterns in volume per tree mirrors amount of growing space available per tree. Adapted from: Daniel et al. 1979, Smith et al. 1997