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PROBABILITY AND MODELS
OF
TREE MORTALITY
Jeetendra Gautam
MSc Forestry (General Forestry)
Roll No. 14
TREE MORTALITY
• PLANT DEATH IS A COMPLEX PROCESS,
INFLUENCED BY
• PHYSIOLOGY,
• ENVIRONMENT

• SUCCESSIONAL DEVELOPMENT,
• AGE, AND,
• CHANCE

- (HARCOMBE, 1987; FRANKLIN ET AL., 1987)
TREE MORTALITY
• GENERAL CAUSES
• LACK OF SUFFICIENT RESOURCES TO RESTRAIN STRESS,
INJURIES OR SUSTAIN LIFE, OR KILLED BY EXTERNAL
FACTOR (WARING, 1987)
• NEGATIVE CARBON BALANCE (RESPIRATION >
PHOTOSYNTHESIS)
• INSECT INFESTATION

• PROLONGED DROUGHT UNDER LOW LIGHT CONDITION
TREE MORTALITY PROCESS
(GAP MODEL)
• BOTKIN (1993) DEFINES TWO GAP MODELS
• INTRINSIC MORTALITY
• OCCUR IN FAVORABLE ENVIRONMENT  WITH/OUT
COMPETITION
• MIGHT INCLUDE NON EPIDEMIC DISEASE, LIGHTNING,
WIND THROW

• GROWTH DEPENDENT MORTALITY
• OFTEN DUE TO COMPETITION FOR RESOURCES
• ASSUMES: SLOW GROWING  LIKELY TO DIE
• OTHER AGENTS: INSECT OR DISEASE, WIND EVENTS OR
ABIOTIC PERTURBATIONS
TREE MORTALITY PROCESS
(GAP MODEL)
• KEANE ET. AL. 2001 DISCUSSES AN ADDITIONAL
CLASS  EXOGENOUS MORTALITY
• WHEN EXTERNAL FACTOR SWEEPS AND
KILLS A PATCH OR STAND OR ALL TREES
• AGENTS: FIRE, PEST OUTBREAKS, SEVERE
WIND
• REPRESENTED AS THE SEVERITY OF THE
INTRINSIC MORTALITY MODEL
• AS AN AGE INDEPENDENT MORTALITY ROUTINE( MAX. AGE
DEPENDENT)
• ASSUMES:
• CHANCE PLAYS MAJOR ROLE (MORTALITY ~ RANDOM,
LOCALIZED)
• CONSTANT PROBABILITY OF DEATH

• STANDS END WITH 1% OR 2% OF ALL TREES SURVIVING
• COMMON EQUATION

• ANNUAL MORTALITY RATE ~ SPECIES~ EACH HAS DIFF.
MAX AGE
GROWTH DEPENDENT
MORTALITY
MODEL

• SLOWED GROWTH RATE  ENHANCED PROB. OF MORTALITY

• ASSUMES: RATE BELOW THRESHOLD  NEGATIVE CARBON
BALANCE

• 1 YEAR OLD STEM WITH STEM RADIAL GROWTH ,0.1 MM/YR 
AFTER 10 YEARS 30% OF VULNERABLE TREES DID NOT SURVIVE
- BOTKIN 1972
• TREEHIGH MOR. RATE ~ ONLY 3 OR MORE CONSECUTIVE YEAR
OF LOW THRESHOLD GROWTH RATE
GROWTH DEPENDENT
MORTALITY
MODEL

• (FORSKA APPROACH, PRENTICE et. al., 1993) ANNUAL
MORTALITY RATE (XM) BASED ON RELATIVE GROWTH
EFFICIENCY (EREL)

• WHERE, U0= INTRINSIC MORTALITY RATE
U1= SPECIES SPECIFIC MORTALITY RATE
q = THRESHOLD VALUE FOR VIGOR INDEX
r = SPECIES SPECIFIC SHAPE PARAMETER
GROWTH DEPENDENT
MORTALITY
MODEL
• SORTIE APPROACH PALACE et. al., 1993 CALCULATES
USING RING WIDTH OF LIVE AND DEAD INDIVIDUALS

• WHERE, PM= PROB. OF MORTALITY
d= AVERAGE RIG WIDTH (mm)
u and v= SPECIES SPECIFIC CONSTANTS
EXOGENOUS MORTALITY MODEL
• RESULT OF LONG TIME SIMULATED PERIODS
• NOT BEEN IMPLEMENTED IN GAP MODELS DUE TO
1.

