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( 2013 Kristin McCully & Myra Finkelstein)
Lead Poisoning of Laysan Albatrosses at Midway Atoll:
From Ecological Model to Conservation Action
Objectives• Learn how to use structured population
models to assess and protect populations
• Appreciate the importance and uses of mathematical models in ecology and conservation
2
3
Outline• Introduction to case study:
Laysan albatrosses at Midway Atoll• Introduction to mathematical models• Structured population models– Assumptions– Model structure
• Mathematical model: Laysan albatrosses at Midway Atoll
• Importance of models to ecology and conservation
4
Outline• Introduction to case study:
Laysan albatrosses at Midway Atoll• Introduction to mathematical models• Structured population models– Assumptions– Model structure
• Mathematical model: Laysan albatrosses at Midway Atoll
• Importance of models to ecology and conservation
5
Case Study: Lead Poisoning of Laysan
Albatrosses at Midway Atoll
Research by Myra Finkelstein
6
Threats to Albatross Populations
• Longline fishing bycatch• Introduced mammals• Marine debris• Lead poisoning
Brenda Zaun/USFWS
7
Where is Midway Atoll?
Papahānaumokuākea Marine National Monument
8
Midway Atoll
Keith Castellano
9
Wisdom: World’s Oldest- Known Wild Bird
Pete Leary/USFWSCheck her out on Facebook: https://www.facebook.com/wisdomthealbatross
10
“Droopwing” syndrome
Peripheral Neuropathy
National Institutes of Health
11
ProblemLead poisoning of Laysan chicks near buildings- Sileo and Fefer (1987) J Wildlife Disease- Sileo et al. (1990) J Wildlife Disease- Work et al (1996) Env Sci & Toxicology
Military base → many sources of leadNeed evidence to link source with poisoning
Away from bldgsn =15
Near bldgsn = 21
Lead
(μg/
dL w
hole
blo
od)
0
200
400
600
800
1000
1200
1400
1600
> 80% of chicks near bldgs. >100 ug/dL
Droopwing or other symptoms
Step 1: Quantify Lead Exposure
Finkelstein et al. (2003) Environmental Science & Technology
13
• Fingerprint source using isotope analysisIsotopes = variants of chemical element with
different numbers of neutrons
• Lead exists as four isotopes:204Pb, 206Pb, 207Pb, 208Pb
• Lead isotope ratios in organism reflects lead isotope ratios in exposure source
Step 2: Identify Source of Lead
3 isotopes of hydrogen:
Image: Dirk Hünniger (Wikimedia)
14→ nest soil is not source of poisoning
Finkelstein et al. (2003) Environmental Science & Technology
Near Buildings: r 2 = 0.363p = 0.152
Step 2: Identify Source of LeadCh
ick
Bloo
d 20
7 Pb/
206 P
b
Nest Soil 207Pb/206Pb
Away from Buildings: r 2 = 0.224p = 0.074
15→ lead-based paint is source of poisoning
Finkelstein et al. (2003) Environmental Science & Technology
r 2 = 0.853p = 0.003
Step 2: Identify Source of LeadCh
ick
Bloo
d 20
7 Pb/
206 P
b
Nest Paint Chips 207Pb/206Pb
16
Step 3: Show Population Consequences
Mathematical Model
17
Outline• Introduction to case study:
Laysan albatrosses at Midway Atoll• Introduction to mathematical models• Structured population models– Assumptions– Model structure
• Mathematical model: Laysan albatrosses at Midway Atoll
• Importance of models to ecology and conservation
18
Discussion: What is a Model?• Simplified representation of object or process• Requires assumptions
USGS
Essentially, all models are wrong, but some are useful.
- George Box, 1987
19
Mathematical Population Models
N
time
Exponential Growth Logistic Growth
N
timeMalthus (1798) Verhulst (1838)
20
Steps to a Mathematical Model1. Ask research questions2. Make assumptions3. Develop conceptual model4. Formulate mathematical model5. Assign values to parameters6. Use model to answer questions
Based on: Soetaert & Herman 2008 A Practical Guide to Ecological Modeling (Fig. 1.7)
21
Outline• Introduction to case study:
Laysan albatrosses at Midway Atoll• Introduction to mathematical models• Structured population models– Assumptions– Model structure
• Mathematical model: Laysan albatrosses at Midway Atoll
• Importance of models to ecology and conservation
22
1. Ask Research Questions• Is the population growing? • Which stage(s) and demographic processes
should managers focus monitoring and conservation effort on?
