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ECOLOGICAL
METHODS
HANDBOOK
by

Chris Butler, Ph.D., and


David Bass, Ph.D
Biology Department
University of Central Oklahoma

Table of Contents

Syllabus

Week 1

Week 2

14

Week 3

24

Week 4

30

Week 5

50

Week 6

68

Week 7

81

Week 8

82

Week 9

83

Week 10 93
Week 11 96
Week 12 101
Week 13 123
Week 14 143
Week 15 144

BIO 3551 Ecological Methods

Description

Objectives
1.
2.
3.
4.
5.

This course introduces students to field, laboratory, and


computer-based methods in ecology. It includes the
study of abiotic and biotic components of terrestrial and
aquatic ecosystems. This course emphasizes common
methods used in modern ecological studies of terrestrial
and aquatic environments. It consists of three hours of
laboratory per week, and many exercises will involve
field trips. For the Central Six, this class will advance
discipline knowledge, problem solving (by engaging in
research) and health and wellness (by engaging in field
biology outside). Prerequisite(s): BIO
1204, 1225, 3543 and STAT 2103 all with a minimum
grade of C.

Learn ecological methods used to collect data in laboratory and field


studies.
Investigate local terrestrial and aquatic environments through the proper
collection of data.
Demonstrate the proper use of laboratory and field equipment.
Analyze data collected during class exercises.
Report results of the analyses.

Instructor
Dr. Chris Butler
301-G Howell Hall
x5782
cbutler11@uco.edu
Class meets
Wednesdays 1:00-3:50 PM. Note that many classes will meet in the field the instructor will
let you know one week in advance where we will meet. Be sure to wear appropriate attire for the field.

Required Reading

Butler, C and D. Bass. 2011. Ecological Methods Handbook. Available from


http://www.biology.uco.edu/personalpages/CButler/BIO3551/Ecological_Methods_Lab_Manua
l.pdf.

Assignments and Grading


Attendance: 50 pts
Mid-Term Exam: 200 pts
Final: 200 pts
Assignments: 440 pts
Total: 900 pts
Course Calendar
Week 1 Introduction, Plant walk
Week 2 Aquatic sampling
Week 3 Comparison of lentic vs. lotic systems
Week 4 Quadrat-based community sampling in a riparian community
Week 5 Quadrat-based community sampling in a cross-timbers community
Week 6 Quarter-based community sampling
Week 7 Mid-term Exam
Week 8 FALL BREAK
Week 9 Terrestrial invertebrates
Week 10 Mammals
Week 11 Birds
Week 12 Ecobeaker Isle Royale
Week 13 Ecobeaker - Keystone
Week 14 THANKSGIVING BREAK
Week 15 Review
Week 16 Final exam

Additional Information

ECOLOG-L listserv
The Ecological Society of America has a listserv devoted to the discussion of ecological issues as well as
posting ecology-related research opportunities.
What is a listserv? It's basically a loose network of e-mail users interested in a particular subject, in this
case ecology. Once you are subscribed (it's free) you will be able to send a ecology-related question,

observation, or message to a single address and it will quickly and automatically be delivered to all other
subscribers to the list. Any replies to your message will also be delivered to all subscribers.
To subscribe, google ECOLOG-L. The first option (https://listserv.umd.edu/archives/ecolog-l.html) will
take you to the archives where you will see a link, Join or leave the list (or change settings). Clicking on
the link will take you to a page where you can subscribe to ECOLOG-L. You may also unsubscribe at any
time.

Name:

Week 1 - Plant Walk


Due at the end of class
Common Name
1.

Field marks

Sketch

2.

3.

4.

Common Name
5.

Field marks

Sketch

6.

7.

8.

9.

Common Name
10.

Field marks

Sketch

11.

12.

13.

14.

Common Name
15.

Field marks

Sketch

16.

17.

18.

19.

Ecological Methods Plant Guide

American Sycamore
Platanus occidentalis

American Sycamore
Platanus occidentalis

Eastern Redcedar
Juniperus virginiana

Eastern Redbud
Cercis canadensis

Pecan
Carya illinoinensis

Pecan
Carya illinoinensis

Slippery Elm
Ulmus rubra

Black Locust
Robinia pseudoacacia

White Mulberry
Morus alba

10

Sugarberry
Celtis laevigata

Honeylocust
Gleditsia triacanthos

Greenbrier
Smilax sp.

Trumpet Creeper
Campsis radicans

Honeylocust
Gleditsia triacanthos

Virginia Creeper
Parthenocissus quinquefolia

Trumpet Creeper
Campsis radicans

Ragweed
Ambrosia sp.

Sandbar Willow
Salix interior

11

Eastern Poison Ivy


Toxicodendron radicans

Grape sp.
Vitis sp.

Eastern Cottonwood
Populus deltoides

Chinkapin Oak
Quercus muehlenbergii

Baldcypress
Taxodium distichum

Blackjack Oak
Quercus marilandica

Post Oak
Quercus stellata

Black Walnut
Juglans nigra

Black Walnut
Juglans nigra

12

Silktree (Mimosa)
Albizia julibrissin

Common Hackberry
Celtis occidentalis

American Elm
Ulmus americana

Roughleaf Dogwood
Cornus drummondii

Red Mulberry
Morus rubra

Gum Bully (Chittamwood)


Sideroxylon lanuginosum

Green Ash
Fraxinus pennsylvanica

13

Lab 2 Aquatic Sampling

COMMON WATER SOURCES IN OKLAHOMA


To understand the ecology of an area or region one must be familiar with the water sources of the area.
Rivers not only provide a ready source of water for the surrounding biota but may also represent a barrier to
dispersal. Large reservoirs may also influence the local climates of an area as well as alter the life forms of
the water course from which they were constructed.
In Oklahoma there are approximately 500 named rivers and creeks, many of them short and intermittent
during most of the year. There are several large streams like the Arkansas, Red, Washita, Cimarron, North
Canadian, Canadian, Grand, and Verdigris, however, that carry millions of cubic feet of water through the
state each year. All of northern Oklahoma and much of the central part of the state is in the drainage basin
of the Arkansas River. The remainder of Oklahoma is in the drainage basin of Red River. Both are long
streams, the source of the Arkansas being in the Rocky Mountains of Colorado and that of the Red on the
High Plains of Texas. Parts of both rivers were used in defining the Boundaries of the Adams-Onis Treaty of
1819, which set definite limits of Spanish and American territory.
Except for the rivers flowing from the Ozark Plateau or the Ouachita Mountains, the streams of Oklahoma
flow in a general eastward direction. The chief tributaries of the Arkansas are the Cimarron and Canadian
from the west, the Verdigris, Grand, and Illinois from the north and northeast, and the Poteau from the
south. The North Canadian, one of the longest streams in the state, is the chief tributary of the Canadian.
Principal tributaries of the Red River are the North Fork, Washita, Blue, Boggy, and Kiamichi.
Rivers have been of primary importance in the development of Oklahoma. The Canadian, Arkansas, North
Canadian, and many others formed parts of the boundaries of the Indian nations. Some served as routes of
travel for early trails and others as sites for the location of pioneer settlements. A few appear in historical
records under different names. The Cimarron has been known as the Red Fork of the Arkansas. The Little
River of Cleveland and Pottawatomie counties were called the Cedar River by the Seminoles. The river now
commonly called the North Canadian has been known as the Rio Nutrio and as the North Fork of the
Canadian. The name North Canadian is often applied to Beaver Creek in the Panhandle, but the earliest
maps show the North Fork of the Canadian to be formed by the confluence of Beaver and Wolf Creeks. The
river now commonly called the South Canadian in Oklahoma is referred to the New Mexico and Texas, as
well as in most government publications, as the Canadian. Gaines Creek, in Pittsburg County, now a part of
the Eufaula Reservoir, was known as the South Canadian by the early Indian settlers.
All of the large lakes in Oklahoma are man-made. They have been developed for such purposes as flood
control, conservation, navigation, irrigation, recreation, production of hydroelectric power, and municipal
water supply. In addition to these large reservoirs, many stock ponds also exist.

14

Physicochemical Analysis
Dissolved Oxygen (YSI Model 55 DO Meter)
1.
2.
3.
4.

Turn on meter, leaving probe in the damp compartment.


Allow meter to remain on for about 5 minutes before calibrating meter.
To calibrate meter, press and release both the Up Arrow and Down Arrow simultaneously.
After calibration is complete, place probe in sample to read both temperature and dissolved oxygen
concentration.

pH (using digital meter)


1. Remove cap and immerse probe into sample.
2. Turn on switch and swirl meter for about 10 seconds, allowing reading to stabilize.
3. Record value and turn off switch.
Specific Conductivity
1.
2.
3.
4.
5.
6.

Connect probe to meter.


Rinse the probe with distilled water.
Switch the meter on.
Immerse the probe in the water sample.
Read the conductivity from the meter in micromhos / cm.
Record the value and turn off the meter.

Plankton Net
1. Tow net through the water at desired sampling depth or pour water from that depth through net.
2. Record distance towed or volume of water poured through the net.
3. Rinse contents of the net into a jar and preserve the sample.
Petite Ponar Bottom Grab
1.
2.
3.
4.
5.
6.

Lock open the jaws of the Ponar grab.


Lower the Ponar grab to the bottom substrate.
As you pull upward the jaws close and a 6" x 6" sample is taken.
Empty the sample into a sieve bucket and wash away the silt.
Carefully transfer the sample to an appropriate container.
Preserve and label the sample.

Dip Net
1.
2.
3.
4.

Sweep the net through the water three times. Make sure to disturb the substrate to a depth of one inch
Empty the sample to a sieve bucket by turning the net inside out in the bucket.
Carefully transfer the sample to an appropriate container.
Preserve and label the sample.
15

1. Complete the following table (physicochemical conditions).


Parameter

Sample 1

Sample 2

Sample 3

Sample 4

Sample 5

Depth (m)
Water
Temperature
(oC)
Dissolved
Oxygen (mg/l)
pH
Conductivity

16

2. Construct a species list of benthic macroinvertebrates


(Morpho)Species
Number

Method

3. Which method resulted in the greatest number of morphospecies?

17

4.) The Shannon-Weiner index is calculated as


= ln( )
For example, imagine a hypothetical scenario
Organisms
# found pi
ln(pi)
Orthetrum
3 0.166666667 -1.791759469
Somatochlora
1 0.055555556 -2.890371758
Corxidae
2 0.111111111 -2.197224577
Enonchis
2 0.111111111 -2.197224577
Peltodytes
7 0.388888889 -0.944461609
Chrionomi
1 0.055555556 -2.890371758
Tonypodinae
2 0.111111111 -2.197224577
18
H'
The Shanon-Weiner index for this site is 1.72.

pi*ln(pi)
-0.298626578
-0.160576209
-0.244136064
-0.244136064
-0.367290626
-0.160576209
-0.244136064
-1.719477814
1.719477814

Calculate the Shannon-weiner Index for your aquatic samples.

5.) Simpsons Diversity Index is calculated as


1 D where =

(1)
(1)

and where n = the total number of organisms of a species while N = the total

number of organisms for all species.


For example, imagine a hypothetical scenario
Organisms
n
n(n-1)
Water boatman
1
0
Hempitera
1
0
Water strider
2
2
Bivalvia
8
56
Gastropoda
3
6
Shiner
5
20
Sunfish
1
0
Sum
21
84
So D =

84
21(20)

= 0.2
18

And Simpsons Diversity Index = 1 D = 0.8.


Calculate Simpsons Diversity Index for your aquatic samples.