IT WAS PREVIOUSLY THOUGHT TO BE UNIMPORTANT TO
THE DYNAMICS OF THE SIMULATED ECOSYSTEM,

2.

THERE WAS LITTLE KNOWN ABOUT THE SPATIAL
MECHANISMS OF THE DISTURBANCE PROCESS

3.

IT WAS DIFFICULT TO SIMULATE BECAUSE OF EXTENSIVE
COMPUTER REQUIREMENTS

4. THERE WERE VERY LITTLE DATA FOR PARAMETERIZATION
AND
5.

THE SIMULATED VARIABLES COULD NOT BE RELATED TO
EXOGENOUS DISTURBANCE EFFECTS
EXOGENOUS MORTALITY MODEL
• KEANE ET. AL., (1990, 1996) SIMULATED FIRE MORTALITY
FROM EMPIRICALLY DERIVED STOCHASTIC EQUATION

where, Pfire= PROB. OF FIRE CAUSED MORTALITY
DBH= TREE DIAMETER
bt= SPECIES SPECIFIC BARK THICKNESS
COEFFICIENTS(cm)
CK= PERCENTAGE CROWN VOLUME
USES OF MODELS
• (FRIDMAN et.al., 2001)THERE ARE AT LEAST THREE
PRINCIPLES FOR APPLICATION OF THE FUNCTIONS:
• DETERMINISTIC,
• MORTALITY IS EVENLY DISTRIBUTED AMONG PLOTS AND
TREES

• STOCHASTIC
• RANDOM SIMULATION DETERMINES WHAT PLOTS AND
TREES WILL BE AFFECTED.

• STOCHASTIC WITH MONTE CARLO SIMULATION
• CAN BE USED TO DERIVE EXPECTED VALUES AND
DISTRIBUTIONS
• GENERAL MODEL USED IN 3 STEP MODELOING OF
SWEDISH FOREST WAS

• THE PERFORMANCE OF THE FUNCTIONS DEVELOPED
INDICATES FAIR PREDICTABILITY IN GENERAL. THUS,
APPLICATION OF THE MODELS IN SWEDISH LONG-TERM
PLANNING SYSTEMS CAN BE RECOMMENDED.
USES OF MODELS
• IN ALBERTA MIXEDWOOD FORESTS FOR ASPEN, WHITE SPRUCE AND
LODGEPOLE PINE, THIS MODEL WAS USED:

WHICH IS THE LOGARITHMIC EXTENSION OF THE EQUATION

WHERE ß = (ß 0, ß 1, ..., ß K ) ARE UNKNOWN PARAMETERS,
YI IS THE RESPONSE VARIABLE FOR THE ITH OBSERVATION,
XI IS THE VECTOR OF THE EXPLANATORY VARIABLE FOR THE ITH
OBSERVATION,
USES OF MODELS
• FINDINGS IN ALBERTA MIXEDWOOD FORESTS
FOR ASPEN, WHITE SPRUCE AND LODGEPOLE
PINE WAS
• ASPEN IS THE SHORTEST LIVED SPECIES
WITH THE LOWEST SURVIVAL PROBABILITY,
WHILE WHITE SPRUCE IS THE LONGEST
LIVED SPECIES WITH THE HIGHEST OVERALL
SURVIVAL PROBABILITY
USES OF MODELS
• HOOD ET. AL. 2007 USES THE FOLLOWING REGRESSION
MODEL TO ESTIMATE DELAYED MORTALITY ON CONIFERS
FOLLOWING FIRE
USES OF MODELS
• THE STUDY FOUND THAT PERCENT CROWN LENGTH KILLED AND THE
NUMBER OF QUADRANTS WITH DEAD CAMBIUM SAMPLES TO BE THE
MOST IMPORTANT VARIABLES FOR PREDICTING POST-FIRE
MORTALITY FOR MIXED CONIFER SPECIES IN CALIFORNIA

• FOR FIRES WHERE ASSESSMENTS HAVE BEEN MADE OVER A
LONGER PERIOD OF TIME, THE MAJORITY OF THE MORTALITY
OCCURRED WITHIN THREE YEARS POST-FIRE. DELAYED MORTALITY,
IN TERMS OF CROWN DEATH, MAY TAKE SEVERAL YEARS TO OCCUR
FOR TREES WITH FATAL LEVELS OF CAMBIUM KILL.

