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![Page 1: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/1.jpg)
Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges
for the next century
Nicholas J. GotelliDepartment of BiologyUniversity of VermontBurlington, VT 05405
U.S.A.
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0
50
100
150
200
1900 1920 1940 1960 1980 2000
Year
Ter
agra
ms
of
Nit
rog
en
Fertilizer NOx
Total anthropogenic N fixed
Natural range
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Effects of N Deposition
• IndividualAltered morphologyChanges in reproduction, survivorship
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Effects of N Deposition
• IndividualAltered morphologyChanges in reproduction, survivorship
• PopulationIncreased long-term extinction riskChanges in short-term dynamics
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Effects of N Deposition• Individual
Altered morphologyChanges in reproduction, survivorship
• Population Increased long-term extinction riskChanges in short-term dynamics
• CommunityChanges in abundance and compositionAltered nutrient transfer and storage
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Effects of N Deposition on Carnivorous Plants
• Life History
• Effects on Individuals
• Effects on Populations
• Effects on Communities
• The Role of Ecologists
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Effects of N Deposition on Carnivorous Plants
• Life History
• Effects on Individuals
• Effects on Populations
• Effects on Communities
• The Role of Ecologists
![Page 9: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/9.jpg)
Carnivorous plants: well-known, but poorly studied
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Carnivory in plants
• Phylogenetically diverse
• Morphological, chemical adaptations for attracting, capturing, digesting arthropods
• Common in low N habitats
• Poor competitors for light, nutrients
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Family Sarraceniaceae
Genus Common Name
Number of Species
Distribution
Darlingtonia Cobra Lilly 1 Northwest USA
Heliamphora Sun Pitchers 5 North-central South America
Sarracenia Pitcher Plants 8 Eastern USA, Canada
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Genus Sarracenia
• 8 described species
• Center of diversity in southeastern US
• Many subvarieties
• Extensive hybridization
• Sarracenia purpurea (New Jersey- Canada)
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The Northern Pitcher Plant Sarracenia purpurea
• Perennial plant of low-N peatlands
• Lifespan 30-50 y
• Arthropod prey capture in water-filled pitchers
• Diverse inquiline community in pitchers
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The Inquilines
Wyeomyia smithii
Metriocnemus knabiHabrotrocha rosa
Blaesoxipha fletcheri
Sarraceniopus gibsoni
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Inquiline food web
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Phyllodia
• Flat leaves
• No prey capture
• High concentration of chlorophyll, stomates
• Photosynthetically more efficient than pitchers
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Flowering Stalks
• Single stalk per rosette
• Flowering after3 to 5 years
• Bumblebee, fly pollinated
• Short-distancedispersal of seeds
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Leaf Senescence
• End-of-season die off
• Production of new leaves in following spring
• Annual increase in rosette diameter
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Effects of N Deposition on Carnivorous Plants
• Life History
• Effects on Individuals
• Effects on Populations
• Effects on Communities
• The Role of Ecologists
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Nutrient Treatments
• Distilled H20
• Micronutrients• Low N (0.1 mg/L)• High N (1.0 mg/L)• Low P (0.025 mg/L)• High P (0.25 mg/L)
• N:P(1) Low N + Low P• N:P(2) Low N + High P• N:P(3) High N + Low P
Nutrient Source:
Micronutrients: Hoaglands
N: NH4Cl
P: NaH2PO4
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Anthropogenic N additions alter growth and morphology
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Anthropogenic N additions alter growth and morphology
Increasing N
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Effects of Anthropogenic N additions