19

Aquatic Sampling Equipment


Know the names and how to use this equipment

Conductivity meter

Dissolved oxygen meter

Dip net

Mechanical flowmeter

Seine

Stream drift net

20

Magnifying loupe

pH meter

Ponar grab

Spectrometer

Plankton net
Kick net

21

Organisms from Arcadia Lake Site

Phylum: Arthropoda
Class: Insecta
Order: Diptera
Midge larva

Phylum: Arthropoda
Class: Insecta
Order: Diptera
Midge larva

Phylum: Arthropoda
Class: Insecta
Order: Hemiptera
Water boatman

Phylum: Arthropoda
Class: Insecta
Order: Coleoptera
Water scavenger beetle

Phylum: Arthropoda
Class: Insecta
Order: Coleoptera
Beetle sp.

Phylum: Arthropoda
Class: Insecta
Order: Coleoptera
Crawling water beetle

Phylum: Arthropoda
Class: Insecta
Order: Odonata
Dragonfly nymph

Phylum: Arthropoda
Class: Insecta
Order: Odonata
Dragonfly nymph

22

Organisms from Chisholm Creek Site

Phylum: Arthropoda
Class: Insecta
Order: Coleoptera
Predaceous Diving Beetle

Phylum: Arthropoda
Class: Insecta
Order: Ephemeroptera
Mayfly nymph

Phylum: Arthropoda
Class: Insecta
Order: Hemiptera
Water strider

Phylum: Arthropoda
Class: Insecta
Order: Hemiptera
Toad bug

Phylum: Mollusca
Class: Gastropoda
Order: Pulmonata
Pond Snail

Phylum: Mollusca
Class: Gastropoda
Order: Pulmonata
Snail sp.

Phylum: Chordata
Class: Actinopterygii
Order: Perciformes
Sunfish

Phylum: Chordata
Class: Actinopterygii
Order: Cypriniformes
Shiner

Phylum: Mollusca
Class: Bivalvia
Order: Veneroida
Asiatic clam

23

Lab 3 Comparison of Lentic and Lotic Systems


Lentic systems are still-water systems such as ponds, lakes, wetlands, etc. Lotic systems are flowingwater systems such as streams, rivers, springs, etc. For this assignment, you are going to compare the
two systems using the data you gathered this week and last week. Before you begin, copy your values
from last week into the tables below.
1.) Does the amount of dissolved oxygen differ between lentic and lotic systems? Use a t-test to
answer this question (cite appropriate statistics!) and create a figure showing means and
standard errors for dissolved oxygen.
a. Appropriate statistics include t-value, df, and p-value)
i. Your sentence will say something like, Dissolved oxygen was significantly higher
in the lotic system (t = ____, df = ______, p = ______; Fig. 1).
1. Note that it is only appropriate to say that there is a significant
difference if your p-value is less than 0.05.
ii. Directions for performing a t-test in Excel can be found at
http://www.excel-easy.com/examples/t-test.html
b. To create a bar graph with standard error bars, you will first need to calculate mean and
standard error.
i. Directions for calculating descriptive statistics (including mean and standard
error) can be found at
http://www.excel-easy.com/examples/descriptive-statistics.html
c. Directions on creating a bar graph with custom error bars can be found at
http://nathanbrixius.wordpress.com/2013/02/11/adding-error-bars-to-charts-in-excel2013/
i. Note that you will be using the standard error values you calculated above,
rather than the 95% confidence intervals in the example.
2.) Does the pH differ between lentic and lotic systems? Use a t-test to answer this question (cite
appropriate statistics!) and create a figure showing means and standard errors
3.) Does the conductivity differ between lentic and lotic systems? Use a t-test to answer this
question (cite appropriate statistics!) and create a figure showing means and standard errors
4.) Does the species diversity differ between the two systems? Calculate the Shannon-Weiner index
for each system to answer this question.
5.) Does the species diversity differ between the two systems? Calculate Simpsons index for each
system to answer this question.

24

Lake Arcadia
Parameter

Sample 1

Sample 2

Sample 3

Sample 4

Sample 5

Chisholm Creek
Parameter
Sample 1

Sample 2

Sample 3

Sample 4

Sample 5

Depth (m)
Water
Temperature
(oC)
Dissolved
Oxygen (mg/l)
pH
Conductivity

Depth (m)
Water
Temperature
(oC)
Dissolved
Oxygen (mg/l)
pH
Conductivity

25

Lake Arcadia
(Morpho)Species

Number

Method

26

Chisholm Creek
(Morpho)Species

Number

Method

27

Lentic
(from
wikipedia.org)

28

Lotic
(from
wikipedia.org)

29

Lab 4 Quadrat-based Community Sampling in a Riparian


Community

One of the most commonly used methods of vegetation sampling is the quadrat method. This method involves
recording all plants within square or rectangular plots in a stand and results in a quantitative description of
vegetation for that stand.
There are two ways that quadrats may be located. One is by superimposing a grid on a map of the site,
numbering each section of the grid, and randomly selecting sections to be sampled. The other method, often
used by experienced field workers, is to subjectively place quadrats in areas that are felt to best represent, and
cover the variation within, the study area. Because of time limitations, we will use the latter method. It is
important that each individual quadrat be fairly homogeneous in its composition. The preferred quadrat size
varies from one region to another, with a 1/40 acre (33 feet by 33 feet) or a hectare (10,000m2) plot being
commonly used in temperate zone forests, and much smaller plots (1m2 or 0.25m2) being used in grasslands or
prairies. Circular quadrats are also used.
Materials
Tape measurers
Compass
Data sheets
Calculators
Field Methods
We will establish several square quadrats, each 10m x 10m (100m2 or 0.01 hectare), in a forest. The class will be
divided into teams, each being responsible for collecting data for one or two quadrats. Measure out 10m using a
tape measure to serve as a side of the square. Then use the tape to measure the remaining three sides. Mark
the corners and edges of your quadrat with stakes or flags.
Record relevant information about the quadrat, including the quadrat location and number, sampling date, and
investigators. Briefly describe the topography and landscape, including the slope, aspect, and location relative to
streams. Soil depth, drainage, texture, soil type, soil pH, and parent material should also be described, but we
will not deal with those during this lab period. Any evidence of disturbance should also be noted, including
logging, fire, or grazing, as well as old fences and roads, tree falls, or obvious diseases of dominant species.
Record all tree species for each quadrat on the data sheet (Table 1). One person in each group should record
data in the field for that group and share it with the rest of the group after returning to the laboratory. Measure
the DBH (circumference) for each individual tree and record it. To be counted as a tree, an individual plant

30

should have a circumference of 5cm and stand >2m tall. For this exercise, we will use a standard tape measurer
and record the circumference (in cm) on the data sheet. If your tape measure is in inches, multiply by 2.54 to
obtain the circumference in centimeters.
There are several things to keep in mind as you collect your data. If a tree is dead, do not measure and record its
circumference -- simply ignore it. If a tree exists on a boundary, count it only if more than one-half of the trunk
lies within your quadrat. If a tree has two or more main trunks, measure the circumference of each trunk
separately, sum those values, and count them as one tree. Take all measurements carefully and accurately!
Calculations
A)

Convert DBH (on Table 1 data sheet) to basal area in cm2 for each tree. Use the equation:
Basal area = Pi x (0.5 x DBH)2
Total the basal area covered by all trees of each species to obtain a total basal area per species.

B) Each member of each team should calculate the following for each tree species (list by scientific name) for
the quadrat they sampled and list those results in Table 2. Be sure to show your work as done in the
example!
1)

Density = Number of trees

2)

Total basal area (totaled in as above)

3)

Relative density = No. of trees of one species / Total no. of trees of all species

4)

Relative dominance = Total basal area of one species / Total basal area of all species

5)

Importance value = (Relative density + Relative dominance) / 2

C) We will pool the data from all quadrats, calculate the following for each tree species, and record those
values on Table 3.
1)

Number of quadrats of occurrence

2)

Density = Number of trees

3)

Total basal area

4) Relative frequency = No. quadrats of occurrence for a species / Total no. of quadrats of occurrence for
all species
5)

Relative density = No. of trees of a species / Total no. of trees of all species

6)

Relative dominance = Total basal area of a species / Total basal area of all species

7)

Importance value = (Relative frequency + Relative density + Relative dominance) / 3


31

8)

Total number of trees of all species combined per hectare (a hectare is 10,000m2)

9)

Total basal area all species combined per hectare

32

Table 1 - Transect Method Field Data (#1)

Location ________________________________________

Number of saplings in Quadrat ______________

Date _______________________________

Number of seedlings in Quadrat ______________

Members of group ___________________________________________________________________________

Number

Species

DBH

Basal Area Coverage


(Circumference)

33

34

Table 1 - Transect Method Field Data (#2)

Location ________________________________________

Number of saplings in Quadrat ______________

Date _______________________________

Number of seedlings in Quadrat ______________

Members of group ___________________________________________________________________________

Number

Species

DBH

Basal Area Coverage


(Circumference)

35

36

Table 1 - Transect Method Field Data (#3)

Location ________________________________________

Number of saplings in Quadrat ______________

Date _______________________________

Number of seedlings in Quadrat ______________

Members of group ___________________________________________________________________________

Number

Species

DBH

Basal Area Coverage


(Circumference)

37

38

Table 2 - Summary of Community Analysis of Single Quadrat by the Quadrat Sampling Technique (#1)

Location ________________________________________

Quadrat Number ______________

Date _______________________________

Quadrat Size ______________

Members of group ___________________________________________________________________________

Species

Density

Total Basal
Area Covered

Relative Density (D)

Relative Dominance (Do)

Importance
(D + Do) / 2

39

Species

Density

Total Basal
Area Covered

Relative Density (D)

Relative Dominance (Do)

Importance
(D + Do) / 2

Totals

40

Table 2 - Summary of Community Analysis of Single Quadrat by the Quadrat Sampling Technique (#2)

Location ________________________________________

Quadrat Number ______________

Date _______________________________

Quadrat Size ______________

Members of group ___________________________________________________________________________

Species

Density

Total Basal

Relative Density (D)

Relative Dominance (Do)

Area Covered

Species

Density

Total Basal

Importance
(D + Do) / 2

Relative Density (D)

Relative Dominance (Do)

Importance

41

Area Covered

(D + Do) / 2

Totals

42

Table 2 - Summary of Community Analysis of Single Quadrat by the Quadrat Sampling Technique (#3)

Location ________________________________________

Quadrat Number ______________

Date _______________________________

Quadrat Size ______________

Members of group ___________________________________________________________________________

Species

Density

Total Basal

Relative Density (D)

Relative Dominance (Do)

Area Covered

Species

Density

Total Basal

Importance
(D + Do) / 2

Relative Density (D)

Relative Dominance (Do)

Importance

43

Area Covered

(D + Do) / 2

Totals

44

Table 3 - Summary of Community Analysis of All Quadrats by the Quadrat Sampling Technique

Location ___________________________

Date _____________________________

Number of Quadrats _____

Quadrat Size __________

Species

Frequency

Density

Total Basal

(No. of quadrats

(No. of

Area Covered

of occurrence)

trees)

Relative

Relative

Relative

Importance

Frequency (F)

Density

Dominance

(F+D+Do)

(D)

45

Species

Frequency

Density

Total Basal

(No. of quadrats

(No. of

Area Covered

of occurrence)

trees)

Relative

Relative

Relative

Importance

Frequency (F)

Density

Dominance

(F+D+Do)

(D)

Totals

Average no. trees per quadrat


Total no. trees / Total no. quadrats =

Total no. trees per hectare =

Average Basal area per quadrat


Total basal area / Total no. quadrats =

Total basal area per hectare =

46

Questions

Do species with the highest relative densities also have the highest relative frequency and relative dominance
values?

