Ground Rules, exams, etc. (no

1
Physiological EcologyHomeostasis maintenance of
a relatively stable internal state under a much
wider range of external environmental
conditions.Temperature regulation
(thermoregulation)Osmoregulation (water, salt
balance)Physiological Optima and Evolution of
Tolerance CurvesTolerance limits, optimaNet
benefitAcclimation Principle of Allocation Time,
Matter, and Energy Budgets Costs and Profits of
Leaf Size, Shape, and Placement Light, Water
availability, Prevailing winds,
Herbivores Multivariate statistics correlated
data Change co-ordinate systems reduce
dimensionality
Lecture 11 27 February 2020

2
In lowland wet tropical rainforest,trees tend to
have large evergreen leaves

• In chaparral, plants tend to have small
  sclerophyllous evergreen leaves
• Arid regions tend to support leafless stem
  succulents such as cacti or plants with entire
  leaf margins
• Cold wet climates tend to support plants with
  notched or lobed leaf margins

Lecture 11 27 February 2020

3
Plant Life FormsEvergreen vs. DeciduousMonolayer
ed vs Multilayered plantsShade
ToleranceXerophytic vs. Mesophytic leavesAlso
Hydrophytes (water lilies)
4
Foraging Tactics and Feeding EfficiencyCosts
and Profits of ForagingAn optimal foraging
tactic maximizes the difference betweenforaging
profits and their costs net benefitFood
matter and energy for maintenance and
reproductionHazards exposure to predators,
loss of time for other activitiesSit-and-Wait
ambush predators (e.g. spiders at webs)Widely
foraging active hunters (go out and find
prey)Search Time (per item eaten) versus
Pursuit Time (per item eaten)Search for all
possible prey items, but pursue them one at a
time Prey items can be ranked from most
preferred to least desirable
5
Economics of Consumer Choice
Assumptionsa) Environmental structure is
repeatable, with statistical expectation of
finding a given resource (habitat, microhabitat,
or prey item)b) Food items can be arranged
along a continuous spectrum, such as by size or
energy rewardc) Similar phenotypes are closely
equivalent in harvesting abilitiesd) Principle
of Allocation applies no one phenotype can be
maximally efficient on all prey typese) An
individuals economic goal is to maximize its
total intake of food resources
Robert MacArthur
6
Economics of Consumer Choice Four Phases
of Foraging 1) deciding where to search 2)
searching for palatable food items 3) upon
locating a potential food item, deciding
whether or not to pursue it 4) pursuit itself,
with possible capture and eatingSearch and
pursuit efficiencies for each food type in each
habitat are entirely determined by preceding
assumptions about morphology and environmental
repeatability. These efficiencies dictate
probabilities associated with search and pursuit
(phases 2 and 4) . Thus, need to consider only
the two decisions where to forage and which prey
items to pursue (phases 1 and 3 above)
Robert MacArthur
7
Economics of Consumer Choice (R. H.
MacArthur)Clearly, an optimal consumer should
forage where its expectationof yield is greatest
an easy decision to make, given knowledge of
efficiency probabilities and the structure of the
environment (of course, in reality, animals are
not omniscient and must make decisions based on
incomplete information).The decision as to
which prey items to pursue is also simple. Upon
finding a potential prey item, a consumer has
just two options either pursue it or go on
searching for a better item and pursue that one
instead. Both decisions end in the forager
beginning a new search, so the best choice is
clearly the one that returns the greatest yield
per unit time.An optimal consumer should opt to
pursue an item only when it cannot expect to
locate, catch and eat a better item during the
time required to capture and ingest the first
prey item.
8
When food is abundant, specialize but when food
is scarce, expand diet
9
From Huey and Pianka 1981 Ecology 62 991-999.
10
C. S. Holling
11
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Energetics of Metabolism and
Movement Ingestion Assimilation
Egestion Assimilation Productivity
Respiration Productivity
Growth Reproduction Ingestion Egestion
Respiration Growth Reproduction Assimi
lation------------------------------------------
------ Homeotherm versus Poikilotherm Endotherm
versus Ectotherm
13
Energetics of Movement
14
Metabolic Cost of Movement
Log,
Log,
15
Metabolic Cost of Movement
Log,
Log,
16
Adaptation and Deterioration of Environment
Ronald A. Fisher
Non-directed (random) changes in either A or B
are equally likely to reduce the level of
adaptation (d ) when small, but as the magnitude
of undirected change increases, the probability
of improvement diminishes. Duality of Fishers
model (A and B can be interchanged)
17
Water Economy in Desert OrganismsOther Limiting
MaterialsSensory Capacities and Environmental
CuesAdaptive SuitesDesign Constraints
18