• FOR WHITE FIR, CAMBIUM KILL WAS A MORE IMPORTANT VARIABLE IN
THE THREE-YEAR MODEL COMPARED TO THE TWO-YEAR MODEL.
CONCLUSION AND
RECOMMENDATION
• DAHLMAN (1985) ~ GAP MODELS ARE POTENTIALLY POWERFUL
TOOLS FOR SIMULATING COMMUNITY-LEVEL RESPONSES TO CO2
INCREASE, BUT ONLY IF PROPERLY PARAMETERIZED AND
VALIDATED. OTHERWISE, ~ JUST EXPLORATORY EXERCISES.

• A QUESTION WHETHER APPROPRIATE MORTALITY DATA ARE
AVAILABLE OR CAN EVER BE COLLECTED TO EXTENSIVELY
VALIDATE THE REPRESENTATION OF PLANT DEATH IN GAP MODELS,
ESPECIALLY SPATIAL GAP MODELS (DEUTSCHMAN ET AL., 1999).
CONCLUSION AND
RECOMMENDATION
• STOCHASTIC MORTALITY FUNCTIONS MUST BE DEVELOPED THAT
USE PROCESS BASED, MECHANISTIC RELATIONSHIPS AS
PREDICTIVE VARIABLES.
E.G., FIRE IGNITION PROBABILITIES  CLIMATE-BASED
VARIABLES.

• RESEARCH - EXPANDED - ATTEMPTS TO MECHANISTICALLY
UNDERSTAND THE RELATIONSHIP BETWEEN ECO-PHYSIOLOGICAL
PROCESSES AND PLANT MORTALITY.

• A COMPREHENSIVE FIELD DATABASE IS NEEDED TO DESIGN,
PARAMETERIZE, AND VALIDATE GAP MODEL MORTALITY ALGORITHMS
22

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Probalility and models of tree mortality advance silviculture