Do the species with the highest importance values contain saplings? What does this indicate about the future of
this forest?

When comparing quadrats, evaluate whether the dominant species, or those with the highest importance values,
are the same or different.

47

How similar are the quadrats to one another in species composition?

How do relative dominance and relative density values compare for different species?

Are there differences between the seedling or sapling numbers in different quadrats?

48

Plant Sampling Equipment


Know the names and how to use this equipment

Compass

Dbh tape

Tape measure

Note that dbh (diameter at breast height) is sampled 1.4 m above the ground.

49

Lab 5 Quadrat-based Community Sampling a Comparison


of Riparian and Cross Timbers Communities

The term Riparian refers to the area adjacent to a river or stream. These areas are typically dominated
by water-loving plants . Riparian areas are important for a number of reasons, not least of which is that
they act as filters to reduce soil runoff into water bodies. The term Cross Timbers refers to the oak
(predominantly Blackjack Oak Q. marilandica and Post Oak Q. stellata) woodlands that extend from
Texas into extreme southeastern Kansas. This area was historically intermixed with tall-grass prairie.

(Image from the EPA)


Due to fire suppression, Eastern Redcedar is invading much of the Cross Timbers.
In todays lab, you will use the techniques that you learned last week (quadrat-based community
sampling) to compare species diversity between riparian and Cross Timbers areas.

50

Table 1 - Transect Method Field Data (#1)

Location ________________________________________

Number of saplings in Quadrat ______________

Date _______________________________

Number of seedlings in Quadrat ______________

Members of group ___________________________________________________________________________

Number

Species

DBH

Basal Area Coverage


(Circumference)

51

52

Table 1 - Transect Method Field Data (#2)

Location ________________________________________

Number of saplings in Quadrat ______________

Date _______________________________

Number of seedlings in Quadrat ______________

Members of group ___________________________________________________________________________

Number

Species

DBH

Basal Area Coverage


(Circumference)

53

54

Table 1 - Transect Method Field Data (#3)

Location ________________________________________

Number of saplings in Quadrat ______________

Date _______________________________

Number of seedlings in Quadrat ______________

Members of group ___________________________________________________________________________

Number

Species

DBH

Basal Area Coverage


(Circumference)

55

56

Table 2 - Summary of Community Analysis of Single Quadrat by the Quadrat Sampling Technique (#1)

Location ________________________________________

Quadrat Number ______________

Date _______________________________

Quadrat Size ______________

Members of group ___________________________________________________________________________

Species

Density

Total Basal
Area Covered

Relative Density (D)

Relative Dominance (Do)

Importance
(D + Do) / 2

57

Species

Density

Total Basal
Area Covered

Relative Density (D)

Relative Dominance (Do)

Importance
(D + Do) / 2

Totals

58

Table 2 - Summary of Community Analysis of Single Quadrat by the Quadrat Sampling Technique (#2)

Location ________________________________________

Quadrat Number ______________

Date _______________________________

Quadrat Size ______________

Members of group ___________________________________________________________________________

Species

Density

Total Basal
Area Covered

Relative Density (D)

Relative Dominance (Do)

Importance
(D + Do) / 2

59

Species

Density

Total Basal
Area Covered

Relative Density (D)

Relative Dominance (Do)

Importance
(D + Do) / 2

Totals

60

Table 2 - Summary of Community Analysis of Single Quadrat by the Quadrat Sampling Technique (#3)

Location ________________________________________

Quadrat Number ______________

Date _______________________________

Quadrat Size ______________

Members of group ___________________________________________________________________________

Species

Density

Total Basal
Area Covered

Relative Density (D)

Relative Dominance (Do)

Importance
(D + Do) / 2

61

Species

Density

Total Basal
Area Covered

Relative Density (D)

Relative Dominance (Do)

Importance
(D + Do) / 2

Totals

62

Table 3 - Summary of Community Analysis of All Quadrats by the Quadrat Sampling Technique

Location ___________________________

Date _____________________________

Number of Quadrats _____

Quadrat Size __________

Species

Frequency

Density

Total Basal

(No. of quadrats

(No. of

Area Covered

of occurrence)

trees)

Relative

Relative

Relative

Importance

Frequency (F)

Density

Dominance

(F+D+Do)

(D)

63

Totals

Average no. trees per quadrat


Total no. trees / Total no. quadrats =

Total no. trees per hectare =

Average Basal area per quadrat


Total basal area / Total no. quadrats =

Total basal area per hectare =

64

Questions

What species had the highest relative frequency, density and dominance in each habitat (riparian vs. crosstimbers forest)?

Which habitat (riparian or cross-timbers) had the highest species richness?

Which habitat (riparian or cross-timbers) had the highest tree abundance?

65

Create a Shannon-weiner and Simpson diversity index for each habitat. Which habitat (riparian or cross-timbers)
had the highest diversity?

Calculating species-area curves.


One way to determine whether you have adequately sampled an area is to create a species-area curve. On the x-

axis you plot your area, and on the y-axis you plot the total number of species recorded. For example, imagine
you recorded the following data:

Plot 1

Plot 3

Plot 4

Species A Species A

Species B

Species A Species A Species A

Species B Species B

Species D Species B

Species C

Plot 2

Plot 5

Species C

Plot 6

Species C

Species D Species E

Species D

Species E

Species E

Species F

Species F

66

Your graph would then look like this:

20
18
16
14
12
10
8
6
4
2
0
1

A total of six species were detected. Because your figure levels off, you have adequately sampled the area. If the
curve doesnt level off, this indicates that you are still finding new species with each new site you visit and you
have not yet adequately sampled the area.

Create species-area curves for each habitat. Did we adequately sample for species richness? How can you tell?

67

Lab 6 Quarter-based Community Sampling


The quarter method is a widely used method of vegetation sampling in forests. It is a distance method, and uses
points rather than quadrats or plots. The points can be 1) randomly located on a grid of the area to be sampled,
2) located by a stratified random method, such as at random points along evenly spaced transects, or 3) located
systematically along a transect. Because of time constraints, we will use the last method.
Materials
Tape measures
DBH tapes
Clipboards
Data sheets
Methods
We will establish several 50m transects through a forest. Each team will sample along one transect. Points will
be located at 10m intervals (0, 10, 20, 30, 40 and 50m) along each transect. At each point, establish a line
perpendicular to the transect, dividing the area around the point into four "quarters". In each quarter, locate the
tree nearest the point that is >5cm DBH (or 15.7cm circumference). On a Table 1 data sheet, record the species,
distance from the point to the center of the tree in meters, and DBH (or circumference) in centimeters for each
of the four individual trees per point. Continue until all points have been sampled.

68

Calculations
Sample data
Sampling Point
0m

10 m

20 m

30 m

40 m

Quarter Number
1
2
3
4
1
2
3
4
1
2
3
4
1
2
3
4
1
2
3
4

Species
Blackjack Oak
Post Oak
Sugarberry
Redbud
Post Oak
Sugarberry
Blackjack Oak
Sugarberry
Blackjack Oak
Blackjack Oak
Blackjack Oak
Blackjack Oak
Redbud
Sugarberry
Sugarberry
Redbud
Blackjack Oak
Blackjack Oak
Post Oak
Post Oak

Distance (m)
1.1
1.6
2.3
3.0
2.8
3.7
0.9
2.2
2.8
1.1
3.2
1.4
1.3
0.8
0.7
3.1
1.5
2.4
3.3
1.7
40.9

DBH
6
48
15
11
65
16
8, 6
9
4
6
6
5
19
22
12
7
7
5
27
36

Total

Youll be doing the same calculations as for the previous two labs. In addition, for quarter-based analysis you
also need to calculate out
1.) =

where n is the number of points sampled

a. So, in this example =

40.9
45

40.9
20

= 2.05

2.) Density per hectare. This is given by =


a. So, in this example, =

1
10,000
2.05

10,000

= 2380 /

And, in case, you need it, here are the formulas from the previous two labs.

Convert DBH (on Table 1 data sheet) to basal area in cm2 for each tree. Use the equation:
Basal area = Pi x (0.5 DBH)2
Total the basal area covered by all trees of each species to obtain a total basal area per species.
Each team should turn in their data sheets to the instructor upon completion.
Each student should then obtain a complete set of data from the instructor for all transects and perform the
following calculations.
A) For each individual species calculate:
1)

Frequency = Number of points of occurrence

69

2)

Density = Number of individuals

3)

Total basal area

4)

Relative frequency = (No. of points at which a species occurs / Total no. of points of occurrence
for all species) x 100

5)

Relative density = (No. of individuals of a species / Total no. individuals of all species) x 100

6) Relative dominance = (Total basal area of a species / Total basal area of all species) x 100
7)

Importance = (Relative frequency + Relative density + Relative dominance) / 3

Summarize these calculations in Table 2, as shown in the example on the last page of this exercise.

70

Table 1 - Data Sheet for Recording Species, Coverage Area, and Point-to-Point Distances in Point-Quarter Sampling (site 1)

Location ___________________________________

Sampling Point
0m

Quarter Number
1

Species

Date _________________

Distance

DBH

2
3
4
10 m

1
2
3
4

20 m

1
2
3
4

30 m

1
2

71

3
4
40 m

1
2
3
4

50 m

1
2
3
4

72

Table 1 - Data Sheet for Recording Species, Coverage Area, and Point-to-Point Distances in Point-Quarter Sampling (site 2)

Location ___________________________________

Sampling Point
0m

Quarter Number
1

Species

Date _________________

Distance

DBH

2
3
4
10 m

1
2
3
4

20 m

1
2
3
4

30 m

1
2
3

73

4
40 m

1
2
3
4

50 m

1
2
3
4

74

Table 1 - Data Sheet for Recording Species, Coverage Area, and Point-to-Point Distances in Point-Quarter Sampling (site 3)

Location ___________________________________

Sampling Point
0m

Quarter Number
1

Species

Date _________________

Distance

DBH

2
3
4
10 m

1
2
3
4

20 m

1
2
3
4

30 m

1
2
3

75

4
40 m

1
2
3
4

50 m

1
2
3
4

76

Table 2 - Summary of Community Analysis of All Points by the Quarter Sampling Technique

Location ______________________________________

Species

Date ___________________

Frequency

Density

Total Basal

Relative

Relative

Relative

Importance

(No. of points

(No. of

Area Covered

Frequency

Density

Dominance

(F+D+Do)

of occurrence)

Trees)

(F)

(D)

(Do)

77

Totals

Total distance (m) =

Trees per hectare =

Average basal area per tree (cm ) =

Mean distance (m) =

Total basal area (cm ) =

Basal area per hectare (cm ) =

78

Questions

Do species with the highest relative densities also have the highest relative frequency and relative dominance
values? If not, explain.

When comparing quadrats, evaluate whether the dominant species, or those with the highest importance
values, are the same or different between the quadrats.

How similar are the quadrats to one another in species composition?

79

How do relative dominance and relative density values compare for different species?

How do these data compare to those obtained by the quadrat?