Heat Budgets and Thermal Ecology
19
Thermoconformers
20
Passive Thermoconformer
Nephrurus laevissimus
21
Active Thermoregulator
Ctenophorus isolepis
22
Active Thermoregulator
Ctenophorus isolepis
Thermoconformer
23
Design Constraints

Thermoregulators
Ctenotus leae
Ctenotus Skinks
Thermoconformers
Ctenotus piankai
24
Thermoregulators
Thermoconformers

25
Design Constraint
Thermoregulators
Thermoconformers

26
Phrynosoma asio (Mexico)
27
Ecological Equivalent Thorny Devil Moloch
horridus (Australia)
28
Phrynosoma platyrhinos
Adaptive Suite
29
Adaptive Suite
30
Vital Statistics of Populations Deme
(Mendelian Population) Demography (assume all
individuals equal) Population
Parameters Mean and Variance Individual Popu
lation Male or Female Sex Ratio Has Babies
or not Birth Rates Alive or Dead Death
Rates Given Age Age Structure Fixed
Genotype Gene Frequencies Growth
Rates Density
31
Life Tables Discrete versus
Continuous Ages Pivotal Age assumption (age
classes) qx force of mortality (fraction
dying during age interval) qx age-specific
death rate Survivorship curves lx fraction
of initial cohort that survives to age x ly /
lx probability of living from age x to age y
Ex Expectation of further life
32
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33
Note 30-fold difference
34
Note 30-fold difference
Uncanny
35
Note 30-fold difference
Saturday Night Special
36
More Personal
(No Guns)
Saturday Night Special
37
More Personal
(No Guns)
Saturday Night Special
38
More Personal
(No Guns)
Saturday Night Special
39
More Personal
(No Guns)
Saturday Night Special
40
Testosterone
No Guns, More Personal
Saturday Night Special Lotsa Lead Flying Around
41
Life tables Horizontal versus vertical samples

Segment

Cohort
Age


Birth
Time

42
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Life Tables Survivorship
curves lx fraction of initial cohort that
survives to age x ly / lx probability of
living from age x to age y Ex Expectation
of further life
ly lx

ly lx

S
?
Ex
__
__
Ex dy
44
Type 1 Rectangular Type 2 Diagonal Type 3
Inverse Hyperbolic
Xantusia vigilis
sheep
Uta stansburiana
45
Xantusia vigilis
Eumeces fasciatus
Eumeces fasciatus
Sceloporus olivaceus
46
Fecundity, Tables of Reproduction mx
age-specific fecundity Two conventions
females only, or count both males and females
but weigh each as one-half (only progeny
entering age class zero are counted) Gross
reproductive rate (GRR) is the sum of mx over all
ages However, because some females will die
before having all their possible babies, must
calculate realized fecundity which is simply
lxmx (the fraction of females surviving times
their fecundity) Realized fecundity, lxmx, is
summed over all ages to get the Net Reproductive
Rate, R0 (also called the Replacement Rate of
the Population)
47
http//en.wikipedia.org/wiki/Quive
rfull http//www.oregonlive.com/kiddo/index.ssf/20
08/05/environmental_moms_stop_at_one.html
48
http//www.zo.utexas.edu/courses/T
HOC/breeders.html http//www.oregonlive.com/kiddo/
index.ssf/2008/05/environmental_moms_stop_at_one.h
tml
49
Capacity for Human Increase Realized (Mormons)
50
Pediculus humanus
51
T, Generation time average time from one
gener- ation to the next (average time
from egg to egg)
vx Reproductive Value Age-specific
expectation of all future offspring
52
In populations that are expanding or contracting,
reproductive value is more complicated. Must
weight progeny produced earlier as being worth
more in expanding populations, but worth less in
declining populations. The verbal definition is
also changed tothe present value of all future
offspring
53
Life tables Age-specific probability
statistics Force of mortality qx Survivorship
lx Fecundity mx Realized fecundity lxmx Net
reproductive rate R0 Generation time T
Reproductive value vx
54
vx mx ? (lt / lx ) mt Residual
reproductive value age-specific
expectation of offspring in distant
future vx ( lx1 / lx ) vx1
55
Illustration of Calculation of Ex, T, R0, and
vx in a Stable Population with Discrete Age
Classes __________________________________________
___________________________


