what is an individual counting...

17
2/24/2014 1 Act II: Single Species Dynamics Scene I: Life, Death, and Life Histories (the chapter from hell) Population β€’ Number of individuals in an area β€’ Density β€’ β„Ž = ++++ What is an individual Unitary Modular Final size Determinate Indeterminate Predictable shape Predictable Less predictable Form of growth Unit Modular Examples Human Tree, corals, hydroids Senescence Predictable Less predictable Ecological individual Genet Ramet or Genet Counting Individuals β€’ Modular organisms – Ramets, genets, biomass β€’ Unitary organisms – Census – Index – Population estimate Births and deaths β€’ Damned difficult β€’ Births – Hard to quantify – 50% of zygotes in rabbits abort – 30% of human zygotes make it full term β€’ Deaths – Rarely find dead individuals – Small plants and animals may get consumed completely Iteroparity and semelparity

Upload: others

Post on 24-Mar-2020

4 views

Category:

Documents


0 download

TRANSCRIPT

Page 1: What is an individual Counting Individualsweb.wilkes.edu/jeffrey.stratford/files/ecology/act2.pdfPopulation β€’ Number of individuals in an area β€’ Density β€’ π‘π‘‘β„Ž =𝑁 +𝑏+

2/24/2014

1

Act II: Single Species Dynamics

Scene I: Life, Death, and Life Histories

(the chapter from hell)

Population

β€’ Number of individuals in an area

β€’ Density

β€’ π‘π‘‘β„Žπ‘’π‘› = π‘π‘›π‘œπ‘€ + 𝑏 + 𝑑 + 𝑖 + 𝑒

What is an individual

Unitary Modular

Final size Determinate Indeterminate

Predictable shape Predictable Less predictable

Form of growth Unit Modular

Examples Human Tree, corals, hydroids

Senescence Predictable Less predictable

Ecological individual Genet Ramet or Genet

Counting Individuals

β€’ Modular organisms

– Ramets, genets, biomass

β€’ Unitary organisms

– Census

– Index

– Population estimate

Births and deaths

β€’ Damned difficult

β€’ Births – Hard to quantify

– 50% of zygotes in rabbits abort

– 30% of human zygotes make it full term

β€’ Deaths – Rarely find dead individuals

– Small plants and animals may get consumed completely

Iteroparity and semelparity

Page 2: What is an individual Counting Individualsweb.wilkes.edu/jeffrey.stratford/files/ecology/act2.pdfPopulation β€’ Number of individuals in an area β€’ Density β€’ π‘π‘‘β„Ž =𝑁 +𝑏+

2/24/2014

2

Climate and reproductive strategies

β€’ Seasonal climates have a greater proportion of semelparous or seasonal iteroparous species

β€’ Seasonality may be induced by precipitation

Examples of diversity in life cycles

Semelparous/Overlapping generations Long-lived

semelparous species

Long-lived (we hope) iteroparous species

Annuals

lx = ax/a0

dx = lx – lx+1

qx = dx/lx*1/days

β€’ dx can be summed

β€’ qx gives intensity in a particular period

Page 3: What is an individual Counting Individualsweb.wilkes.edu/jeffrey.stratford/files/ecology/act2.pdfPopulation β€’ Number of individuals in an area β€’ Density β€’ π‘π‘‘β„Ž =𝑁 +𝑏+

2/24/2014

3

Seed Banks

Static vs. Cohort Life Tables

β€’ Cohort

– Strengths

– Disadvantages

β€’ Static

– Strengths

– Disadvantages

Page 4: What is an individual Counting Individualsweb.wilkes.edu/jeffrey.stratford/files/ecology/act2.pdfPopulation β€’ Number of individuals in an area β€’ Density β€’ π‘π‘‘β„Ž =𝑁 +𝑏+

2/24/2014

4

Basic reproductive rate

β€’ 𝑅0 = 𝑙π‘₯π‘šπ‘₯

β€’ When non-overlapping generations R = R0

β€’ 𝑁1 = 𝑁0𝑅

β€’ 𝑁2 = 𝑁1𝑅 = 𝑁0𝑅𝑅 = 𝑁0𝑅2

β€’ 𝑁3 = 𝑁2𝑅𝑅 = 𝑁0𝑅𝑅𝑅 = 𝑁0𝑅3

β€’ So, generally, 𝑁𝑑 = ?