  • 1.
  • 2. A Term Paper Presentation On PROBABILITY AND MODELS OF TREE MORTALITY Jeetendra Gautam MSc Forestry (General Forestry) Roll No. 14
  • 3. TREE MORTALITY • PLANT DEATH IS A COMPLEX PROCESS, INFLUENCED BY • PHYSIOLOGY, • ENVIRONMENT • SUCCESSIONAL DEVELOPMENT, • AGE, AND, • CHANCE - (HARCOMBE, 1987; FRANKLIN ET AL., 1987)
  • 4. TREE MORTALITY • GENERAL CAUSES • LACK OF SUFFICIENT RESOURCES TO RESTRAIN STRESS, INJURIES OR SUSTAIN LIFE, OR KILLED BY EXTERNAL FACTOR (WARING, 1987) • NEGATIVE CARBON BALANCE (RESPIRATION > PHOTOSYNTHESIS) • INSECT INFESTATION • PROLONGED DROUGHT UNDER LOW LIGHT CONDITION
  • 5. TREE MORTALITY PROCESS (GAP MODEL) • BOTKIN (1993) DEFINES TWO GAP MODELS • INTRINSIC MORTALITY • OCCUR IN FAVORABLE ENVIRONMENT  WITH/OUT COMPETITION • MIGHT INCLUDE NON EPIDEMIC DISEASE, LIGHTNING, WIND THROW • GROWTH DEPENDENT MORTALITY • OFTEN DUE TO COMPETITION FOR RESOURCES • ASSUMES: SLOW GROWING  LIKELY TO DIE • OTHER AGENTS: INSECT OR DISEASE, WIND EVENTS OR ABIOTIC PERTURBATIONS
  • 6. TREE MORTALITY PROCESS (GAP MODEL) • KEANE ET. AL. 2001 DISCUSSES AN ADDITIONAL CLASS  EXOGENOUS MORTALITY • WHEN EXTERNAL FACTOR SWEEPS AND KILLS A PATCH OR STAND OR ALL TREES • AGENTS: FIRE, PEST OUTBREAKS, SEVERE WIND • REPRESENTED AS THE SEVERITY OF THE
  • 7. INTRINSIC MORTALITY MODEL • AS AN AGE INDEPENDENT MORTALITY ROUTINE( MAX. AGE DEPENDENT) • ASSUMES: • CHANCE PLAYS MAJOR ROLE (MORTALITY ~ RANDOM, LOCALIZED) • CONSTANT PROBABILITY OF DEATH • STANDS END WITH 1% OR 2% OF ALL TREES SURVIVING • COMMON EQUATION • ANNUAL MORTALITY RATE ~ SPECIES~ EACH HAS DIFF. MAX AGE
  • 8. GROWTH DEPENDENT MORTALITY MODEL • SLOWED GROWTH RATE  ENHANCED PROB. OF MORTALITY • ASSUMES: RATE BELOW THRESHOLD  NEGATIVE CARBON BALANCE • 1 YEAR OLD STEM WITH STEM RADIAL GROWTH ,0.1 MM/YR  AFTER 10 YEARS 30% OF VULNERABLE TREES DID NOT SURVIVE - BOTKIN 1972 • TREEHIGH MOR. RATE ~ ONLY 3 OR MORE CONSECUTIVE YEAR OF LOW THRESHOLD GROWTH RATE
  • 9. GROWTH DEPENDENT MORTALITY MODEL • (FORSKA APPROACH, PRENTICE et. al., 1993) ANNUAL MORTALITY RATE (XM) BASED ON RELATIVE GROWTH EFFICIENCY (EREL) • WHERE, U0= INTRINSIC MORTALITY RATE U1= SPECIES SPECIFIC MORTALITY RATE q = THRESHOLD VALUE FOR VIGOR INDEX r = SPECIES SPECIFIC SHAPE PARAMETER
  • 10. GROWTH DEPENDENT MORTALITY MODEL • SORTIE APPROACH PALACE et. al., 1993 CALCULATES USING RING WIDTH OF LIVE AND DEAD INDIVIDUALS • WHERE, PM= PROB. OF MORTALITY d= AVERAGE RIG WIDTH (mm) u and v= SPECIES SPECIFIC CONSTANTS
  • 11. EXOGENOUS MORTALITY MODEL • RESULT OF LONG TIME SIMULATED PERIODS • NOT BEEN IMPLEMENTED IN GAP MODELS DUE TO 1. IT WAS PREVIOUSLY THOUGHT TO BE UNIMPORTANT TO THE DYNAMICS OF THE SIMULATED ECOSYSTEM, 2. THERE WAS LITTLE KNOWN ABOUT THE SPATIAL MECHANISMS OF THE DISTURBANCE PROCESS 3. IT WAS DIFFICULT TO SIMULATE BECAUSE OF EXTENSIVE COMPUTER REQUIREMENTS 4. THERE WERE VERY LITTLE DATA FOR PARAMETERIZATION AND 5. THE SIMULATED VARIABLES COULD NOT BE RELATED TO EXOGENOUS DISTURBANCE EFFECTS
  • 12. EXOGENOUS MORTALITY MODEL • KEANE ET. AL., (1990, 1996) SIMULATED FIRE MORTALITY FROM EMPIRICALLY DERIVED STOCHASTIC EQUATION where, Pfire= PROB. OF FIRE CAUSED MORTALITY DBH= TREE DIAMETER bt= SPECIES SPECIFIC BARK THICKNESS COEFFICIENTS(cm) CK= PERCENTAGE CROWN VOLUME