• Increased production of phyllodiaPhenotypic shift from carnivory to
photosynthesis
• Increased probability of flowering
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Contrasting effects of anthropogenic N vs. N derived
from prey
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Food Addition Experiment
• Ecological “press” experiment
• Food supplemented with house flies• Treatments: 0, 2, 4 ,6, 8,10,12, 14 flies/week
• Plants harvested after one field season
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Food additions do not alter growth and morphology
Increasing prey
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N uptake increases with food level
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P uptake increases with food level
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N:P ratio decreases with increasing food supply
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Altered N:P ratios suggest P limitation under ambient conditions
P limitation (Koerselman & Meuleman 1996, Olde Venternik et al. in press)
Ambient
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Food additions do not alter growth and morphology
Increasing prey
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Anthropogenic N additions alter growth and morphology
Increasing N
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Contrasting effects of anthropogenic and natural sources of N
Anthropogenic NAltered N:P ratiosMorphological shiftReduction in prey uptake
Prey NUptake, storage of N & PNo morphological shiftsContinued prey uptake
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Effects of N Deposition on Carnivorous Plants
• Life History
• Effects on Individuals
• Effects on Populations
• Effects on Communities
• The Role of Ecologists
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Study Sites
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Demography survey
• 100 adult, juvenile plants tagged at each site
• Plants censused and measured each year
• Seed plantings to estimate recruitment functions
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Recruits Juveniles Adults Flowering Adults
Sarracenia matrix model
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Recruits Juveniles Adults Flowering Adults
Hawley Bog Transitions
0.10
4.00
0.04
0.09
0.18
0.83
0.95 0.70 0.17
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Recruits Juveniles Adults Flowering Adults
Molly Bog Transitions
0.10
4.00
0.13
0.17
0.10
0.66
0.85 0.71 0.31
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Matrix Transition Model(stationary)
nt+1 = Ant
Population vector at time (t + 1)
Transition matrixPopulation vector at time (t)
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Population Projections
Site r individuals/individual•year Doubling Time
Hawley Bog 0.00456 152 y
Molly Bog 0.00554 125 y
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Deterministic Model: Results
• Growth, survivorship, and reproduction are closely balanced in both sites
• Doubling times > 100 y
• Juvenile, adult persistence contribute most to population growth rate
• Sexual reproduction, recruitment relatively unimportant
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Matrix Transition Model(stochastic)
nt+1 = Atnt
Population vector at time (t + 1)
Random transition matrix at time (t)
Population vector at time (t)
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Hawley Bog
-0.30-0.20-0.100.000.100.200.30
Deter
min
istic
ME (m
ild)
ME (s
ever
e)
ES (mild
)
ES (sev
ere)
r
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Hawley Bog
200
400
600
800
1000
0 5 10 15 20 25Time Step
n
DeterministicES (severe)ES (mild)
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Stochastic Model: Results
• Cannot reject H0 (r = 0.0)
• Environmental variation can lead to a substantial risk of long-term extinction (0.3 < p(ext) < 0.6)
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How do N and P concentrations affect population growth of
Sarracenia?
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Nutrient Addition Experiment
• 10 juveniles, 10 adults/treatment
• Nutrients added to leaves twice/month
• Nutrient concentrations bracket observed field values
• Nutrient treatments maintained 1998, 1999
• “Press” experiment
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Nutrient Treatments