β€’ And 𝑅0 = 𝑅𝑑 -> take log of both sides gives:

Intrinsic rate of increase

β€’ If Ln R = r

β€’ Then π‘Ÿ =ln 𝑅0

𝑇

β€’ When population are overlapping then use

β€’ π‘Ÿ =ln 𝑅0

𝑇𝑐 where Tc is the cohort generation

time

β€’ 𝑇𝑐 = π‘₯𝑙π‘₯π‘šπ‘₯

𝑅0

Stable Age Distribution

Life history

β€’ Lifetime pattern of growth, differentiation, storage, and reproduction

p.s. don’t Google β€œorchid seed” = w.t.h.!

Two majors elements

β€’ Optimization

β€’ Bet-hedging

β€’ Trade-offs

Page 5: What is an individual Counting Individualsweb.wilkes.edu/jeffrey.stratford/files/ecology/act2.pdfPopulation β€’ Number of individuals in an area β€’ Density β€’ π‘π‘‘β„Ž =𝑁 +𝑏+

2/24/2014

5

Allocation of limited resources

Page 6: What is an individual Counting Individualsweb.wilkes.edu/jeffrey.stratford/files/ecology/act2.pdfPopulation β€’ Number of individuals in an area β€’ Density β€’ π‘π‘‘β„Ž =𝑁 +𝑏+

2/24/2014

6

Clutch size vs. litter size

β€’ In general, if more produced then smaller produced

Survivorship and reproduction

β€’ When survivorship is low, reproduction is high

β€’ Age of maturity

β€’ Semelparity

r-K spectrum

β€’ Developed by MacArthur and Wilson 1967 in the Theory of Island Biogeography

β€’ r species: maximize reproduction

β€’ K species: maximize competitive edge

β€’ Make a table that includes habitat, propagule size, time to maturity, clutch size, parental care, semelparity

β€’ Support

Phenotypic Plasticity

β€’ P = G

β€’ P = G + E

β€’ P = G + E + G*E

β€’ Plasticity is the aspect of the phenotype of the phenotype that is determined by the environment

β€’ Are phenotypic responses optimal?

Page 7: What is an individual Counting Individualsweb.wilkes.edu/jeffrey.stratford/files/ecology/act2.pdfPopulation β€’ Number of individuals in an area β€’ Density β€’ π‘π‘‘β„Ž =𝑁 +𝑏+

2/24/2014

7

Constraints

β€’ Phylogenetic

β€’ Physics

β€’ Example: Allometric relationships

– At the cell

– At the organismal

Phylogenetic effects

β€’ The comparative method – Harvey and Pagel 1991

More on single species dynamics

Leslie Matrix

L =

F0 F1 F2 F3

S0 0 0 0

0 S1 0 0

0 0 S2 0

Γ©

Γ«

ΓͺΓͺΓͺΓͺΓͺ

ΓΉ

Γ»

ΓΊΓΊΓΊΓΊΓΊ

Page 8: What is an individual Counting Individualsweb.wilkes.edu/jeffrey.stratford/files/ecology/act2.pdfPopulation β€’ Number of individuals in an area β€’ Density β€’ π‘π‘‘β„Ž =𝑁 +𝑏+

2/24/2014

8

Elements of Leslie Matrix (L)

Fx – Age-specific Fecundity Γ— age-specific survival

Sx –Age-specific Survival

Fx = Sx mx+1

How does the Leslie Matrix estimate Population Growth?