Age Expectation Reproductive Weighted
of Life Value Survivor- Realized by
Realized Ex vx Age (x)
ship Fecundity Fecundity Fecundity
lx mx lxmx x
lxmx _____________________________________________
________________________ 0 1.0 0.0 0.00
0.00 3.40 1.00 1 0.8
0.2 0.16 0.16 3.00
1.25 2 0.6 0.3 0.18 0.36
2.67 1.40 3 0.4 1.0 0.40
1.20 2.50 1.65 4 0.4 0.6
0.24 0.96 1.50 0.65 5
0.2 0.1 0.02 0.10 1.00
0.10 6 0.0 0.0 0.00 0.00
0.00 0.00 Sums 2.2 (GRR) 1.00 (R0)
2.78 (T)
__________________________________________________
___________________ E0 (l0 l1 l2 l3 l4
l5)/l0 (1.0 0.8 0.6 0.4 0.4 0.2) /
1.0 3.4 / 1.0 E1 (l1 l2 l3 l4 l5)/l1
(0.8 0.6 0.4 0.4 0.2) / 0.8 2.4 / 0.8
3.0 E2 (l2 l3 l4 l5)/l2 (0.6 0.4
0.4 0.2) / 0.6 1.6 / 0.6 2.67 E3 (l3 l4
l5)/l3 (0.4 0.4 0.2) /0.4 1.0 / 0.4
2.5 E4 (l4 l5)/l4 (0.4 0.2) /0.4 0.6 /
0.4 1.5 E5 (l5) /l5 0.2 /0.2 1.0 v1
(l1/l1)m1(l2/l1)m2(l3/l1)m3(l4/l1)m4(l5/l1)m5
0.20.2250.500.30.025 1.25 v2
(l2/l2)m2 (l3/l2)m3 (l4/l2)m4 (l5/l2)m5
0.300.670.40 0.03 1.40 v3 (l3/l3)m3
(l4/l3)m4 (l5/l3)m5 1.0 0.6 0.05
1.65 v4 (l4/l4)m4 (l5/l4)m5 0.60 0.05
0.65 v5 (l5/l5)m5 0.1 _____________________
__________________________________________________
____
56
Illustration of Calculation of Ex, T, R0, and
vx in a Stable Population with Discrete Age
Classes __________________________________________
___________________________


Age Expectation Reproductive Weighted
of Life Value Survivor- Realized by
Realized Ex vx Age (x)
ship Fecundity Fecundity Fecundity
lx mx lxmx x
lxmx _____________________________________________
________________________ 0 1.0 0.0 0.00
0.00 3.40 1.00 1 0.8
0.2 0.16 0.16 3.00
1.25 2 0.6 0.3 0.18 0.36
2.67 1.40 3 0.4 1.0 0.40
1.20 2.50 1.65 4 0.4 0.6
0.24 0.96 1.50 0.65 5
0.2 0.1 0.02 0.10 1.00
0.10 6 0.0 0.0 0.00 0.00
0.00 0.00 Sums 2.2 (GRR) 1.00 (R0)
2.78 (T)
__________________________________________________
___________________ E0 (l0 l1 l2 l3 l4
l5)/l0 (1.0 0.8 0.6 0.4 0.4 0.2) /
1.0 3.4 / 1.0 E1 (l1 l2 l3 l4 l5)/l1
(0.8 0.6 0.4 0.4 0.2) / 0.8 2.4 / 0.8
3.0 E2 (l2 l3 l4 l5)/l2 (0.6 0.4
0.4 0.2) / 0.6 1.6 / 0.6 2.67 E3 (l3 l4
l5)/l3 (error extra terms) 0.4 0.4 0.2)
/0.4 1.0 / 0.4 2.5 E4 (l4 l5)/l4
(error extra terms) 0.4 0.2) /0.4 0.6 / 0.4
1.5 E5 (l5) /l5 0.2 /0.2 1.0 v1
(l1/l1)m1(l2/l1)m2(l3/l1)m3(l4/l1)m4(l5/l1)m5
0.20.2250.500.30.025 1.25 v2
(l2/l2)m2 (l3/l2)m3 (l4/l2)m4 (l5/l2)m5
0.300.670.40 0.03 1.40 v3 (l3/l3)m3
(l4/l3)m4 (l5/l3)m5 1.0 0.6 0.05
1.65 v4 (l4/l4)m4 (l5/l4)m5 0.60 0.05
0.65 v5 (l5/l5)m5 0.1 _____________________
__________________________________________________
____
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Life tables Age-specific probability
statistics Force of mortality qx Survivorship
lx ly / lx probability of living from age x
to age y Fecundity mx Realized fecundity at
age x lxmx Net reproductive rate R0 ? lxmx
Generation time T ? xlxmx Reproductive value
vx ? (lt / lx ) mt
Ex Expectation of further life
60
Life tables and tables of fecundity
Age-specific (and sex-specific) probability
statistics Force of mortality qx Survivorship
lx Fecundity mx Realized fecundity lxmx Net
reproductive rate R0 Generation time T S xlxmx
Reproductive value vx (stable, non-stable
poplns) Residual reproductive value vx Stable
age distributions
61
Stable age distribution Stationary age
distribution
62
Intrinsic rate of natural increase (per
capita) r b dwhen birth rate exceeds
death rate (b gt d), r is positivewhen death
rate exceeds birth rate (d gt b), r is
negativeEulers implicit equation S?e-rx lxmx
1(solved by iteration)If the Net
Reproductive Rate R0 is near one, r loge R0
/T
63
When R0 equals one, r is zero When R0 is
greater than one, r is positiveWhen R0 is less
than one, r is negative Maximal rate of
natural increase, rmax
64
Estimated Maximal Instantaneous Rates of
Increase (rmax, per capita per day) and Mean
Generation Times ( in days) for a Variety of
Organisms ________________________________________
___________________________ Taxon Species
rmax Generation Time (T) ----------------------
--------------------------------------------------
----------------------------- Bacterium Escherich
ia coli ca. 60.0 0.014 Protozoa Par
amecium aurelia 1.24
0.330.50 Protozoa Paramecium caudatum 0.94
0.100.50 Insect Tribolium confusum
0.120 ca.
80 Insect Calandra oryzae 0.110(.08.11)
58 Insect Rhizopertha dominica 0.085(.07.10)
ca. 100 Insect Ptinus tectus 0.057 102 Inse
ct Gibbum psylloides 0.034 129 Insect Trigonog
enius globulosus 0.032 119 Insect Stethomezium
squamosum 0.025 147 Insect Mezium
affine 0.022 183 Insect Ptinus
fur 0.014 179 Insect Eurostus
hilleri 0.010 110 Insect Ptinus
sexpunctatus 0.006 215 Insect Niptus
hololeucus 0.006 154 Mammal Rattus
norwegicus 0.015 150 Mammal Microtus
aggrestis 0.013 171 Mammal Canis
domesticus 0.009 ca.
1000 Insect Magicicada septendecim 0.001
6050 Mammal Homo sapiens 0.0003
ca. 7000 _______________________________________
___________________________ _
65
J - shaped exponential population growth
66
Instantaneous rate of change of N at time t is
total births (bN) minus total deaths (dN)dN/dt
bN dN (b d )N rNNt N0 ert
(integrated version of dN/dt rN)log Nt log
N0 log ert log N0 rtlog R0 log 1 rtr
log l or l er
67
y c mx
log Nt log N0 rt

