Ch

5: Popula,on Ecology

Learning Objec,ves
A"er studying this chapter, you should be able to:
1. Know what a popula,on is, dierent ways popula,ons can be distributed, and factors aec,ng distribu,on. 2. Understand the various types of popula,on growth and how to calculate growth rates and popula,on size. 3. Know the factors that limit popula,on growth. 4. Understand the dierent paJerns of survival, reproduc,on and reproduc,ve rates and tools ecologists use to study these.

Introduc,on
Popula,on
A group of individuals of a single species that co-occur in space and ,me.

Characterized by
Distribu,on: size, loca,on Density: the number of individuals per unit area
Addi,onal characteris,cs of a popula,on include age distribu,ons, growth rates, and abundance.

Distribu,on Limits
Physical environment limits geographic distribu,on of a species.
Organisms can only compensate so much for environmental varia,on.

Distribu,on of Barnacles along an Inter,dal Exposure Gradient

Organisms living in an inter,dal zone have evolved to dierent degrees of resistance to drying. Barnacles show dis,nc,ve paJerns of zona,on within inter,dal zone.
Connell found Chthamalus stellatus restricted to upper levels while Balanus balanoides is limited to middle and lower levels.

Distribu,ons of Barnacles along an Inter,dal Gradient

Balanus appears to be more vulnerable to desicca,on, excluding it from the upper inter,dal zone. Chthamalus adults appear to be excluded from lower areas by compe,,on with Balanus.
Barnacle-covered rocks at low ,de, Monterey Na,onal Seashore

Photo: J. Kerr

Distribu,on of Individuals
Random: Equal chance of being anywhere.
Uniform distribu,on of resources.

Regular: Uniformly spaced.

Exclusive use of areas. Individuals avoid one another.

Clumped: Unequal chance of being anywhere.

Mutual aJrac,on between individuals. Patchy resource distribu,on.

Distribu,on of Individuals

Poisson Distribu,on

# squares 1 2 3 4 5 6 7 8 9 10 # confeZ

Clumped -Probably around certain habitat features -Deviates from Poisson

Uniform -Possibly represen,ng territories -Deviates from Poisson

Creosote bush root distribu,ons

Do they compete below ground? Brisson and Trynolds (1994) Careful excava,ons and root mapping

biology.ed.ac.uk

Dr. Susan Dudley

Popula,on Growth or Decline

Popula,ons change. Popula,ons may increase or decrease in the short term and in the long term. Understanding factors behind popula,on growth and decline is important for nature conserva,on and resource management.

Closed and Open Popula,ons

In closed popula,ons, changes in abundance (N) are determined by births (M) and deaths (D). In open popula,ons, changes are further inuenced by emigra,on (E) and immigra,on (I). Nt+1 = Nt + (Mt Dt) + (It Et) Change in abundance over ,me is described by a rate. Time frame of measurement can be con,nuous or discrete.

Con,nuous Time Popula,on Growth

When ,me dierence between (t+1) and (t) is innitesimally short. Birth rate (m) = M/N (#births/popula,on size) = e.g. 100/500 = 0.2 births/ind/unit ,me Death rate (d) = D/N (#deaths/popula,on size) = e.g. 200/500 = 0.4 deaths/ind/unit ,me

Con,nuous Time Popula,on Growth

Popula,on growth rate determined by m and d. r = m d (intrinsic rate of popula,on growth)

For an exponen,ally growing popula,on r does not change with either 2me or popula2on size.
Termed the maximum per capita rate of growth (rmax)

For an exponen,ally growing popula,on r = rmax

15.9

1.78 1.08 0.60

0.27 0.12 Year (t)

dN/dt = rmaxN

Calcula,ng the popula,on at some later ,me (t)

For a popula,on growing con2nuously at an exponen,al rate, the popula,on size at any ,me t can be calculated as:

Nt = N0ermaxt

Should be able to subs,tute with numbers and calculate popula,on size at specied ,me.

Where: Nt = popula,on size at some later ,me (t) N0 = ini,al popula,on size rmax = intrinsic rate of increase t = ,me

Doubling ,me
Can rearrange the previous equa,on to determine the doubling ,me
Time for the popula,on to go from N0 to 2N0.

tdouble = ln(2)/rmax

Time for the popula,on to go from N0 to 3N0.

ttriple = ln(3)/rmax

Discrete Time Exponen,al Growth

Discrete pulses of births and deaths Reproduc,on and mortality omen seasonal. N t = N0t

Nt = number of individuals at ,me t N0 = ini,al number of individuals = nite rate of increase t = number of ,me intervals or genera,ons

 r < 0 declining r = 0 stable r > 0 increasing

< 1 declining = 1 stable > 1 increasing

Grey Seals on Sable Island

Grey seals were severely diminished by bounty hun,ng and other factors. Recovery on Sable Islands, Nova Sco,a, since 1960. Exponen,al growth observed.