80

Week 7 MidTerm Exam


Study guide for Ecological Methods Mid-term Exam
Plant walk lab

Be able to visually identify the 19 species that were covered in the lab
Know the difference between simple and compound leaves
Be familiar with deciduous vs. evergreen

Aquatic ecosystems

Be able to describe the difference between a lentic and lotic system


Be able to describe the difference between riffles and pools
Be able to identify the organisms captured in class
Be able to identify and use a dissolved oxygen meter, pH meter, conductivity meter
Be able to identify and describe how to use a ponar trap, dipnet, kick net, and plankton net
Be able to calculate the Shannon-Weiner index and the Simpson index
Be able to read the output for t-test results in Excel

Vegetation lab

Be able to identify all 34 plant species


Be able to use dbh tape, compass, tape measure
Be able to read rarefaction curves
Compare riparian and cross timbers systems

81

Week 9 Fall Break

82

Week 8 Terrestrial Invertebrates


It is difficult to accurately sample terrestrial invertebrate populations. These beasts occupy many different
microhabitats and this requires several collecting techniques to be employed to make obtain suitable
samples. All collecting methods are limited and some species will be missed. Therefore, several methods of
collecting should be used. By comparing samples of the same kind, useful information, such as population
changes and community patterns may be observed. We will sample two areas (prairie and forest) and
compare the samples collected from each area. The collecting techniques we will use are rather general and
may by modified for specific sampling requirements.
Ground Macroinvertebrates
1.
2.
3.
4.
5.
6.

Place the 0.1m 2 wire loop on the ground.


Carefully remove all contents within the loop, including both litter and detritus, down to a depth of
5cm.
Place this sample into a jar, record its number, add preservative, and return it to the lab for
sorting, identifying, and counting organisms.
Repeat this procedure.
Note: An alternative to field preservation is the use of Berlese-Tulgren funnels. Upon returning to
the lab, place the sample into the Berlese-Tulgren funnel. After one week, some of the animals will
have crawled away from the heat and drying soil, and fallen into the 80% ethanol in the jar below.
The animals in the sample may be sorted, identified, and counted.

Cryptozoans
1.
2.
3.
4.
5.

Drop boards (0.1m2) may be placed to collect animals that prefer cool, moist microhabitats.
Each drop board is left undisturbed for at least one week.
When sampling, lift each board carefully, but quickly.
Collect animals observed on the surface of the soil and leaf debris.
Place this sample into a jar, record its number, add preservative, and return it to the lab for
sorting, identifying, and counting organisms.

Foliage Invertebrates
1.
2.
3.

Using an insect net, take 10 steps through the vegetation (do not sweep the same area more than
once).
Place this sample into a jar, record its number, add preservative, and return it to the lab for
sorting, identifying, and counting organisms.
Repeat this procedure.

83

Ground Macroinvertebrates (Prairie)

Taxa

1A

3A

5A

7A

9A

No. of individuals
No. of species
Species diversity

84

Ground Macroinvertebrates (Forest)

Taxa

2A

4A

6A

8A

10A

No. of individuals
No. of species
Species diversity

85

Cryptozoans (Prairie)

Taxa

1B

3B

5B

7B

9B

No. of individuals
No. of species
Species diversity

86

Cryptozoans (Forest)

Taxa

2B

4B

6B

8B

10B

No. of individuals
No. of species
Species diversity

87

Foliage Invertebrates (Prairie)

Taxa

1C

3C

5C

7C

9C

No. of individuals
No. of species
Species diversity

88

Foliage Invertebrates (Forest)

Taxa

2C

4C

6C

8C

10C

No. of individuals
No. of species
Species diversity

89

Questions

1.

Which area (prairie or forest) contained the most number of individuals?

2.

Which area (prairie or forest) contained the most number of species?

3.

Which area (prairie or forest) had the highest species diversity value?

4.

Which microhabitat contained the most number of individuals?

5.

Which microhabitat contained the most number of species?

6.

Which microhabitat contained the highest species diversity value?

7.

Explain the above results.

90

Terrestrial invertebrates list

Phylum Athropoda
Class Insecta
Order Odonata (dragonflies, damselflies)
Order Blattodea (cockroaches)
Order Mantodea (praying mantis)
Order Orthoptera (cricket, grasshopper, katydid)
Order Hemiptera (true bugs)
Order Coleoptera (beetles)
Order Diptera (flies)
Order Lepidoptera (butterflies, moths)
Order Hymenoptera (ants, bees, wasps)
Class Arachnida
Order Araneae (spiders)

91

Terrestrial Invertebrate Sampling Equipment


Know the names and how to use this equipment

Berlese-Tulgren funnel

Insect net

Wire loop

Drop board

92

Week 10 Mammals
There are several different techniques for studying mammals. The three techniques you will be exposed
to in todays lab include Sherman live mammal traps, plaster tracks and radiotelemetry.
Sherman live mammal traps are essentially box traps that are used to catch mammals. They come in a
variety of sizes, but are most frequently used to catch small mammals (typically rodents). The traps are
usually baited with oats or peanut butter and then left overnight. They are checked in the morning and
the mammals that have been caught are either released or sacrificed.
Many mammals are too large or infrequent to be easily captured with a Sherman live mammal trap. In
this case, an easy way to determine their presence is to look for tracks. Plaster tracks are made using
plaster of paris and can be used as references. To make a plaster track, mix two parts of water to one
part plaster of paris. Put a plastic circle around the track and pour the resulting mixture into the circle.
As the plaster is hardening, push a stick (or small pole) through the plaster, leaving a small hole. (This
hole is used to attach a tag to the plaster track.) The plaster should harden within approximately hour.
Another common method for studying mammals is using radiotelemetry. A radio tag is attached to the
mammal. The researcher will have a Yagi antenna connected to a receiver. The intensity (i.e.
loudness) of the beep that emerges from the receiver will depend upon both distance and direction
to the radio tag. If you are pointed at the radio tag and close the beep will be relatively loud. If you are
further away, the beep will be relatively soft. Researchers adjust both the gain and volume on the
receiver to help identify the direction to the tag.

Homework
1.) Get plaster of Paris casts from three different mammal species (domestic mammals excluded!)
(30 pts)
2.) Describe five methods to determine what mammal species are present in an area (10 pts)

93

Mammal Study Guide


For Ecological Methods, you will need to know how to operate the traps and the common names of the
following species.

Sherman Live Mammal trap

Hispid Cotton Rat


Sigmodon hispidus

North American Deermouse


Peromyscus maniculatus

Plains Harvest Mouse


Reithrodontomys montanus

Nine-banded Amradillo
Dasypus novemcinctus

Virginia Oppossum
Didelphis virginiana

North American Porcupine


Erethizon dorsatum

White-tailed Deer
Odocoileus virginianus

Raccoon
Procyon lotor

Gray Fox
Urocyon cinereoargenteus

American Beaver
Castor canadensis

House Mouse
Baeolophus bicolor

94

Coyote
Canis latrans

Cougar
Puma concolor

White-tailed Deer
Odocoileus virginianus

Virginia Oppossum
Didelphis virginiana

Raccoon
Procyon lotor

Raccoon
Procyon lotor

Nine-banded Amradillo
Dasypus novemcinctus

Cougar
Puma concolor

95

Week 11 Birds
Although most birds are diurnal, they tend to avoid coming in close to people. Consequently, studying
birds in the field typically involves the use of binoculars. Binoculars vary in both their magnification and
their field of view. Most binoculars will have numbers separated by an x on them, such as 10 x 50 or 7
x 42. The first number is the magnification. If the bird is 35 m away, a 10x binocular will make it appear
that it is only 3.5 m away, while a 7x binocular will make it appear as though it was 5 m away. The field
of view is how much of an area you can see when looking through the binoculars and it is inversely
related to magnification. The higher the magnification, the smaller the field of view. Consequently,
people who watch small, active birds in forests (such as warblers) tend to use 7x and 8x binoculars. In
contrast, people who watch big, distant birds on the beach tend to use 10x binoculars and/or telescopes
(called spotting scopes in birding circles).
The second number on the binoculars is the diameter of the objective lens (the big lenses). This affects
brightness. The best measure of the binoculars brightness is the exit pupil diameter. This is found by
dividing the diameter of the objective lens by the magnification. So for a 10 x 40 binocular, the exit pupil
size is 4 mm. The pupil of the human eye changes in response to light conditions. In bright light, it
contracts down to ~2 mm, while in very dim conditions it expands to ~7 mm. Consequently, while a 10 x
40 binocular would be perfectly fine for use on a bright, sunny day, images would appear to be very dark
indeed as the sun sets. Astronomers and other people who watch the night sky tend to use 7 x 50
binoculars.
In some cases, it is possible to attract birds by pishing, squeaking or using a soundbox. Pishing involves
going pish pish pish can be fairly effective at attracting small Passerines. It is speculated that it
mimics the alarm call of the Tufted Titmouse (Baeolophus bicolor) and other birds will fly in to see what
the fuss is about. Squeaking involves making a high-pitched squeak by kissing the back of your hand.
This may also mimic an alarm call and tends to be fairly effective at getting sparrows (and other birds) to
appear for a few seconds. A soundbox is created by cupping your two hands together and blowing
across the knuckles on your thumb, causing the air inside to resonate. This is a good technique for
imitating owls and doves.
Birds can also be studied in the hand. Much of our knowledge of avian migration comes from bird
banding, where licensed bird banders attach a metal ring to the leg of a bird. This band has a unique
numeric code as well as contact information for the national Bird Banding Laboratory (BBL). Banders
submit their records annually to the BBL. The BBL will then inform banders when their birds are reencountered. Because all native birds are protected by law (the Migratory Bird Treaty Act of 1918), it is
necessary to obtain permits to band birds. A common method of catching birds is using mist nets (which

96

superficially resemble volleyball nets). The birds become entangled in the nets and are then carefully
removed by the bird bander. Because birds are small and fragile (due in part to their pneumatized
bones) it requires great delicacy to safely extract a bird captured in a mist net.

97

Bird Study Guide


For Ecological Methods, you will need to know the common names of the following ten bird species.

Blue Jay
Cyanocitta cristata

Mourning Dove
Zenaida macroura

Carolina Chickadee
Poecile carolinensis

Eurasian Collared-Dove
Streptopelia decaocto

Tufted Titmouse
Baeolophus bicolor

Northern Cardinal
Cardinalis cardinalis

98

European Starling
Sturnus vulgaris

Red-tailed Hawk
Buteo jamaicensis

Eastern Screech-Owl
Megascops asio

House Sparrow
Passer domesticus

99

In lab today, you learned how to pish and squeak. For your homework, visit four locations
(separated by at least 100 m). At each location, record the number of birds that you see during two
minutes. Then pish for one minutes and record the number of birds that you see during the next two
minutes. Finally, squeak for one minute and record the number of birds that you see.
Is there a significant difference in the number of birds you see? Use a one-way ANOVA to test this and
create a figure to illustrate the average number of birds detected using each method.