NT
Slope r
Log N
N0
t
Time
68
Demographic and Environmental Stochasticity
random walks, especially important in small
populations Evolution of Reproductive
Tactics Semelparous versus Interoparous
Big Bang versus Repeated Reproduction
Reproductive Effort (parental investment)
Age of First Reproduction, alpha, a Age of
Last Reproduction, omega, v
69
Mola mola (Ocean Sunfish) 200 million
eggs!
Poppy (Papaver rhoeas) produces only 4 seeds
when stressed, but as many as 330,000 under
ideal conditions
70
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71
How much should an organism invest in any given
act of reproduction? R. A. Fisher (1930)
anticipated this question long ago It would
be instructive to know not only by what
physiological mechanism a just apportionment is
made between the nutriment devoted to the gonads
and that devoted to the rest of the parental
organism, but also what circumstances in the life
history and environment would render profitable
the diversion of a greater or lesser share of
available resources towards reproduction.
Italics added for emphasis.
Reproductive Effort
Ronald A. Fisher
72
Asplanchna (Rotifer)
73
Trade-offs between present progeny and
expectation of future offspring
74
Iteroparous organism
75
Iteroparous organism
76
a
w
77
http//www.commondreams.org/view/2011/03/07-0
Microtus
78
Chittys Genetic Control Hypothesis
Could optimal reproductive tactics be involved in
driving population cycles?
Dennis Chitty
79
Semelparous organism
80
Semelparous organism
81
mx
Age, x
82
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83
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84
Patterns in Avian Clutch SizesAltrical versus
Precocial
85
Patterns in Avian Clutch SizesAltrical versus
PrecocialNidicolous vs. NidifugousDeterminant
vs. Indeterminant Layers
N 5290 Species
86
Patterns in Avian Clutch SizesOpen Ground
Nesters Open Bush Nesters Open Tree Nesters
Hole Nesters
MALE
(From Martin and Ghalambor 1999)
87
Daylength HypothesisPrey Diversity
HypothesisSpring Bloom or Competition
HypothesisNest Predation Hypothesis
(Skutch)Hazards of Migration HypothesisEvolution
of Senescence

• recession of time of expression of the overt
  effects of a detrimental allele
• precession of time of expression of the effects
  of a
• beneficial allelle

88
Joint Evolution of Rates of Reproduction and
Mortality
Sceloporus
Donald Tinkle
89
Inverse relationship between rmax and generation
time, T Threshold of Annuality
90
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