• Distilled H20
• Micronutrients• Low N (0.1 mg/L)• High N (1.0 mg/L)• Low P (0.025 mg/L)• High P (0.25 mg/L)
• N:P(1) Low N + Low P• N:P(2) Low N + High P• N:P(3) High N + Low P
Nutrient Source:
Micronutrients: Hoaglands
N: NH4Cl
P: NaH2PO4
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Effects of N additions
• Increased production of phyllodia
• Increased probability of flowering
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Effects of N additions
• Increased production of phyllodia
• Increased probability of flowering
• Decreased juvenile survivorship
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Population Growth Rate(Deterministic)
-0.15
-0.10
-0.05
0.00
0.05
r
Distilled Micros Low N
High N Low P High P
NP (2) NP (1) NP (3)
LLM
H
H
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Population Growth Rate(Measurement Error)
-0.40
-0.35
-0.30
-0.25
-0.20
-0.15
-0.10
-0.05
0.00
0.05
0.10
Distil
led
Mic
ros
Low N
High
N
Low P
High
P
NP (2)
NP (1)
NP (3)
r
H
ML
H
L
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Effects of Nitrogen on Demography: Results
• Population growth rates respond to different N and P regimes
• Population growth rate decreases in response to increasing N
• Population growth rate decreases in responses to increasing N:P
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Modeling Long-term Environmental Change
Observed N Deposition
Long-termForecast
N(t)
Transition Matrix (t)
PopulationStructure (t)
Time Series Modeling
Transition Function
Population Time SeriesExtinction RiskTime to Extinction
MatrixMultiplication
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Modeling Long-term Environmental Change
Observed N Deposition
Long-termForecast
N(t)
Transition Matrix (t)
PopulationStructure (t)
Time Series Modeling
Transition Function
Population Time SeriesExtinction RiskTime to Extinction
MatrixMultiplication
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N monitoring
• National Atmospheric Deposition Program
• NH4, NO3 measured as mg/l/yr
• Annual data 1984-1998
• Monitoring sitesShelburne, VTQuabbin, MA
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0.00
0.05
0.10
0.15
0.20
0.25
0.30
0.35
1984
1986
1988
1990
1992
1994
1996
1998
0.00
0.05
0.10
0.15
0.20
0.25
0.30
0.35
1984
1986
1988
1990
1992
1994
1996
1998
0.0
0.5
1.0
1.5
2.0
2.5
1984
1986
1988
1990
1992
1994
1996
1998
0.0
0.5
1.0
1.5
2.0
2.5
1984
1986
1988
1990
1992
1994
1996
1998
Quabbin, MA Shelburne, VT
NH4
N03
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0.00
0.05
0.10
0.15
0.20
0.25
0.30
0.35
1984
1986
1988
1990
1992
1994
1996
1998
0.00
0.05
0.10
0.15
0.20
0.25
0.30
0.35
1984
1986
1988
1990
1992
1994
1996
1998
0.0
0.5
1.0
1.5
2.0
2.5
1984
1986
1988
1990
1992
1994
1996
1998
0.0
0.5
1.0
1.5
2.0
2.5
1984
1986
1988
1990
1992
1994
1996
1998
Quabbin, MA Shelburne, VT
NH4
N03
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Regression Models
Ordinary Least Squares (OLS)
Nt = a + bt + e
First-order auto-regressive (AR-1)
Nt = a +bNt-1 + e
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Quabbin (AR-1)
0.01
0.1
1
10
1 10 19 28 37 46
Year
N (m
g/l/y
r) b = 0.947
b = 1.000
b = 1.053
Quabbin (OLS)
00.10.20.30.40.50.60.7
1 10 19 28 37 46
Year
N (m
g/l/y
) b = -0.004
b = 0.000
b = 0.004
Shelburne (AR-1)
0.01
0.1
1
10
1 10 19 28 37 46
Year
N (m
g/l/y
)
b = 0.978
b = 1.000
b = 1.022
Shelburne (OLS)
0.2
0.3
0.4
0.5
0.6
0.7
1 10 19 28 37 46
Year
N (m
g/l/y
) b = -0.001
b = 0.000
b = 0.001
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Modeling Long-term Environmental Change
Observed N Deposition
Long-termForecast
N(t)
Transition Matrix (t)
PopulationStructure (t)
Time Series Modeling
Transition Function
Population Time SeriesExtinction RiskTime to Extinction
MatrixMultiplication
![Page 75: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/75.jpg)
Establishing Transition Functions
• Linear interpolation between observed data points (n = 3)
• Asymptotic transitions at extreme levels of nitrogen:pij = observed pij if [N] < 0.01 mg/l/yr
pij = 0.0 if [N] > 10.0 mg/l/yr
• Logarithmic response curve
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0
0.2
0.4
0.6
0.8
1
-3 -2 -1 0 1 2
Log10 [N]
Ad
ult
s →
Ad
ult
s
ExtrapolatedObserved
Modeling Demographic Transitions as a Function of Nitrogen
![Page 77: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/77.jpg)