• How might threats or management actions impact the population?
23
2. Make Assumptions• Constant environment (Deterministic)• Unlimited resources
Constant birth & death rates• No immigration or emigration• No time lags• All individuals are identical
N
time
Exponential Growth
From Gotelli (2001) A Primer of Ecology
24
2. Make Assumptions• Constant environment (Deterministic)• Unlimited resources
Constant birth & death rates• No immigration or emigration• No time lags• All individuals are identical• All individuals in each stage are identical
From Gotelli (2001) A Primer of Ecology
vital rates
→ structured population model
Complex Life Histories
Transition Matrix
Structured Population Model
25
n1 P1 F2 F3 F4 F5 F6 n1
n2 G1 P2 0 0 0 0 n2
n3 =
0 G2 P3 0 0 0 n3
n4 0 0 G3 P4 0 0 n4
n5 0 0 0 G4 P5 0 n5
n6 0 0 0 0 G5 P6 n6
tt+1
• Age • Size • Sex • Developmental stageStructured by
Transition Matrix
Structured Population Model
26
n1 P1 F2 F3 F4 F5 F6 n1
n2 G1 P2 0 0 0 0 n2
n3 =
0 G2 P3 0 0 0 n3
n4 0 0 G3 P4 0 0 n4
n5 0 0 0 G4 P5 0 n5
n6 0 0 0 0 G5 P6 n6
t
1 2 3 4 5 6
1
2
=3
4
5
6t+1
1 2 3 4 5 6
1 P1
2 P2
3 P3
4 P4
5 P5
6 P6
1 2 3 4 5 6
1
2 G1
3 G2
4 G3
5 G4
6 G5
1 2 3 4 5 6
1 F2 F3 F4 F5 F6
2
3
4
5
6
0 0 0 0
=0 0 0 0
0 0 0 0
0 0 0 0
0 0 0 0
P = Persistence G = Growth F = FecundityFROM
TO
27
PJ
GL GJ
PA
P = Persistence
Simpler Example: Lionfish (Pterois volitans)
NOAA SE Fisheries Science Center
Todd Gardner
L J A
Larvae Juveniles Adults
FAG = Growth
F = Fecundity
Morris et al. (2011) Biological Invasions
Time step: 1 month
28
Larvae Juveniles Adults
Larvae
Juveniles
Adults
ToStage
From Stage
GL
GJ
PL
PA
FA
Simpler Example: Lionfish (Pterois volitans)
0
0
0
0
PL
GL GJ
PA
P = Persistence L J AFAG = Growth
F = Fecundity
Time step: 1 month
Seeds
Immature Plants
Mature Plants
F3
G1P1
P2
P3G2
F = Fecundity G = Growth P = PersistenceEx2: Life Cycle Diagram → Transition Matrix
Ex2: Life Cycle Diagram → Transition Matrix
Stage-Based ModelF = Fecundity G = Growth P = Persistence
Seeds Immature Plants
Mature Plants
Seeds
Immature Plants
MaturePlants
ToStage
From Stage
P1
G1
0
F
0
P3
0
P2
G2
Seeds
Immature Plants
Mature Plants
F
G1P1
P2
P3G2
Ex3: Transition Matrix → Life Cycle Diagram
0 0 F3 F4
G1 P2 0 00 G2 P3 00 0 G3 P4
1 2 3 4P2 P3 P4
G3G2G1
F4F3
32
Outline• Introduction to case study:
Laysan albatrosses at Midway Atoll• Introduction to mathematical models• Structured population models– Assumptions– Model structure
• Mathematical model: Laysan albatrosses at Midway Atoll
• Importance of models to ecology and conservation
33
Steps to a Mathematical Model1. Ask research questions2. Make assumptions3. Develop conceptual model4. Formulate mathematical model5. Assign values to parameters6. Use model to answer questions
Based on: Soetaert & Herman 2008 A Practical Guide to Ecological Modeling (Fig. 1.7)
34
1. Determine Research Questions
• How does lead poisoning of chicks affect population growth?