  • 13. USES OF MODELS • (FRIDMAN et.al., 2001)THERE ARE AT LEAST THREE PRINCIPLES FOR APPLICATION OF THE FUNCTIONS: • DETERMINISTIC, • MORTALITY IS EVENLY DISTRIBUTED AMONG PLOTS AND TREES • STOCHASTIC • RANDOM SIMULATION DETERMINES WHAT PLOTS AND TREES WILL BE AFFECTED. • STOCHASTIC WITH MONTE CARLO SIMULATION • CAN BE USED TO DERIVE EXPECTED VALUES AND DISTRIBUTIONS
  • 14. • GENERAL MODEL USED IN 3 STEP MODELOING OF SWEDISH FOREST WAS • THE PERFORMANCE OF THE FUNCTIONS DEVELOPED INDICATES FAIR PREDICTABILITY IN GENERAL. THUS, APPLICATION OF THE MODELS IN SWEDISH LONG-TERM PLANNING SYSTEMS CAN BE RECOMMENDED.
  • 15. USES OF MODELS • IN ALBERTA MIXEDWOOD FORESTS FOR ASPEN, WHITE SPRUCE AND LODGEPOLE PINE, THIS MODEL WAS USED: WHICH IS THE LOGARITHMIC EXTENSION OF THE EQUATION WHERE ß = (ß 0, ß 1, ..., ß K ) ARE UNKNOWN PARAMETERS, YI IS THE RESPONSE VARIABLE FOR THE ITH OBSERVATION, XI IS THE VECTOR OF THE EXPLANATORY VARIABLE FOR THE ITH OBSERVATION,
  • 16. USES OF MODELS • FINDINGS IN ALBERTA MIXEDWOOD FORESTS FOR ASPEN, WHITE SPRUCE AND LODGEPOLE PINE WAS • ASPEN IS THE SHORTEST LIVED SPECIES WITH THE LOWEST SURVIVAL PROBABILITY, WHILE WHITE SPRUCE IS THE LONGEST LIVED SPECIES WITH THE HIGHEST OVERALL SURVIVAL PROBABILITY
  • 17. USES OF MODELS • HOOD ET. AL. 2007 USES THE FOLLOWING REGRESSION MODEL TO ESTIMATE DELAYED MORTALITY ON CONIFERS FOLLOWING FIRE
  • 18. USES OF MODELS • THE STUDY FOUND THAT PERCENT CROWN LENGTH KILLED AND THE NUMBER OF QUADRANTS WITH DEAD CAMBIUM SAMPLES TO BE THE MOST IMPORTANT VARIABLES FOR PREDICTING POST-FIRE MORTALITY FOR MIXED CONIFER SPECIES IN CALIFORNIA • FOR FIRES WHERE ASSESSMENTS HAVE BEEN MADE OVER A LONGER PERIOD OF TIME, THE MAJORITY OF THE MORTALITY OCCURRED WITHIN THREE YEARS POST-FIRE. DELAYED MORTALITY, IN TERMS OF CROWN DEATH, MAY TAKE SEVERAL YEARS TO OCCUR FOR TREES WITH FATAL LEVELS OF CAMBIUM KILL. • FOR WHITE FIR, CAMBIUM KILL WAS A MORE IMPORTANT VARIABLE IN THE THREE-YEAR MODEL COMPARED TO THE TWO-YEAR MODEL.
  • 19. CONCLUSION AND RECOMMENDATION • DAHLMAN (1985) ~ GAP MODELS ARE POTENTIALLY POWERFUL TOOLS FOR SIMULATING COMMUNITY-LEVEL RESPONSES TO CO2 INCREASE, BUT ONLY IF PROPERLY PARAMETERIZED AND VALIDATED. OTHERWISE, ~ JUST EXPLORATORY EXERCISES. • A QUESTION WHETHER APPROPRIATE MORTALITY DATA ARE AVAILABLE OR CAN EVER BE COLLECTED TO EXTENSIVELY VALIDATE THE REPRESENTATION OF PLANT DEATH IN GAP MODELS, ESPECIALLY SPATIAL GAP MODELS (DEUTSCHMAN ET AL., 1999).
  • 20. CONCLUSION AND RECOMMENDATION • STOCHASTIC MORTALITY FUNCTIONS MUST BE DEVELOPED THAT USE PROCESS BASED, MECHANISTIC RELATIONSHIPS AS PREDICTIVE VARIABLES. E.G., FIRE IGNITION PROBABILITIES  CLIMATE-BASED VARIABLES. • RESEARCH - EXPANDED - ATTEMPTS TO MECHANISTICALLY UNDERSTAND THE RELATIONSHIP BETWEEN ECO-PHYSIOLOGICAL PROCESSES AND PLANT MORTALITY. • A COMPREHENSIVE FIELD DATABASE IS NEEDED TO DESIGN, PARAMETERIZE, AND VALIDATE GAP MODEL MORTALITY ALGORITHMS
  • 21.
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