Site

Control

Pish

Squeak

100

Week 12 Ecobeaker Isle Royale


The Wolves and Moose of Isle Royale
If you were to travel on Route 61 to the farthest
reaches of Minnesota and stand on the shore of
Lake Superior looking east, on a clear day you
would see Isle Royale. This remote, forested
island sits isolated and uninhabited 15 miles off
of the northern shore of Lake Superior, just south
of the border between Canada and the USA. If
you had been standing in a similar spot by the
lake in the early 1900s, you may have witnessed
a small group of hardy, pioneering moose
swimming from the mainland across open water,
eventually landing on the island. These fortunate
moose arrived to find a veritable paradise,
devoid of predators and full of grass, shrubs, and trees to eat. Over the next 30 years, the moose
population exploded, reaching several thousand individuals at its peak. The moose paradise didnt last
for long, however.
Lake Superior rarely freezes. In the 1940s, however, conditions were cold and calm enough for an ice
bridge to form between the mainland and Isle Royale. A small pack of wolves found the bridge and
made the long trek across it to the island. Once on Isle Royale, the hungry wolves found their own
paradise a huge population of moose. The moose had eaten most of the available plant food, and
many of them were severely undernourished. These slow-moving, starving moose were easy prey for
wolves.
The Isle Royale Natural Experiment
The study of moose and wolves on Isle Royale began in 1958 and is thought to be the longest-running
study of its kind. The isolation of the island provides conditions for a unique natural experiment to study
the predator-prey system. Isle Royale is large enough to support a wolf population, but small enough to
allow scientists to keep track of all of the wolves and most of the moose on the island in any given year.
Apart from occasionally eating beaver in the summer months, the wolves subsist entirely on a diet of
moose. This relative lack of complicating factors on Isle Royale compared to the mainland has made the
island a very useful study system for ecologists.
The EcoBeaker Version of Isle Royale
During this lab, you will perform your own experiments to study population dynamics using a computer
simulation based on a simplified version of the Isle Royale community. The underlying model includes
five species: three plants (grasses, maple trees, and balsam fir trees), moose, and wolves. If you were

2010, SimBiotic Software for Teaching and Research, Inc. All Rights Reserved 101

actually watching a large patch of moose-free grass through time, you would observe it slowly
transforming into forest. Likewise, the simulated plant community exhibits a simple succession from
grasses to trees.
While the animal species in the Isle Royale simulation are also simplified compared with their real-world
counterparts, their most relevant behaviors are included in the model. Moose prefer to eat grass and fir
trees. Wolves eat moose, more easily catching the slower, weaker moose. Each individual animal of both
species has a store of fat reserves that decreases as the individual moves around and reproduces, and
increases when food is consumed. Both moose and wolves reproduce; however, for simplicity, the
simulation ignores gender. Any individual with enough energy simply duplicates itself, passing on a
fraction of its energy to its offspring. Death occurs when an individuals energy level drops too low.
Because weaker moose move at slower speeds, they take longer to find food and move away from
predators, so their chance of survival is lower than for healthier moose. In the EcoBeaker simulation,
wolves hunt alone, whereas in the real world, wolves are social animals that hunt in packs. These
simplifications make the simulation tractable, while still retaining the basic qualitative nature of how
these species interact.
Some Important Terms and Concepts
Population Ecology
Population ecology is the study of changes in the size and composition of populations and the factors
that cause those changes.
Population Growth
Many different factors influence how a population grows. Mathematical models of population growth
provide helpful frameworks for understanding the complexity involved, and also (if the models are
accurate) for predicting how populations will change through time. The simplest model of population
growth considers a situation in which limitations to the populations growth do not exist (that is, all
necessary resources for survival and reproduction are present in continual excess). Under these
conditions, the larger a population becomes, the faster it will grow. If each successive generation has
more offspring, the more individuals there will be to have even more offspring, and so on. This type of
population growth is described with the exponential growth model.
The exponential growth model assumes that a population is increasing at its maximum per capita rate of
growth (represented by rmax) also known as the intrinsic rate of increase. If population size is N and
time is t, then:

The notation dN/dt represents the instantaneous change in population size with respect to time. In
this context, instantaneous change simply means how fast the population is growing or shrinking at
any particular instant in time. The equation indicates that at larger values of N (the population size), the
rate at which the population size increases will be greater.
The following graph depicts an example of exponential population growth. Notice how the curve starts
out gradually moving upwards and then becomes steeper over time. This graph illustrates that when the
population size is small, it can only increase in size slowly, but as it grows, it can increase more quickly.

2010, SimBiotic Software for Teaching and Research, Inc. All Rights Reserved 102

Exponential Population Growth

Carrying Capacity
In the real world, conditions are generally not so favorable as those assumed for the exponential growth
model. Population growth is normally limited by the availability of important resources such as food,
nutrients, or space. A populations carrying capacity (symbolized by K) is the maximum number of
individuals of that species that the local environment can support at any particular time. When a
population is small, such as during the early stages of colonization, it may grow exponentially (or nearly
so) as described above. As resources start to run out, however, population growth typically slows down
and eventually the population size levels off at the populations carrying capacity.
To incorporate the influence of carrying capacity in projections of population growth rate, ecologists use
the logistic growth model. In this model, the per capita growth rate (r) decreases as the population
density increases. When the population is at its carrying capacity (i.e., when N = K ) the population will
no longer grow. Again, using the dN/dt notation, if the maximum per capita rate of growth is rmax,
population size is N, time is t, and carrying capacity is K, then:

When the population size (N) is near the carrying capacity (K), K-N will be small and hence, (K-N)/K will
also be small. The change in the population size through time (dN/dt) will therefore decrease and
approach zero (meaning the population size stops changing) as N gets closer to K.
The following graph depicts an example of logistic growth. Notice how it initially looks like the
exponential growth graph but then levels off as N (population size) approaches K (carrying capacity).

2010, SimBiotic Software for Teaching and Research, Inc. All Rights Reserved 103

Logistic Population Growth

While the logistic model is more realistic than the exponential growth model for most populations,
many other factors can also influence how populations change in size through time. For example, the
growth curve for a recently-introduced species might temporarily overshoot the populations carrying
capacity. This would happen if the abundance of resources encountered by the colonizing individuals
stimulated a high rate of reproduction, but the pressures of limited resources were soon felt (i.e.,
individuals might not start dying off until after a period of rapid reproduction has already taken place).
Graphs based on real population data are never such smooth, neat curves as the ones above. Random
events almost always cause population sizes and carrying capacities to fluctuate through time.
Interactions with other species, such as predators, prey, or competitors, also cause the size of
populations to change erratically. To estimate carrying capacity in situations such as these, one generally
calculates the median value around which the population size is fluctuating.
More Information
Links to additional terms and topics relevant to this laboratory can be found in the SimBio Virtual Labs
Library which is accessible via the programs interface.
Starting Up
[ 1 ] Read the introductory sections of the workbook, which will help you understand whats going on in
the simulation and answer questions.
[ 2 ] Start SimBio Virtual Labs by double-clicking the program icon on your computer or by selecting it
from the Start Menu.
[ 3 ] When the program opens, select the Isle Royale lab from the EcoBeaker suite.
IMPORTANT!
Before you continue, make sure you are using the SimBio Virtual Labs version of Isle Royale. The splash
screen for SimBio Virtual Labs looks like this:

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If the splash screen you see does not look like this, please close the application (EcoBeaker 2.5)
and launch SimBio Virtual Labs.
When the Isle Royale lab opens, you will see several panels:
The ISLAND VIEW panel (upper left section) shows a birds eye view of northeastern
Isle Royale, which hosts ideal moose habitat.
The DATA & GRAPHS panel to the right displays a graph of population sizes of
moose and wolves through time.
The SPECIES LEGEND panel above the graph indicates the species in the simulation; the buttons link
to the SimBio Virtual Labs Library where you can find more information about each.
[ 4 ] Click Moose in the SPECIES LEGEND panel to read about moose natural history, and then answer
the following question (you can read about other species too, if you wish)
[ 4.1 ] Based on what you find in the Library, answer the following: could a moose swim fast
enough to win a swimming medal in the Olympics (where the fastest speeds are
around 5 miles / hour)?
Yes No (Circle one)
[ 5 ] Examine the bottom row of buttons on your screen. You will use the CONTROL PANEL buttons to
control the simulation and the TOOLS buttons (to the right) to conduct your experiments. These will be
explained as you need them; if you become confused, position your mouse over an active button and a
tool tip will appear.

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Exercise 1: The Moose Arrive


In this first exercise, you will study the moose on Isle Royale
before the arrival of wolves. The lab simulates the arrival of
the group of moose that swam to the island and rapidly
reproduced to form a large population.

[ 1 ] Click the GO button in the CONTROL PANEL at the bottom of the screen to begin the simulation.
You will see the plants on Isle Royale starting to spread, slowly filling up most of this area of the
island.
Grass starts out as the most abundant plant species, but is soon replaced with maple and
balsam fir trees. The Isle Royale simulation incorporates simplified vegetation succession to
mimic the more complex succession of plant species that occurs in the real world. After about 5
simulated years, the first moose swim over to the island from the mainland and start munching
voraciously on the plants.
[ 2 ] You can zoom in or out using the ZOOM LEVEL SELECTOR at the top of the ISLAND VIEW panel. Click
different Zoom Level circles to view the action up closer or further away. After watching for a
bit, click on the left circle to zoom back out. You can zoom in and out at any time.
[ 3 ] Reset the simulation by clicking the RESET button in the CONTROL PANEL. Confirm that the
simulation has been reset by checking that the TIME ELAPSED box to the right of the CONTROL
PANEL reads 0 Years.
[ 4 ] Click the STEP 50 button on the CONTROL PANEL, and the simulation will run for 50 years and
automatically stop. Watch the graph to confirm that the size of the moose population changes
dramatically when the moose first arrive, and then eventually stabilizes (levels out).
You can adjust how fast the simulation runs with the SPEED slider to the right of the
CONTROL PANEL.
[ 5 ] Once 50 years have passed (model years not real years!), examine the moose population graph
and answer the questions below. (NOTE: if you cant see the whole graph, use the scroll bar at
the bottom of the graph panel to change the field of view.)
[ 5.1 ] What is the approximate size of the stable moose population? ________
[ 5.2 ] What was the (approximate) maximum size the moose population attained? ________
[ 5.3 ] Using the horizontal and vertical axes below, roughly sketch the population size graph showing
the simulated moose population changing over time. Label one axis POPULATION SIZE (N) and the

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other one TIME (years). You do not need to worry about exact numerical values; just try to capture
the shape of the line.

[ 5.4 ] Examine your graph and determine the part that corresponds to the moose population
growing exponentially. Draw a circle around that part of the moose population curve
you drew above.
[ 5.5 ] The moose population grew fastest when it was:
Smallest Medium-sized Largest (Circle one)
[ 5.6 ] What is the approximate carrying capacity of moose? Draw an arrow on your graph
that indicates where the carrying capacity is (label it K) and then write your
answer in the space below:

[ 6 ] The following logistic growth equation should look familiar (if not, revisit the Introduction):

[ 6.1 ] What does dN/dt mean, in words?

[ 6.2 ] Think about what happens to dN/dt in the equation above when the population size
(N) approaches the carrying capacity (K)? Think about the case when the two
numbers are the same (N = K). Rewrite the right-hand side of the equation above,
substituting K for N. Write this new version of the equation below:
dN/dt = _________________ when N=K

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[ 6.3 ] Look at the equation you just wrote and figure out what happens to the right-hand
side of the equation. Then complete the following sentence by circling the correct
choices.
According to the logistic growth equation, when a growing population reaches its
carrying capacity (N = K),
dN/dt = 0 / 1 / K / N / rmax (Circle one),
and the population will
grow more rapidly / stop growing / shrink (Circle one)
[ 7 ] Look at the graph that depicts an example of logistic growth and compare that to your moose
population growth graph.
[ 7.1 ] Sketch both curves in the spaces provided below. (Dont worry about the exact numbers; just
show the shapes of the curves. Be sure to label the axes!)

[ 7.2 ] How do the shapes of the curves differ? Describe the differences in terms of population sizes
and carrying capacities.

[7.3 ] Provide a biological explanation for why the moose population overshoots its carrying capacity
when moose first colonize Isle Royale. (HINT: consulting the Introduction might help.)