• Will lead remediation help protect the population?
Finkelstein et al. 2009 Animal Conservation
35
2. Make Assumptions• Constant environment (Deterministic)• Unlimited resources
Constant birth & death rates• All individuals are identical• No immigration or emigration• No time lags• All individuals in each stage are identical
From Gotelli (2001) A Primer of Ecology
vital rates
→ structured population model
Complex Life Histories
36
3. Develop Conceptual ModelVideo: Albatross Life History
3. Develop Conceptual Model
37
E 5421 3 6 7 8
Pete Leary/USFWS
SubadultImmatureJuvenileEggs
Chicks
AdultsGina
Ruttle
Time step: 1 year
4. Formulate Math Model
38
E 5421 3 6 7 8
F
Gone Gj Gj Gj Gi Gi Gsa Pa
Gsa
G = GrowthP = PersistenceF = Fecundity
4. Formulate Math Model
39
F
E 1 2 3 4 5 6 7 8E
1
2
3
4
5
6
7
8
E 5421 3 6 7 8Gone Gj Gj Gj Gi Gi Gsa Pa
Gsa
4. Formulate Math Model
40
F
E 1 2 3 4 5 6 7 8E F
1 Gone
2 Gj
3 Gj
4 Gj
5 Gi
6 Gi
7 Gsa
8 Gsa Pa
E 5421 3 6 7 8Gone Gj Gj Gj Gi Gi Gsa Pa
Gsa
4. Formulate Math Model
41
F
E 1 2 3 4 5 6 7 8E 0 0 0 0 0 0 0 0 F
1 Gone 0 0 0 0 0 0 0 0
2 0 Gj 0 0 0 0 0 0 0
3 0 0 Gj 0 0 0 0 0 0
4 0 0 0 Gj 0 0 0 0 0
5 0 0 0 0 Gi 0 0 0 0
6 0 0 0 0 0 Gi 0 0 0
7 0 0 0 0 0 0 Gsa 0 0
8 0 0 0 0 0 0 0 Gsa Pa
E 5421 3 6 7 8Gone Gj Gj Gj Gi Gi Gsa Pa
Gsa
42
5. Assign Values to Parameters
Pete Leary/USFWS
Andrew Derocher (IUCN)
NOAA SEFSC
NOAA
5. Assign Values to Parameters
43
E 5421 3 6 7 8
F
Gone Gj Gj Gj Gi Gi Gsa Pa
Gsa
G = GrowthP = PersistenceF = Fecundity
0.482 0.84 0.84 0.84 0.906 0.906 0.9450.916
0.945
= 0.366
5. Assign Values to Parameters
44
F
E 1 2 3 4 5 6 7 8E 0 0 0 0 0 0 0 0 F
1 Gone 0 0 0 0 0 0 0 0
2 0 Gj 0 0 0 0 0 0 0
3 0 0 Gj 0 0 0 0 0 0
4 0 0 0 Gj 0 0 0 0 0
5 0 0 0 0 Gi 0 0 0 0
6 0 0 0 0 0 Gi 0 0 0
7 0 0 0 0 0 0 Gsa 0 0
8 0 0 0 0 0 0 0 Gsa Pa
E 5421 3 6 7 8Gone Gj Gj Gj Gi Gi Gsa Pa
Gsa
5. Assign Values to Parameters
45
F
E 1 2 3 4 5 6 7 8E 0 0 0 0 0 0 0 0 0.3661 0.482 0 0 0 0 0 0 0 02 0 0.840 0 0 0 0 0 0 03 0 0 0.840 0 0 0 0 0 04 0 0 0 0.840 0 0 0 0 05 0 0 0 0 0.906 0 0 0 06 0 0 0 0 0 0.906 0 0 07 0 0 0 0 0 0 0.945 0 08 0 0 0 0 0 0 0 0.945 0.916
E 5421 3 6 7 8Gone Gj Gj Gj Gi Gi Gsa Pa
Gsa
46
Steps to a Mathematical Model1. Ask research questions2. Make assumptions3. Develop conceptual model4. Formulate mathematical model5. Assign values to parameters6. Use model to answer questions
Based on: Soetaert & Herman 2008 A Practical Guide to Ecological Modeling (Fig. 1.7)
