nicholas j. gotelli department of biology university of vermont burlington, vt 05405 u.s.a
DESCRIPTION
The Ecological Impacts Of Nitrogen Deposition: Insights From The Carnivorous Pitcher Plant Sarracenia purpurea. Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A. Effects of N Deposition. Individual Altered morphology - PowerPoint PPT PresentationTRANSCRIPT
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The Ecological Impacts Of Nitrogen Deposition: Insights From The Carnivorous
Pitcher Plant Sarracenia purpurea
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
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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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Wakefield, A. E., N. J. Gotelli, S. E. Wittman, and A. M. Ellison. 2005. Prey addition alters nutrient stoichiometry of the carnivorous plant Sarracenia purpurea. Ecology 86: 1737-1743.
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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 added food
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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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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
Although Sarracenia has evolved adaptations for low N environments, chronic N deposition may have caused populations to be currently limited by P, not N.
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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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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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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
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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
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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 74: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/74.jpg)
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)
![Page 75: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/75.jpg)
Estimated population size at Hawley bog
Stage Number of individuals
Recruits 1500
Juveniles 23,500
Non-flowering Adults 1400
Flowering Adults 500
![Page 76: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/76.jpg)
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 77: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/77.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
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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
![Page 80: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/80.jpg)
Effects of N Deposition on Carnivorous Plants
• Life History
• Effects on Individuals
• Effects on Populations
• Effects on Communities
• The Role of Ecologists
![Page 81: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/81.jpg)
Sarracenia Nutrient Feedback Loop
PitcherNutrient
Pool [N,P]
InquilineCommunity
ArthropodPrey
PlantGrowth
AtmosphericDeposition
![Page 82: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/82.jpg)
Sarracenia Nutrient Feedback Loop
PitcherNutrient
Pool [N,P]
InquilineCommunity
ArthropodPrey
PlantGrowth
AtmosphericDeposition
![Page 83: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/83.jpg)
Sarracenia Nutrient Feedback Loop
PitcherNutrient
Pool [N,P]
InquilineCommunity
ArthropodPrey
PlantGrowth
AtmosphericDeposition
![Page 84: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/84.jpg)
Sarracenia Nutrient Feedback Loop
PitcherNutrient
Pool [N,P]
InquilineCommunity
ArthropodPrey
PlantGrowth
AtmosphericDeposition
![Page 85: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/85.jpg)
Four-level Multi-Factorial Experiment
• Atmospheric N (8 levels)
• Prey supplement (yes,no)
• Top predator removal (yes,no)
![Page 86: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/86.jpg)
Four-level Multi-Factorial Experiment
• Atmospheric N (8 levels)
• Prey supplement (yes,no)
• Top predator removal (yes,no)
• Nutrient exchange with plant(unmanipulated, isolated, control)
![Page 87: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/87.jpg)
Quantifying Trophic Structure• Food web saturation is our response variable. Each taxon in the food web is • given a binary value representing its presence (1xxxx) or absence (0xxxx):
• Taxon Binary value Decimal value• Metriocnemus 1 1• Habrotrocha 10 2• Sarraceniopus 100 4• Wyeomyia 1000 8• Fletcherimyia 10000 16
• The saturation of the food web in a given pitcher is the sum of the equivalent• decimal values of each taxon present. Food webs with higher saturation values have • both more trophic levels present and more trophic links present. There are 32 possible • food webs that can be assembled with these 5 taxa; the decimal value for each food • web ranges from 0 – 31, with increasing numbers indicating more saturated food webs.
![Page 88: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/88.jpg)
Nutrient exchange with the plant and top predators affect food web structure
![Page 89: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/89.jpg)
Sarracenia Nutrient Feedback Loop
PitcherNutrient
Pool [N,P]
InquilineCommunity
ArthropodPrey
PlantGrowth
AtmosphericDeposition
![Page 90: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/90.jpg)
Inquilines → Nutrients
• Manipulate [N], [P] in leaves
• Orthogonal “regression” design
• Establish initial [] in a “pulse” experiment
![Page 91: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/91.jpg)
Response Surface Experimenal Design
[N]0 1 2 3 4 5 6 7
[P]
0
1
2
3
4
5
6
7
![Page 92: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/92.jpg)
Null Hypothesis
[N]0 1 2 3 4 5 6 7
[P]
0
1
2
3
4
5
6
7
![Page 93: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/93.jpg)
Community Regulation of Nutrients
[N]0 1 2 3 4 5 6 7
[P]
0
1
2
3
4
5
6
7
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Sarracenia Nutrient Feedback Loop
PitcherNutrient
Pool [N,P]
InquilineCommunity
ArthropodPrey
PlantGrowth
AtmosphericDeposition
![Page 96: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/96.jpg)
Nutrients ↔ Inquilines
),,(
),,(
tNIgdt
dI
tINfdt
dN
![Page 97: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/97.jpg)
![Page 98: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/98.jpg)
![Page 99: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/99.jpg)
Effects of N Deposition on Carnivorous Plants
• Life History
• Effects on Individuals
• Effects on Populations
• Effects on Communities
• The Role of Ecologists
![Page 100: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/100.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 101: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/101.jpg)
Ecology ≠
Environmental Science
![Page 102: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/102.jpg)
Reasons for Studying Ecology
![Page 103: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/103.jpg)
Reasons for Studying Ecology
• Natural History
![Page 104: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/104.jpg)
![Page 105: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/105.jpg)
Reasons for Studying Ecology
• Natural History
• Field Studies & Experiments
![Page 106: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/106.jpg)
![Page 107: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/107.jpg)
Reasons for Studying Ecology
• Natural History
• Field Studies & Experiments
• Statistics & Data Analysis
![Page 108: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/108.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 109: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/109.jpg)
Reasons for Studying Ecology
• Natural History
• Field Studies & Experiments
• Statistics & Data Analysis
• Modeling
![Page 110: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/110.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 111: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/111.jpg)
Reasons for Studying Ecology
• Natural History
• Field Studies & Experiments
• Statistics & Data Analysis
• Modeling
• Collaboration
![Page 112: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/112.jpg)
Aaron M. Ellison
Harvard Forest
![Page 113: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/113.jpg)
Conclusions• Anthropogenic deposition of N is a major ecological
challenge
![Page 114: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/114.jpg)
Conclusions• Anthropogenic deposition of N is a major ecological
challenge• Carnivorous plants in ombrotrophic bogs are a model
system
![Page 115: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/115.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 116: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/116.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 117: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/117.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
![Page 118: Nicholas J. Gotelli Department of Biology University of Vermont Burlington, VT 05405 U.S.A](https://reader035.vdocuments.us/reader035/viewer/2022062801/56814320550346895daf875f/html5/thumbnails/118.jpg)