[ 7.4 ] At year 50 or later, with the moose population at its carrying capacity, what would happen if an
extra 200 moose suddenly arrived on Isle Royale? How would this change the population graph over
the next 20 to 30 years? In the space provided, draw a rough sketch of what you think the graph
would look like under these conditions. Be sure to label the axes.
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[ 8 ] Now you will test your prediction by increasing the number of moose on the island. Click the ADD
MOOSE button in the TOOLS panel. With the ADD MOOSE button selected, move your mouse
to the ISLAND VIEW, click and hold down the mouse to draw a small rectangle. As you draw, a
number at the top of the rectangle tells you how many moose will be added. When you release
the mouse, the new moose appear inside your rectangle. Add approximately 200-300 moose.
HINT: To obtain the exact moose population size from the graph, click the graph to see
the x and y data values at any point (population size is the y value).
[ 9 ] Click GO to continue running the simulation for 20 to 30 more years and watch what happens to the
moose population. Click STOP to pause the simulation. Then answer the following questions:
[ 9.1 ] Did you predict correctly in question 8.3? ________
[ 9.2 ] What is the carrying capacity of moose on Isle Royale after adding 200-300 new moose?
________

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Exercise 2: The Wolves Arrive


One especially cold and harsh winter in the late 1940s, Lake
Superior froze between the mainland and Isle Royale. A small
pack of wolves travelled across the ice from Canada and reached
the island. In this second exercise, you will investigate how the
presence of predators affects the moose population through
time.
[ 1 ] To load the next exercise, select The Wolves Arrive from the SELECT AN EXERCISE menu at the
top of the screen.
[ 2 ] Click STEP 50 to advance the simulation 50 years. You will see moose arrive and run around the
island eating plants as before. Next, you will add some wolves to the island, but first answer the
following question:
[ 2.1 ] How do you predict the moose population graph will change with predatory wolves in
the system? Will the moose population grow or shrink?

[ 3 ] Activate the ADD WOLF button in the TOOLS panel by clicking it. Add 20-40 wolves to Isle Royale by
drawing small rectangles on the island (they will fill with wolves) until you have succeeded in
helping the wolf population to get established.
[ 4 ] Run the simulation for about 200 years (you can click STEP 50 four or five times). Observe how the
moose and wolves interact, and how the population graph changes through time. (To better
observe the system you can try changing the simulation speed or zoom level.)
[ 4.1 ] In the space below, copy the moose-wolf population graph starting with the time when wolves
were established. Make sure you label the axes.

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NOTE: if you have trouble estimating the wolf population size from the graph, hold
down your mouse button and move the pointer along the graph line to see the x and y
values represented.
[ 4.2 ] Did the introduction of wolves cause the moose population size to decrease or
increase? If so, how much smaller or larger (on average) is the moose population
when wolves are present?

[ 4.3 ] You should have noticed that the populations of moose and wolves go through cycles.
(If not, run the simulation for another 100 years.) Describe the pattern and provide a
biological explanation for what you observe. Does the moose or the wolf population
climb first in each cycle? Which population drops first in each cycle?

[ 5 ] If you havent already, click STOP.


[ 6 ] The MICROSCOPE tool lets you sample animals to determine their current energy reserves. Activate
the MICROSCOPE tool by clicking it. Then click several moose to confirm that you can measure
their Fat Stores. These reserves are important health indicators for moose; the greater a
mooses fat stores, the more likely it will survive the winter and produce healthy, viable
offspring.
[ 6.1 ] All else being equal, which do you think would be healthier (on average), moose on an
island with wolves or moose on an island without wolves? Explain your reasoning.

[ 7 ] You will now test your prediction. RESET the simulation and then click GO to run the simulation
without wolves until the moose population has stabilized at its carrying capacity. Click STOP so you can
collect and record data. Decrease your zoom level to see as much of the island as possible.
[ 8 ] Randomly select 10 adult moose and use the MICROSCOPE tool to sample their fat stores. Record
your data on the left-hand side of the table below. Do NOT sample baby moose; they are still
growing and so do not store fat as adults do.

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[ 9 ] When you are done, activate the ADD WOLVES button as before, and add 10-20 wolves. Click GO
and run the simulation until the moose and wolf populations have cycled several times. STOP
the simulation when the moose population is about midway between a low and high point (i.e.
at its approximate average size).
[ 10 ] Randomly select another 10 adult moose and use the MICROSCOPE tool to sample their fat stores.
[ 10.1 ] Record the values on the right-hand side of the table.

[ 10.2 ] Calculate and record the mean fat stores of adult moose with wolves absent and present in
the table above. (You can open your computers calculator by clicking the CALCULATOR button near
the lower right corner of your screen.) Provide a biological explanation for any differences you have
observed.

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Exercise 3: Changes in the


Weather
You have probably heard that scientists are concerned
about climate change and the effects of global
warming due to increasing atmospheric greenhouse
gases. Recent evidence suggests that temperatures
around the world are rising. In particular, the average
yearly temperature in northern temperate regions is
expected to increase significantly. This change will lead to longer, warmer spring and summer seasons in
places like Isle Royale. The duration of the growing season for plants will therefore be extended,
resulting in more plant food for moose living on the island.
How would a longer growing season affect the moose and wolf populations on Isle Royale? Would they
be relatively unaffected? Would the number of moose and wolves both increase indefinitely with higher
and higher temperatures, and longer and longer growing seasons?
One way ecologists make predictions about the impacts of global warming is by testing different
scenarios using computer models similar to the one youve been using in this lab. Even though
simulation models are simplifications of the real world, they can be very useful for investigating how
things might change in the future. In this exercise, you will use the Isle Royale simulation to investigate
how changes in average yearly temperature due to global warming may affect the plant-moose-wolf
system on the island.
[ 1 ] Use the SELECT AN EXERCISE menu to launch Changes in the Weather.
[ 2 ] Click STEP 50 to advance the simulation 50 years. You can zoom in to view the action up close. The
moose population should level out before the simulation stops.
[ 3 ] Activate the ADD WOLF button in the TOOLS panel. Add about 100 wolves by holding down your
mouse button and drawing rectangular patches of wolves. Remember to look at the number at
the top of the rectangle to determine how many wolves are added.
[ 4 ] Advance the simulation 150 more years by clicking STEP 50 three times. Watch the action. The
simulation should stop at Year 200.
[ 4.1 ] Estimate the average and maximum sizes for moose and wolf populations after the
wolves have become established. Record these values below:
Maximum moose population size: _________________
Maximum wolf population size: _________________
Average moose population size: _________________

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Average wolf population size: _________________


[ 5 ] In the PARAMETERS panel below the ISLAND VIEW you will see Duration of Growing Season
options where you can select different scenarios. The default is Normal, which serves as your
baseline this is the option you have been using thus far.
The Short option simulates a decrease in the average annual temperature on Isle
Royale. The growing season is shorter than the baseline scenario, which results in
annual plant productivity that is about half that of Normal.
The Long option simulates a warming scenario in which the growing season begins
earlier in the spring and extends later in the autumn. Plant productivity is almost
double that of Normal.
[ 5.1 ] Predict how moose and wolf population trends will differ with the Short growing
season compared to the Normal scenario. Will average population sizes be smaller
or larger? Why?
[ 6 ] Without resetting the model, select the Short growing season option.
[ 7 ] Advance the simulation another 100 years by clicking STEP 50 twice (total time elapsed should be
~300 years).
[ 7.1 ] Estimate the maximum and average sizes for moose and wolf populations after several
cycles with a Short growing season. Record these values below:
Maximum moose population size: _________________
Maximum wolf population size: _________________
Average moose population size: _________________
Average wolf population size: _________________
[ 7.2 ] How do these numbers compare to those you observed with the Normal growing season (Step 4
above)?

[ 8 ] In the Short growing season, the plant growth is half of what it was before.
[ 8.1 ] Based on your measurements, how much do you think the moose carrying capacity
changed, and why?

[ 9 ] Now its time to consider the warming scenario.

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[ 9.1 ] How do you predict that moose and wolf population trends will differ with a Long
growing season, and why?

[ 10 ] Without resetting the model, select the Long growing season option from the PARAMETERS
panel.
[ 11 ] Click GO and monitor the graph as the populations cycle. If you watch for a while you should
notice something dramatically different about this scenario, in which the plant productivity is
high.
[ 12 ] Click STOP and estimate the maximum and average size for moose and wolf populations under the
Long growing season scenario
[ 12.1 ] Record these values below:
Maximum moose population size: _________________
Maximum wolf population size: _________________
Average moose population size: _________________
Average wolf population size: _________________
[ 12.2 ] If you watched for a while, you probably saw some species go extinct. If you didnt observe
extinctions, you can continue to run the simulation until you see this dramatic phenomenon. Explain
why you think extinction is more likely in this scenario than the other two (this is known as the
paradox of enrichment).

[ 12.3 ] Looking at your results from running the simulation under the normal climate conditions and
the two alternative scenarios, were your predictions correct? Provide biological explanations for the
trends and differences that you observed. Pay particular attention to how the population cycles
changed (e.g., increased, decreased, became less stable) as the rate of plant growth changed.

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[ 12.4 ] [Optional] If you have already talked about global warming and climate change in class,
provide another example of how increased yearly temperature can affect an animal or plant
population. In particular, think about pests, invasive species, disease, or species of agricultural
importance.

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Extension Exercise: Whats the Difference?


In Exercise 2 you conducted an experiment comparing health of moose with wolves absent to health of
moose with wolves present. You probably observed at least a small difference between the samples, but
does that really indicate that moose have greater fat stores when wolves are present? The difference
could be related to wolves, but it could also have arisen simply by chance. You might have accidentally
selected very healthy moose one time and unhealthy moose the other. How can you know whether the
difference in means between two samples is real?
The short answer is that you cant. But you can make a good guess using statistics. In fact, inferential
statistics were invented to allow us to better uncover the truth and answer these sorts of questions. In
this section, you will perform a simple statistical test, called a t-test, to decide whether or not the
wolves presence had a significant effect on moose fat stores. If we were to be very thorough and formal
in our t-test lesson, we would include a lengthy discussion of such concepts as random variables,
sampling distributions, standard errors, and alpha levels. These are important, but to keep this short, we
will just focus on the core ideas underlying the t-test.
You start with a question: Is the mean moose fat stores different when wolves are present versus
absent? The null hypothesis is a negative answer: there is no real difference. Under the null hypothesis,
the difference in your samples arises from chance. The alternative hypothesis is that there is an effect
of wolves on moose fat stores. In order to know which hypothesis your samples support, we examine
the difference in means relative to the variability you observed.
[ 1 ] Look back at Exercise 2 where you measured the fat stores of adult moose with wolves absent and
present, and record those values here. Note that the subscript p represents samples with
wolves present, while a represents those with wolves absent.
[ 1.1 ] Mean fat stores of adult moose, wolves present ( ) : _________
[ 1.2 ] Mean fat stores of adult moose, wolves absent ( ) : __________
[ 1.3 ] Calculate the difference in mean fat stores ( ) : ___________
[ 2 ] Look at the following three hypothetical graphs. Each graph shows two distributions of moose fat
stores, one with wolves present (lighter gray line) and one with wolves absent (darker line). Note that in
each graph, mean moose fat stores are represented by dashed vertical lines, and the difference in
means is the same for all three. However, the variation in fat stores is smaller in the distributions on
the left, and larger in those on the right.