Grey seal (Halichoerus grypus)

Fig. 5.2a, p. 100

Human Popula,on
1 billion for the rst ,me in 1825. Adding 1 billion people every 13 years.

Human Popula,on Growth

In 1975, the popula,on was growing at a rate of nearly 2% every year (doubling every 35 years).
At this rate, we would reach 32 billion by 2080.

But growth rate has slowed recently, to about 1.21% per year.
If this rate is maintained, there would be roughly 16 billion people on Earth in 2080.

Could Earth support 16 billion people? Ecological maxim: No popula2on can increase in size forever
There are limits imposed by a nite planet that restricts growth Ecologists try to understand the factors that limit or promote popula,on growth

Logis,c Popula,on Growth (S-curve)

As resources are depleted, popula,on growth rate slows and eventually stops: logisHc populaHon growth Sigmoid (S-shaped) popula,on growth curve. Carrying capacity (K) is the number of individuals of a popula,on that the environment can support. Finite amount of resources can only support a nite number of individuals.

Logis,c Popula,on Growth

dN/dt = rmaxN(1 N/K)
rmax = maximum per-capita rate of increase under ideal condi,ons. When N nears K, the right side of the equa,on nears zero. As popula,on size increases, popula,on growth rate becomes smaller. Highest when N = K/2 (seen in the logis,c curve).

Limits to Popula,on Growth

Good to recognize that this is only a mathema,cal model not a law The environment limits popula,on growth by altering birth and death rates through Density-dependent factors
disease, resource compe,,on

Density-independent factors
natural disasters

In nature, both types of factors constantly at play (e.g. case study of Galapagos nches)

Galapagos Finch Popula,on Growth

Boag and Grant: Geospiza for2s was numerically dominant nch (1,200). Amer drought of 1977, popula,on fell to 180.
Food plants failed to produce seed crop.

In 1983, 10x normal rainfall caused popula,on to grow (1,100) due to abundance of seeds and caterpillars.

Galapagos Finch Popula,on Growth

Age-Specic Fecundity
Fecundity = the number of ospring produced by an individual during a breeding season. Birth rates are age-specic Popula,on birth rate depends on popula,on structure Determinate growth (stops growing amer certain age): fecundity is constant Indeterminate growth (con,nues to grow): fecundity increases with age, un,l senescence

Bighorn sheep ewes give birth to one lamb, but not every year. Popula,on birth rate is calculated based on age- specic probabili,es of breeding.
Iteroparous vs. Semelparous

Age-Specic Survival
Probability of death (and survival) depends on age of organism Three general types of survivorship curves:
Type I: Low mortality except at old age Type II: Constant rate of mortality Type III: High mortality rate for young individuals
Type I Survival ln [lx]

Type II Type III

Age (x)
Examples: (I) Bighorn sheep, (II) black- capped chickadee and (III) Atlan,c cod.

Fig. 5.12, p. 108

Es,ma,ng PaJerns of Survival

Cohort life table
Iden,fy individuals born at same ,me and keep records from birth.

Sta,c life table

Data collected from individuals of all ages at one point in ,me.
E.g. Sample all individuals of all age classes at once (age distribu,on).

High Survival Among the Young

Murie collected Dall sheep skulls, Ovis dalli.
Major Assump,on: Propor,on of skulls in each age class represented typical propor,on of individuals dying at that age.
Reasonable given sample size of 608.

Constructed a survivorship curve.

Discovered bi-modal mortality.
<1 yr. 913 yrs.

Age Distribu,on
Reects a popula,ons history of survival, reproduc,on, and growth poten,al. Ecological popula,on dynamic models improve forecasts of how human popula,ons change
A countrys age distribu,on, birth rate, and death rate strongly aect its projected growth rate Popula,on pyramids

China
China popula,on 600 million to 1 billion 1950-1975 Government wanted zero popula,on growth by 2000 One-child policy of the late 1980s Encouragement (1 child couples)
Extra food, housing, pensions, medicare Free contracep,ves, steriliza,on, abor,ons Enforced abor,on of 3rd child

Results
Female infan,cide/spoiled boys Social Human rights abuse

Age Distribu,on

Using Life Tables to Es,mate Popn Growth

Survival (lx)
Propor,on surviving at start of age x.

Age-specic fecundity (mx)

Number of young born per female. In a period of ,me.

Es,ma,ng Rates for an Annual Plant

Phlox drummondii
R0 = net reproduc,ve rate; average number of ospring (seeds) produced by an individual in a popula,on during its life,me.

R0= lxmx
usbg.gov

x = age interval in days lx = propor,on pop. surviving to each age (x) mx = average number seeds produced by each individual in each age category

Es,ma,ng Rates when Genera,ons Overlap

Northern water snake (Nerodia sipedon) Average genera,on ,me: T = xlxmx R0 x = age in years Per Capita Rate of Increase: r

ln = base of the natural logarithm

ln R0 T

Per capita rate of increase

r = ln R0 T r = ln (0.787)/5.04 r = -0.048

r < 0 r = 0 r > 0

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