Nt+1 = LΒ΄Nt

Population Projection

Nt+1 =

F0 F1 F2 F3

S0 0 0 0

0 S1 0 0

0 0 S2 0

Γ©

Γ«

ΓͺΓͺΓͺΓͺΓͺ

ΓΉ

Γ»

ΓΊΓΊΓΊΓΊΓΊ

Β΄Nt

Population Projection

N0,t+1

N1.t+1

N2,t+1

N3,t+1

Γ©

Γ«

ΓͺΓͺΓͺΓͺΓͺ

ΓΉ

Γ»

ΓΊΓΊΓΊΓΊΓΊ

=

F0 F1 F2 F3

S0 0 0 0

0 S1 0 0

0 0 S2 0

Γ©

Γ«

ΓͺΓͺΓͺΓͺΓͺ

ΓΉ

Γ»

ΓΊΓΊΓΊΓΊΓΊ

Β΄

N0,t

N1,t

N2,t

N3,t

Γ©

Γ«

ΓͺΓͺΓͺΓͺΓͺ

ΓΉ

Γ»

ΓΊΓΊΓΊΓΊΓΊ

Assumptions

β€’ Individuals can be aged reliably

β€’ No age-effects in vital rates

β€’ Vital rates are constant

– Constant environment

– No density dependence

– stochastic Leslie Matrices possible

β€’ Sex ratio at birth is 1:1

– i.e., male and female vital rates are congruent

Advantages of Leslie Matrix

β€’ Stable-age distribution not assumed

β€’ Sensitivity analyses –

– can identify main age-specific vital rates that affect abundance and age structure

β€’ Modify the analyses to include density-dependence

β€’ Derive finite rate of population change (Ξ») and SAD

Page 9: What is an individual Counting Individualsweb.wilkes.edu/jeffrey.stratford/files/ecology/act2.pdfPopulation β€’ Number of individuals in an area β€’ Density β€’ π‘π‘‘β„Ž =𝑁 +𝑏+

2/24/2014

9

Disadvantage of Leslie Matrix

β€’ See assumptions

β€’ Age data may not be available

– can use stage-based Lefkovitch Matrix

β€’ Fecundity data may not be available for all ages

EigenAnalysis of L

β€’ Eigenvalues – – dominant = population growth rate

β€’ asymptotic growth rate at Stable Age Distribution

β€’ Stable Age Structure – right eigenvector

β€’ Reproductive Value – left eigenvector

Other Statistics β€’ Sensitivities

– how Ξ» varies with a change in matrix elements β€’ absolute changes in matrix elements

β€’ Elasticities – how Ξ» varies with a change in a vital rate holding

other rates constant

–

β€’ Damping ratio – rate population approaches equilibrium - SAD

r =

l1

l2

Act II Scene II

β€’ Darwin

β€’ Competition

February 22, 1809

Happy Darwin Day

Darwin’s Childhood

Page 10: What is an individual Counting Individualsweb.wilkes.edu/jeffrey.stratford/files/ecology/act2.pdfPopulation β€’ Number of individuals in an area β€’ Density β€’ π‘π‘‘β„Ž =𝑁 +𝑏+

2/24/2014

10

Darwin’s Education Darwin’s Education

Voyage of the Beagle Voyage of the Beagle

Voyage of the Beagle Voyage of the Beagle

β€’ Immensity of Time

– Charles Lyell

Page 11: What is an individual Counting Individualsweb.wilkes.edu/jeffrey.stratford/files/ecology/act2.pdfPopulation β€’ Number of individuals in an area β€’ Density β€’ π‘π‘‘β„Ž =𝑁 +𝑏+

2/24/2014

11

Voyage of the Beagle

β€’ Thomas Malthus

– Struggle for existence

Voyage of the Beagle

Back home Back home

Back home Development of natural selection

β€’ Deep time

β€’ Struggle for existence

β€’ Genealogical connection with past creatures

β€’ Isolation

β€’ Variation is important

Page 12: What is an individual Counting Individualsweb.wilkes.edu/jeffrey.stratford/files/ecology/act2.pdfPopulation β€’ Number of individuals in an area β€’ Density β€’ π‘π‘‘β„Ž =𝑁 +𝑏+

2/24/2014

12

Alfred Russell Wallace Origin of Species

Other stuff Darwin did

β€’ Expression of emotion

β€’ Human history

β€’ Action of earthworms

β€’ Barnacles

β€’ Pollination

Darwin’s children

Up to now

β€’ How organisms interact with the environment

β€’ What has shaped the organisms (natural selection + constraints)

β€’ Simple populations

Intraspecific Competition

β€’ Exploitation

β€’ Interference

– Includes mates

Page 13: What is an individual Counting Individualsweb.wilkes.edu/jeffrey.stratford/files/ecology/act2.pdfPopulation β€’ Number of individuals in an area β€’ Density β€’ π‘π‘‘β„Ž =𝑁 +𝑏+