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[ 2.1 ] Which of the above graphs (A, B, or C) would make the most convincing argument that
the difference in fat stores is real, and not just due to chance?
[ 2.2 ] Explain your choice:
If there is a lot of variability in the data sets you are comparing, you will more likely see a difference in
their means just by chance, supporting the null hypothesis. Only if the difference in means is large
compared to the amount of variability in the data do you suppose that the difference might be real. A
statistic called t formalizes this intuition in fact, t is calculated as a ratio of difference in means to
amount of variability. Here is its formula (with the p and a subscripts referring to moose energy with
wolves present vs. absent):

In the formula above, the mean values of the two samples is given by and . The variability of values
within the sampled data sets is incorporated into the denominator, where SE stands for the standard
error of the sample-mean difference (a fancy-sounding phrase for a simple concept: variability).
Calculating this value is straightforward but requires a few steps if you are doing it by hand; the
formula is:

Here, varp and vara are the variances for each sample, a measure of the amount of variability in the
values. Finally, np and na are the number of samples in each data set. If you have never calculated
variance before, dont fret this exercise will walk you through the calculation. Combining the two
above equations yields the following formula for t:

[ 3 ] Examine the formula for t.


[ 3.1 ] Draw a square around the part of the formula for t that compares the means of the
two data sets.
[ 3.2 ] Draw a circle around the part of the formula for t that describes the amount of
variability in the data.
[ 3.3 ] If the means are close together, and the variability is high (so that the difference in means could
more easily have arisen by chance), will the value of t be low or high?

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[ 4 ] You probably noticed a difference in the health of moose when wolves were present versus when
they were absent. To find out whether this difference is large enough to distinguish it from the
null hypothesis, you have to calculate the t statistic for your moose fat stores data. Start by
estimating the variance in each population (with and without wolves) as follows.
[ 4.1 ] Go back to Section 2 and look at your table of adult moose fat stores. Copy the values
from that table into the table below, in the column labeled Fat Store. (Do this for
both samples with and without wolves.)
[ 4.2 ] Focus first on your samples WITHOUT WOLVES. For each fat store value in that sample,
a from step [1.2] above), and enter
subtract the mean fat store with wolves absent (
a. Remember you can click
this difference from the mean in the column labeled x -
the CALCULATOR button near the lower right corner to open your computers
calculator.
[ 4.3 ] Square each difference from the mean and enter the squared value in the column
a)2.
labeled (x -

[ 4.4 ] Sum the squared differences. Enter the sum of squares at the bottom of the table.

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[ 4.5 ] Divide the sum of squares by the sample size minus 1 (na-1). Here, na is the number of moose
whose fat stores you sampled. (Note that sample size is different than population size.) You will use
the estimated variance in the t-test:
vara = (sum squared differences)a /(na-1) = __________
[ 4.6 ] Repeat the above steps ([4.2] through [4.5]) to calculate the variance for moose fat stores WITH
WOLVES present. Remember this time to use the mean fat store with wolves present ( from step [1.1]
above).
varp = (sum squared differences)p /(np-1) = __________
[ 4.7 ] Now that you have calculated variances, plug these values into the equation for the standard
error of the sample-mean difference to calculate an overall measure of variability in your samples.
(And yes, you will divide by the sample sizes again!)

[ 4.8 ] What is the value t of the t-test, given the difference in means and the standard error of the
sample-mean difference you calculated above?

The higher the value of t, the more confident you can be that the difference did not result from chance.
But how confident are you? A common protocol is to call something significant if the probability is less
than 0.05 that the difference is due to chance alone. This probability is dubbed the p-value.
[ 5 ] Given the value of t, and something called the degrees of freedom in your data, you can
determine the p-value (the probability of the difference occurring by chance) using a statistical
table, or, better yet, using SimBio Virtual Labs handy-dandy t-test p-value calculator.
[ 5.1 ] The number of degrees of freedom in your t-test is equal to the total number of
samples (20 in this case) minus 2. That is, degrees of freedom=np+na-2. How many
degrees of freedom do your moose fat stores data have? __________.
[ 5.2 ] Launch the t-test p-value calculator by clicking the t-test button on the TOOLS panel
(very bottom right of your screen).

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[ 5.3 ] In the dialog that appears, type in your t value and the degrees of freedom, and press
the CALCULATE button. What is the probability of the null hypothesis being correct
(i.e., that the difference was due to chance alone)?
[ 5.4 ] What can you say about moose fat stores with wolves absent vs. present, after performing the
t-test?

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Key Publications
A few researchers have studied the population dynamics of wolves and moose on Isle Royale for a very
long time, resulting in an exceptional continuity in research approach and data collection. The research
program is currently directed out of Michigan Tech by John Vucetich and Rolf Peterson, both of whom
have published extensively on moose-wolf population dynamics. Below are a few references regarding
moose and wolves on Isle Royale, the contribution of Isle Royale studies to broader ecological issues,
and the scientific and conservation challenges involved.
Peterson, R.O., & Page, R.E.. 1988. The Rise and Fall of Isle Royale Wolves, 1975-1986. Journal of
Mammology, 69: 89-99.
Peterson, R.O. 1995. The Wolves of Isle Royale: A Broken Balance. Willow Creek Press, Minocqua,
WI.
Vucetich, J.A., R.O. Peterson, & C.L. Schaefer. 2002. The Effect of Prey and Predator Densities on
Wolf Predation. Ecology, 83(11): 3003-3013.
Vucetich, J.A., & R.O. Peterson. 2004. Long-Term Population and Predation Dynamics of Wolves on Isle
Royale. In: D. Macdonald & C. Sillero-Zubiri (eds.), Biology and Conservation of Wild Canids, Oxford
University Press, pp. 281-292.

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Week 13 Ecobeaker Keystone

Introduction

ecology.

A diversity of strange-looking creatures makes


their home in the tidal pools along the edge of
rocky beaches. If you walk out on the rocks at
low tide, youll see a colorful variety of crusty,
slimy, and squishy-looking organisms scuttling
along and clinging to rock surfaces. Their
inhabitants may not be as glamorous as the
megafauna of the Serengeti or the bird life of
Borneo, but these rocky intertidal areas turn
out to be great places to study community

An ecological community is a group of species that live together and interact with each other. Some
species eat others, some provide shelter for their neighbors, and some compete with each other for
food and/or space. These relationships bind a community together and determine the local community
structure: the composition and relative abundance of the different types of organisms present. The
intertidal community is comprised of organisms living in the area covered by water at high tide and
exposed to the air at low tide.
This laboratory is based on a series of famous experiments that were conducted in the 1960s along the
rocky shore of Washington state, in the northwestern United States. Similar intertidal communities
occur throughout the Pacific Northwest from Oregon to British Columbia in Canada. The nine species in
this laboratorys simulated rocky intertidal area include three different algae (including one you may
have eaten in a Japanese restaurant); three stationary (or sessile) filter-feeders; and three mobile
consumers.
Ecological communities are complicated, and the rocky intertidal community is no exception.
Fortunately, carefully designed experiments can help us tease apart these complexities, providing
insight into how communities function. As will become apparent, understanding the factors that govern
community structure can have serious implications for management. In this laboratory, youll use
simulated experiments to elucidate how interactions between species can play a major role in
determining community structure. You will apply techniques similar to those used in the original studies,
in order to experimentally determine which species in the simulated rocky intertidal are competitively

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dominant over which others. Youll then analyze gut contents and use your data to construct a food web
diagram. Finally, youll conduct removal experiments, observing how the elimination of particular
species influences the rest of the community. When youve completed this lab, you should have a
greater appreciation for the underlying complexity of communities, and for how the loss of single
species can have surprisingly profound impacts.

Food Chains, Food Webs and Trophic Levels

You probably know that herbivores eat plants and that predators eat herbivores. The progression of
what eats what, from plant to herbivore to predator, is an example of a food chain. Omnivores eat both
plants and animals. Within a community, producers, herbivores, predators, and omnivores are linked
through their feeding relationships. If you create a diagram that connects different species and food
chains together based on these relationships, the result is called a food web diagram.
Ecosystems can also be represented by a pyramid comprising a series of trophic levels. A species
trophic level indicates its relative position in the ecosystems food chain. Producers (including algae and
green plants) use energy from the sun to produce their own food rather than consuming other
organisms, thus they occupy the lowest trophic level. Since herbivores consume the producers, they
occupy the next trophic level. Predators eat the herbivores, thus occupying the next higher trophic level.
Omnivores occupy multiple trophic levels. The highest level is occupied by top-level predators, which are
not eaten by anything (until they die). Generally, but not always, lower trophic levels have more species
than do higher levels within a community.

Competition

Among community relationships, predation is perhaps the most obvious but certainly not the most
important. Two species may also compete with each other for space or food. Stationary organisms in
particular must often compete intensively for limited space. When one species is better at obtaining or
holding space than another, or is able to displace the second species, the winner is said to be
competitively dominant. In the same way that you can draw a food web, you can also construct a
diagram to illustrate which species are superior competitors within a community, called a competitive
dominance hierarchy. In this lab, you will create competition dominancy hierarchy and food web
diagrams to help you understand the community structure of the intertidal zone.

Dominant versus Keystone Species


In many communities, there is one species that is more abundant in number or biomass than any other,
often referred to as the dominant species. For example, in a dense, old-growth forest, one type of late
successional tree is often the dominant species. As you might imagine, such species greatly affect the
nature and composition of the community.
However, a species does not have to be the most abundant to have the greatest impact on the
community. Imagine an archway made of stones. The one stone at the top center of the arch supports
all the other stones. If you remove that stone, called the keystone, the arch crumbles. In some
communities, the presence of a single species controls community structure even though that species
may have relatively low abundance. These organisms are known as keystone species. An important
characteristic of a keystone species is that its decline or removal will drastically alter the structure of the
local community. For example, many keystone species are top predators that keep the populations of
lower-level consumers in check. If top predators are removed, populations of the lower-level organisms
can grow, dramatically changing species diversity and overall community structure, sometimes resulting
in the collapse of the entire community.

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The EcoBeaker Model

If youre curious about how the simulated intertidal community works, heres the basic idea. In
EcoBeaker models, each individual belongs to a species which is defined by a collection of rules that
determine that species behavior. For example, species that are mobile consumers follow rules that
dictate how far they can move in a time step, what they can eat, how much energy they obtain from
their prey, how much energy they use when they move, etc. When an individual consumers energy runs
out, it dies. Individuals within species all follow the same rules, but because the rules defining species
include some random chance (e.g., which direction to turn), you will notice variability in what individuals
are doing at any given time. Different species behave differently because they dont have the same
parameters assigned for their rules (e.g., they might eat different species or move slower or faster).
The six stationary species in the modelthe three algae and the three filter feeders (or sessile
consumers)are modeled differently than the mobile consumers. The simulation uses a transition
matrix for these six stationary species. The transition matrix is a set of probabilities that determine what
happens from one time step to the next on a particular space on the rock. For each species, the
transition matrix lists the probability of an individual of that species settling on top of bare rock, the
probabilities of being replaced by each of the other species, and the probability of dying (and being
replaced by bare rock). In addition, the transition matrix includes the probability of bare rock remaining
bare. For example, a patch of rock that contains Nori Seaweed (Porphyra) may do one of three things
each time step: host a different species (that is, another organism displaces Nori Seaweed), continue to
be occupied by Nori Seaweed, or become bare rock (the Nori Seaweed dies and is not replaced). Each of
these changes, or transitions, has a probability associated with it included within the transition matrix. If
one species out-competes another for space, this will be reflected in the relevant transition probability.

More Information

Links to additional terms and topics relevant to this laboratory can be found in the Keystone Predator
Library accessible via the SimBio Virtual Labs program interface.