6. Use Model to Answer QuestionsA. Is the population growing?
B. Which vital rate(s) should managers focus monitoring and conservation effort on?
C. Which stage(s) should managers focus monitoring and conservation effort on?
D. Which stage(s) contain most of the population?
E. How might management actions impact population?
Transition Matrix
48
n1 P1 F2 F3 F4 F5 F6 n1
n2 G1 P2 0 0 0 0 n2
n3 =
0 G2 P3 0 0 0 n3
n4 0 0 G3 P4 0 0 n4
n5 0 0 0 G4 P5 0 n5
n6 0 0 0 0 G5 P6 n6
tt+1
A. Is the population growing?
Math: multiplication of transition matrix & initial population vectorPopTools: Matrix Tools → Matrix Projection
49
Popu
latio
n Si
zeA. Is the population growing?
Time
50
A. Is the population growing?
Population Growth Rate (λ) • λ = Nt+1 / Nt
• If λ = 1 , Nt+1 = Nt → population size
• If λ > 1, Nt+1 > Nt → population size
• If λ < 1, Nt+1 < Nt → population size
Math: dominant eigenvalue of transition matrixPopTools: = DomEig(TM)
is constant
is increasing
is decreasing
51
Time (Months)
# Li
onfis
h
Morris et al. (2011) Biological Invasions
λ = 1.134
A. Is the population growing?
0 4 8 12 16 20 24 28 32 36 40 44 480
250,000
500,000
750,000
1,000,000
1,250,000
1,500,000
1,750,000
Time (years)
Num
ber o
f Ind
ivid
uals
in P
opu-
latio
n
A. Is the population growing? [After many years with these vital rates]
With initial population vector based on stable stage distributionWith initial population vector not based on stable stage distribution
λ = 0.995
Elasticity= Effect that a change in each one of the vital
rates has on population growth rate λ • Used to determine which vital rate changes
the population growth rate λ most
B. Which vital rate(s) should managers focus monitoring and conservation effort on?
Math: , where <> denotes the scalar productPopTools: Matrix Tools → Elasticity
B. Which vital rate(s) should managers focus monitoring and conservation effort on?
Elasticity:
Corresponds to Pa
0 0 0 0 0 0 0 0 0.048720.04872 0 0 0 0 0 0 0 0
0 0.04872 0 0 0 0 0 0 00 0 0.04872 0 0 0 0 0 00 0 0 0.04872 0 0 0 0 00 0 0 0 0.04872 0 0 0 00 0 0 0 0 0.04872 0 0 00 0 0 0 0 0 0.04872 0 00 0 0 0 0 0 0 0.04872 0.56151
00.10.20.30.40.50.6
Elas
ticity
Vital Rate
Elasticity
Gone 0.04872Gj 0.04872Gi 0.04872Gsa 0.04872Ga 0.56152F 0.04872
B. Which vital rate(s) should managers focus monitoring and conservation effort on?
Gone Gj Gi Gsa Ga F
Reproductive Value= expected number of offspring that remain to be born
to each individual of a stage
• Essentially which stages are most valuable for future population growth
• Math: left eigenvector of transition matrix• PopTools: Matrix Tools → Age Distribution
v(x) =
# of offspring produced by individuals of age x or older (discounted by likelihood of surviving to reproduce)
# of individuals of age x
C. Which stage(s) should managers focus monitoring and conservation effort on?
Egg 1 2 3 4 5 6 7 80.000.020.040.060.080.100.120.140.160.18
Stage
Repr
oduc
tive
Valu
e
Stage Reproductive Value
Egg 0.035
1 0.0718
2 0.0851
3 0.1009
4 0.1195
5 0.1314
6 0.1443
7 0.1520
8 0.1601
C. Which stage(s) should managers focus monitoring and conservation effort on?
Reproductive Value• Humans?