Modeling Long-term Environmental Change
Observed N Deposition
Long-termForecast
N(t)
Transition Matrix (t)
PopulationStructure (t)
Time Series Modeling
Transition Function
Population Time SeriesExtinction RiskTime to Extinction
MatrixMultiplication
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Matrix Transition Model(changing environment)
nt+1 = Atnt
Population vector at time (t + 1)
Sequentially changing transition matrix at time (t)
Population vector at time (t)
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Estimated population size at Hawley bog
Stage Number of individuals
Recruits 1500
Juveniles 23,500
Non-flowering Adults 1400
Flowering Adults 500
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Quabbin Exponential Forecast Models (AR-1)
Scenario Annual % Change
P (ext) at 100 y
Time to ext (p = 0.95)
Best case -4.7% 0.00 > 10,000 y
No change 0.0% 0.038 650 y
Small increase
1% 0.378 290 y
Worst case 4.7% 0.996 70 y
![Page 81: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/81.jpg)
Shelburne Exponential Forecast Models (AR-1)
Scenario Annual % Change
P (ext) at 100 y
Time to ext (p = 0.95)
Best case -2.2% 0.158 > 10,000 y
No change 0.0% 0.510 230 y
Small increase
1.0% 0.694 200 y
Worst case 2.2% 0.838 140 y
![Page 82: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/82.jpg)
Shelburne Nitrogen Forecast Model
0
5000
10000
15000
20000
25000
30000
1 10 19 28 37 46
Year
Po
pu
lati
on
Siz
e
AR
OLS
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Forecasting Models for Nitrogen Deposition: Results
• Increasing or stationary models of Nitrogen deposition drive Sarracenia populations to extinction
• Extinction risk declines with reduced nitrogen
• Correlated nitrogen series can induce cycles and complex population dynamics
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Effects of N Deposition on Carnivorous Plants
• Life History
• Effects on Individuals
• Effects on Populations
• Effects on Communities
• The Role of Ecologists
![Page 85: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/85.jpg)
Sarracenia Nutrient Feedback Loop
PitcherNutrient
Pool [N,P]
InquilineCommunity
ArthropodPrey
PlantGrowth
AtmosphericDeposition
![Page 86: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/86.jpg)
Sarracenia Nutrient Feedback Loop
PitcherNutrient
Pool [N,P]
InquilineCommunity
ArthropodPrey
PlantGrowth
AtmosphericDeposition
![Page 87: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/87.jpg)
Sarracenia Nutrient Feedback Loop
PitcherNutrient
Pool [N,P]
InquilineCommunity
ArthropodPrey
PlantGrowth
AtmosphericDeposition
![Page 88: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/88.jpg)
Sarracenia Nutrient Feedback Loop
PitcherNutrient
Pool [N,P]
InquilineCommunity
ArthropodPrey
PlantGrowth
AtmosphericDeposition
![Page 89: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/89.jpg)
Nutrients → Inquilines
• Manipulate [N], [P] in leaves
• Orthogonal “regression” design
• Maintain [] in a “press” experiment
![Page 90: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/90.jpg)
Response Surface Experimenal Design
[N]0 1 2 3 4 5 6 7
[P]
0
1
2
3
4
5
6
7
![Page 91: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/91.jpg)
Effects of [N,P] on Inquiline Abundance
0
10
20
30
40
01
23
45
6
0123456
Rot
ifer
Abu
ndan
ce
[P]
[N]
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Sarracenia Nutrient Feedback Loop
PitcherNutrient
Pool [N,P]
InquilineCommunity
ArthropodPrey
PlantGrowth
AtmosphericDeposition
![Page 93: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/93.jpg)
Inquilines → Nutrients
• Manipulate [N], [P] in leaves
• Orthogonal “regression” design
• Establish initial [] in a “pulse” experiment
![Page 94: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/94.jpg)
Response Surface Experimenal Design
[N]0 1 2 3 4 5 6 7
[P]
0
1
2
3
4
5
6
7
![Page 95: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/95.jpg)
Null Hypothesis
[N]0 1 2 3 4 5 6 7
[P]
0
1
2
3
4
5
6
7
![Page 96: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/96.jpg)
Community Regulation of Nutrients
[N]0 1 2 3 4 5 6 7
[P]
0
1
2
3
4
5
6
7
![Page 97: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/97.jpg)
![Page 98: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/98.jpg)
Sarracenia Nutrient Feedback Loop
PitcherNutrient
Pool [N,P]
InquilineCommunity