2/24/2014

13

So, increased density =

β€’ Increased mortality

β€’ Decreased fertility

β€’ Caveats

– Group selection

Density

β€’ Individuals per unit area

β€’ Individuals per unit resource

β€’ Caveats

– Group size

Page 14: What is an individual Counting Individualsweb.wilkes.edu/jeffrey.stratford/files/ecology/act2.pdfPopulation β€’ Number of individuals in an area β€’ Density β€’ π‘π‘‘β„Ž =𝑁 +𝑏+

2/24/2014

14

Carrying Capacity

β€’ b = d

What does this mean for K?

Sigmoidal Growth Curve Sigmoidal Growth Curve

β€’ Growth rates along the curve

β€’ Caveat

– Beware the asymptote!

– Competition requires

– What else happens to populations

Page 15: What is an individual Counting Individualsweb.wilkes.edu/jeffrey.stratford/files/ecology/act2.pdfPopulation β€’ Number of individuals in an area β€’ Density β€’ π‘π‘‘β„Ž =𝑁 +𝑏+

2/24/2014

15

Models

β€’ What is a model?

β€’ All models are wrong

β€’ But some are useful

Discrete breeding seasons

β€’ R is the fundamental net reproductive rate

β€’ 𝑁𝑑+1 = 𝑁𝑑𝑅

β€’ 𝑁𝑑 = 𝑁0𝑅𝑑

β€’ Assumes R is constant

Discrete with competition

β€’ At a small population size the effect of competition is small so

‒𝑁𝑑

𝑁𝑑+1=

1

𝑅

β€’ To add competition

β€’ 𝑁𝑑+1 =𝑁𝑑𝑅

1+(π‘…βˆ’1)𝑁𝑑

𝐾

but, if (R-1)/K=a then

β€’ 𝑁𝑑+1 =𝑁𝑑𝑅

1+π‘Žπ‘π‘‘ what is the effect of a on Nt+1 as

a increases

Time lags

β€’ Models presented so far represent populations that respond instantaneously to competition

β€’ A time lag implies that the population responds to previous competition – Time lags can be -1 generations or more – Time lags induce oscillations – Dampening is the loss of oscillations

β€’ These models can be called time-dependent or autoregressive models

β€’ The effect of neighbors on a data point is called autocorrelation (spatial and temporal.. Or both)

Time lag

β€’ 𝑁𝑑+1 =𝑁𝑑𝑅

1+π‘Žπ‘π‘‘βˆ’1

β€’ 𝑁𝑑+1 =𝑁𝑑𝑅

1+ π‘Žπ‘π‘‘π‘

Page 16: What is an individual Counting Individualsweb.wilkes.edu/jeffrey.stratford/files/ecology/act2.pdfPopulation β€’ Number of individuals in an area β€’ Density β€’ π‘π‘‘β„Ž =𝑁 +𝑏+

2/24/2014

16

Implications

β€’ When R is large populations are likely to oscillate without extrinsic forces

Continuous breeding

‒𝑑𝑁

𝑑𝑑= π‘Ÿπ‘

β€’ r = ln(R)

β€’ Note that this is not the population size but what?

β€’ With intraspecific competition

‒𝑑𝑁

𝑑𝑑= π‘Ÿπ‘

πΎβˆ’π‘

𝐾

β€’ Logistic equation (Verhulst 1838)

β€’ Why logistic?

Competition: Effects on individuals

β€’ Body size: body size decreases but skewness increases with increasing density

Asymmetry in competition Asymmetry in intraspecific

competition

β€’IS NATURAL SELECTION

Page 17: What is an individual Counting Individualsweb.wilkes.edu/jeffrey.stratford/files/ecology/act2.pdfPopulation β€’ Number of individuals in an area β€’ Density β€’ π‘π‘‘β„Ž =𝑁 +𝑏+

2/24/2014

17

Territoriality

β€’ Resources (usually space) is defended β€’ Costs

– Defense, moving

β€’ Benefits – Mates – Predictable resources

β€’ When does territoriality break down? – Unpredictable resources – Superabundant resources – Simply put c > b

Self-thinning