(based on Daly and Wilson 1988 Homicide)
• To maximize sustainable harvest yield, harvest stages with reproductive value
• To maximize impact of restoration, transplant individuals with reproductive value
• Natural selection will act most strongly on stages with reproductive value
low
high
high
RV
Age
Gotelli 2008 A Primer of Ecology, Caswell 1980 Ecology
D. Which stage(s) contain most of the population?
0 3 6 9 12 15 18 21 24 27 300
200,000
400,000
600,000
800,000
1,000,000
1,200,000
Stage E
Stage 1
Stage 2
Stage 3
Stage 4
Stage 5
Stage 6
Stage 7
Stage 8Time (years)
Num
ber o
f Ind
ivid
uals
in E
ach
Stag
e
PopTools: Matrix Tools → Matrix Projection
0 3 6 9 12 15 18 21 24 27 300
0.1
0.2
0.3
0.4
0.5
0.6
0.7
0.8
Stage EStage 1Stage 2Stage 3Stage 4Stage 5Stage 6Stage 7Stage 8Time (years)
Prop
ortio
n of
Indi
vidu
als
in
Each
Sta
geD. Which stage(s) contain most of the population?
[After many years with these vital rates]
Stable Stage Distribution
Stable Stage Distribution= constant proportion of individuals in each stage
(derived from constant vital rates)
• Math: Right eigenvector of transition matrix• PopTools: Matrix Tools → Age Distribution
w(x) = # of individuals of age x in SSD
total # of individuals
D. Which stage(s) contain most of the population? [After many years with these vital rates]
Stage Stable Stage Distribution
Egg 0.1671 0.0812 0.0683 0.0584 0.0495 0.0446 0.0417 0.0388 0.455
D. Which stage(s) contain most of the population? [After many years with these vital rates]
Egg
1 2 3 4 5 6 7 80
0.1
0.2
0.3
0.4
0.5
Stage
Stab
le S
tage
Dis
trib
ution
63
• How does lead poisoning of chicks affect population growth?
• Will lead remediation help protect the population?
E. How might management actions impact population?
SIMULATION – What if . . .
64
How does lead poisoning of chicks affect population growth?
E 5421 3 6 7 8
65Finkelstein et al. 2009 Animal Conservation
How does lead poisoning of chicks affect population growth?
3.5%7%14%
Lead-induced chick mortality
0%
66
starting in 21 years
Will lead remediation help protect the population?
Lead Remediation
1% Reduction in Adult Bycatch Mortality
Finkelstein et al. 2009 Animal Conservation
No Action
67
Result: Conservation Action
For more info: http://www.fws.gov/midway/lpa.html
Pete Leary / USFWS USFWS
68
Outline• Introduction to case study:
Laysan albatrosses at Midway Atoll• Introduction to mathematical models• Structured population models– Assumptions– Model structure
• Mathematical model: Laysan albatrosses at Midway Atoll
• Importance of models to ecology and conservation
69
Discussion: How do we use and why do we need mathematical models in ecology and
conservation?
Objectives• Learn how scientists use structured population
models to assess and protect populations
• Appreciate the importance and uses of mathematical models in ecology and conservation
70
71
Lab: Cooperative Activity Using PopTools
Dwayne Meadows, NOAA/NMFS/OPR
John Snow
Chris Muenzer
Using PopTools Add-on to MS Excelhttp://www.poptools.org/
Resources• Crouse, D. T., Crowder, L. B., & Caswell, H. (1987). A stage-
based population model for loggerhead sea turtles and implications for conservation. Ecology 68(5):1412-1423. http://dx.doi.org/10.2307/1939225
• Finkelstein, M. E., Doak, D. F., Nakagawa, M., Sievert, P. R., & Klavitter, J. (2010). Assessment of demographic risk factors and management priorities: impacts on juveniles substantially affect population viability of a long-lived seabird. Animal Conservation, 13(2), 148-156. http://dx.doi.org/10.1111/j.1469-1795.2009.00311.x
• Gotelli, N. (Various) A Primer of Ecology. Chapter 3: Age-Structured Population Growth. Sinauer.
74
Questions?
Prettier photo?