ArthropodPrey
PlantGrowth
AtmosphericDeposition
![Page 99: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/99.jpg)
Nutrients ↔ Inquilines
),,(
),,(
tNIgdt
dI
tINfdt
dN
![Page 100: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/100.jpg)
![Page 101: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/101.jpg)
![Page 102: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/102.jpg)
Effects of N Deposition on Carnivorous Plants
• Life History
• Effects on Individuals
• Effects on Populations
• Effects on Communities
• The Role of Ecologists
![Page 103: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/103.jpg)
0
50
100
150
200
1900 1920 1940 1960 1980 2000
Year
Ter
agra
ms
of
Nit
rog
en
Fertilizer NOx
Total anthropogenic N fixed
Natural range
![Page 104: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/104.jpg)
Ecology ≠
Environmental Science
![Page 105: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/105.jpg)
Reasons for Studying Ecology
![Page 106: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/106.jpg)
Reasons for Studying Ecology
• Natural History
![Page 107: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/107.jpg)
![Page 108: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/108.jpg)
Reasons for Studying Ecology
• Natural History
• Field Studies & Experiments
![Page 109: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/109.jpg)
![Page 110: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/110.jpg)
Reasons for Studying Ecology
• Natural History
• Field Studies & Experiments
• Statistics & Data Analysis
![Page 111: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/111.jpg)
Population Growth Rate(Deterministic)
-0.15
-0.10
-0.05
0.00
0.05
r
Distilled Micros Low N
High N Low P High P
NP (2) NP (1) NP (3)
0.00
0.05
0.10
0.15
0.20
0.25
0.30
0.35
1984
1986
1988
1990
1992
1994
1996
1998
![Page 112: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/112.jpg)
Reasons for Studying Ecology
• Natural History
• Field Studies & Experiments
• Statistics & Data Analysis
• Modeling
![Page 113: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/113.jpg)
),,(
),,(
tNIgdt
dI
tINfdt
dN
PitcherNutrient
Pool [N,P]
InquilineCommunity
ArthropodPrey
PlantGrowth
AtmosphericDeposition
0
5000
10000
15000
20000
25000
30000
1 10 19 28 37 46
Year
Po
pu
lati
on
Siz
e
AR
OLS
![Page 114: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/114.jpg)
Reasons for Studying Ecology
• Natural History
• Field Studies & Experiments
• Statistics & Data Analysis
• Modeling
• Collaboration
![Page 115: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/115.jpg)
Aaron M. Ellison
Harvard Forest
![Page 116: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/116.jpg)
Conclusions• Anthropogenic deposition of N is a major ecological
challenge
![Page 117: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/117.jpg)
Conclusions• Anthropogenic deposition of N is a major ecological
challenge• Carnivorous plants in ombrotrophic bogs are a model
system
![Page 118: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/118.jpg)
Conclusions• Anthropogenic deposition of N is a major ecological
challenge• Carnivorous plants in ombrotrophic bogs are a model
system• Individual response
plants alter morphology and growth in response to N:P ratios
![Page 119: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/119.jpg)
Conclusions• Anthropogenic deposition of N is a major ecological
challenge• Carnivorous plants in ombrotrophic bogs are a model
system• Individual response
plants alter morphology and growth in response to N:P ratios
• Population response N and P environments affect population growth rate
![Page 120: Nitrogen deposition and extinction risk in carnivorous plants: ecological challenges for the next century Nicholas J. Gotelli Department of Biology University.](https://reader038.fdocuments.in/reader038/viewer/2022110320/56649cad5503460f9496f0a8/html5/thumbnails/120.jpg)
Conclusions• Anthropogenic deposition of N is a major ecological
challenge• Carnivorous plants in ombrotrophic bogs are a model
system• Individual response
plants alter morphology and growth in response to N:P ratios
• Population response N and P environments affect population growth rate
• Community response Further study of nutrient ↔ inquiline feedback loop
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