Starting Up
If youve explored tide pools (a fun thing to do if you visit a rocky coast), you likely know that many of
the plants and animals living in them are unusual. This section will introduce you to the different species
youll encounter in this lab.
[ 1 ] Make sure that you have read the introductory section of the workbook. The background
information and introduction to ecological concepts will help you understand the simulation
model and answer questions correctly.
[ 2 ] Start the program by double-clicking the SimBio Virtual Labs icon on your computer or by selecting
it from the Start Menu.
[ 3 ] When SimBio Virtual Labs opens, select the Keystone Predator lab from the EcoBeaker suite.
IMPORTANT!
Before you continue, make sure you are using the SimBio Virtual Labs version of Keystone Predator. The
splash screen for SimBio Virtual Labs looks similar to this:

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If the splash screen you see does not look like this, please close the application (EcoBeaker 2.5) and
launch SimBio Virtual Labs.
You will see a number of different panels on the screen:
The left side of the screen shows a view of an intertidal zone area. This is where you
will be conducting your experiments and observing the action.
A bar graph on the right shows the population sizes of all of species in the intertidal
area.
Above the graph is a list of the plant and animal species included in the simulation.
You can switch between Latin and common names of each species using the tabs
above the species list. The workbook will refer to common names.
In the bottom left corner of the screen is the CONTROL PANEL. To the right of the
Control Panel is a set of TOOLS that you will use for doing your experiments. These
will be described in the following exercises, as you need them.
[ 4 ] Click on the names in the SPECIES LEGEND in the upper right corner of the screen to bring up library
pages for each species. Use the library to answer the following question:
[ 4.1 ] If you slipped on a rock while exploring a tide pool and your knee became inflamed,
which of the three algal species might help reduce the swelling?
Nori Black Pine Coral Weed (Circle one)
[ 4.2 ] Use the information in the Introduction and Library pages to fill in the blank spaces in
the table at the top of the next page.
HELPFUL HINT: producers are organisms that generate their own food using energy from the sun,
filter-feeders are consumers that extract food particles out of the water, and stationary (or sessile)
organisms do not actively move around (at least not as adults) they permanently adhere to substrates
such as rocks.

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[ 5 ] When you are done completing the above table, start the simulation by clicking the GO button in
the CONTROL PANEL. Watch the action for a bit. Notice how the mobile consumers clear off
areas of rock by eating, and how the stationary species recolonize those areas.
[ 6 ] Click the STOP button to pause the simulation.
Look at the population graph. The Population Size Index represents the number of
individuals of each species present in the simulation. For the three algal species, a more
appropriate measure of relative abundance might be percent cover or biomass, because in the
real world, a single alga can grow quite large. The EcoBeaker model simulates algal growth as
individuals multiplying, which, though not exactly realistic, makes possible the comparison of
population sizes for the three algal and six animal species.
[ 7 ] Click the RESET button to return the community to its initial state. Then move your mouse over to
the population graph and click on one of the bars. You will see the population size for that
species displayed.
[ 7.1 ] Which species in the simulation has the largest population? ___________
[ 7.2 ] What is the size of the population for that species? ___________
[ 7.3 ] Which species in the simulation has the smallest population? ___________
[ 7.4 ] What is the size of the population for that species? ___________

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Exercise 1: Flexing Your Mussels


In the competitive dominance hierarchy diagram below, the arrows point from weaker to
stronger competitors. For example, the arrow pointing from Nori Seaweed to Black Pine indicates that
Black Pine is dominant over (i.e., can displace) Nori Seaweed. The diagram also shows that any of the
sessile consumers (the barnacles and the mussel) can out-compete any of the algae.

[ 1 ] According to the figure above, which algal species is the strongest competitor?
Nori Seaweed Black Pine Coral Weed (Circle one)
The diagram does not indicate the dominance hierarchy among the three sessile consumers.
Fortunately, you have some tools that will let you experimentally determine their competitive
relationships. Your approach will involve creating patches of each sessile consumer on an intertidal rock
and observing which of the other two sessile consumers successfully invades those patches.
[ 2 ] Select Flexing Your Mussels from the Select an Exercise menu at the top of the screen to load the
experimental system.
You should now see only the six stationary species in the simulation (three algae and three
sessile consumers)this is because your assistant is patrolling the shore and keeping the

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mobile consumers out of your experimental area. Excluding these predators will help you
determine the competitive relationships among the sessile consumers.
[ 3 ] Find and click the ADD SESSILE CONSUMER button in the TOOLS PANEL. If you have trouble finding
a particular button in the lab, move your mouse over buttons and tool tips will appear. The
ADD SESSILE CONSUMER button will initially have an Acorn Barnacle picture on it. If a different
species appears, click the downward arrow next to the picture and select the Acorn Barnacle.
[ 4 ] Click somewhere in the Intertidal Zone area, hold down the mouse button, and then drag out a
rectangle to create a solid patch of Acorn Barnacles. The rectangle should fill at least a third of
the Intertidal Zone area and have no other species inside.
[ 5 ] Use the STEP button to advance the simulation one week at a time for ten weeks and monitor
which other species displace Acorn Barnacles through time. These species are stealing rock
space from the Acorn Barnacles, and thus are competitively dominant.
[ 5.1 ] Circle the species which are competitively dominant over Acorn Barnacles:

[ 6 ] RESET the simulation. Click the ADD SESSILE CONSUMER button and select the Goose Neck
Barnacle.
[ 7 ] Repeat steps 4 and 5 for Goose Neck Barnacles.
[ 7.1 ] Circle the species that are competitively dominant over Goose Neck Barnacles:

[ 8 ] RESET the simulation. Click the ADD SESSILE CONSUMER button and select the Mussel.
[ 9 ] Repeat steps 4 and 5 for Mussels.
[ 9.1 ] Circle the species that are competitively dominant over Mussels:

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[ 10 ] Construct a competitive dominance hierarchy diagram using the information you have gathered
from your experiments. Next to each species name, indicate how many arrows point to that species. The
highest number indicates the most highly ranked and aggressive, or best, competitor. If you do this
correctly, each species should have a different rank.

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Exercise 2: You Are What You Eat


Now you will continue to investigate this intertidal community by determining what each of the mobile
consumer species eats. One trick ecologists use to find out what creatures eat is to look at whats in
their guts or excrement. If you eat something, it hangs around in your stomach for a little while, and
then (later) the undigested parts come out the other end. Normally, when researchers look at gut
contents they have to kill the animal, cut it open, and examine what is inside (people get paid to do
this!). Within SimBio Virtual Labs there is a kinder and gentler method for determining gut contents.
[ 1 ] Select You Are What You Eat from the Select an Exercise menu at the top of the screen.
[ 2 ] You will now see all nine species in the simulation: three algae, three sessile consumers, and three
mobile consumers: Starfish, Whelk, and Chiton.
[ 3 ] Start running the simulation (click GO).
[ 4 ] Watch the action for about 100 weeks and monitor the abundance of species in the population
graph. Notice how the population index for each species fluctuates and eventually settles at a
relatively stable level.
[ 5 ] STOP the simulation.
[ 6 ] Click the MICROSCOPE (VIEW ORGANISM) button in the TOOLS PANEL to activate your mobile
Gut-o-Scope (patent pending).
[ 7 ] Click your favorite Starfish, Whelk, or Chiton your choice!
A window will appear with gut content information for that individual, either identifying the
predators last prey item or indicating that the gut is empty (because the creature has not
eaten recently). Note that if you click on organisms that dont have guts (algae or filter
feeders), you wont see gut contents.
[ 8 ] You will now conduct a survey of the three mobile consumers to learn which species they eat. The
data forms below will help you record and summarize your findings. The forms are divided into three
sections, one for each mobile consumer.
[ 8.1 ] In each data table section, record gut content data for 10 randomly-selected individuals
of that species. Ignore individuals that have not eaten recently (indicated by gut contents labeled
empty). If recorded correctly, each row of the data form should have one species circled. At the
bottom of each section, record the total number of individuals circled in each column.

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[ 8.2 ] For each mobile consumer species below, record its prey and the percentage of diet each prey
species comprises for that consumer (e.g., 4 out of 10 samples = 40%). The numbers in each column
should add up to 100%.

[ 9 ] You now have enough information to construct a food web diagram from your findings. Consider
the hypothetical example below. In a forest, both deer and rabbits eat the plants. Wolves, the predators
in the system, eat both deer and rabbits. We can draw these feeding relationships like this, with arrows
pointing to the consumer:

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[ 9.1 ] Use your data on feeding relationships to construct a food web diagram for the organisms that
live in the simulated intertidal zone. Link the species names below with arrows that point from prey
to consumer. (Unlike the simple four-species example on the previous page, your nine-species
diagram will look more complicated, with many crossing lines.)

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Experiment 3: Who Rules the Rock?


Based on your studies so far, you know important details about the competitive and feeding
relationships among the species in your simulated intertidal community, and these relationships, when
integrated, define the role played by each. One way to more fully elucidate the importance of a species
to its community structure is to remove it from the environment and observe what happens. In this
exercise, you will experimentally determine how removing each of the highest trophic level species (the
mobile consumers) affects the rocky intertidal community structure.
[ 1 ] Before you start your experiments, first make some predictions. Refer back to your data to inform
your answers. [NOTE: only one species will be removed in each experiment.]
[ 1.1 ] In the spaces provided below, predict which other species in the community will be
impacted the most by each removal and explain your reasoning.
Predicted impact of removing Whelk and explanation:

Predicted impact of removing Chiton and explanation:

Predicted impact of removing Starfish and explanation:

[ 1.2 ] One removal experiment will have a more dramatic impact than the other two. Write down
which one you predict this will be, and why:

[ 2 ] Select Who Rules the Rock? from the Select an Exercise menu.
[ 3 ] Your first step is to record population sizes BEFORE REMOVALS. To make sure the simulation is
initialized correctly, click the RESET button. A data table is provided on the next page for
recording your results.
HELFUL HINT: if you click on the colored bars in the Population Size graph, the numbers
(population sizes) that the bars represent will pop up!

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[ 3.1 ] In the table on the next page, record the population size of each species at Time
Elapsed = 0 Weeks in the BEFORE REMOVALS column.
[ 4 ] After recording data BEFORE REMOVALS, you are ready to remove mobile consumers. Find the
REMOVE WHELK button (which is round and depicts a Whelk with a slash through it) in the
TOOLS PANEL. When you click this button, all Whelk will vanish from the Intertidal Zone.
[ 5 ] For each removal experiment, you will run the simulation for 200 weeks (in model time, not real
time!). To do this, first make sure that the Time Elapsed = 0 weeks (RESET if not), and then click
the STEP 200 button in the CONTROL PANEL..
HELPFUL HINT: If your computer is a little slow, you can speed things up using the
Speed Slider to the right of the CONTROL PANEL
[ 6 ] Confirm that the simulation stopped at (or near) 200 weeks. If so, click the bars in the Population
Size graph and record the abundance of each species.
[ 6.1 ] In the data table, record the population size of each species in the AFTER WHELK
REMOVAL column.
[ 7 ] RESET the simulation and confirm that Time Elapsed = 0 weeks. Then click the REMOVE CHITON
button to remove all Chiton from the Intertidal Zone.
[ 8 ] Click the STEP 200 button to run the simulation for 200 weeks.
[ 8.1 ] When Time Elapsed = 200 weeks, record the population size of each species in the
AFTER CHITON REMOVAL column.
[ 9 ] Finally, RESET the simulation and use the REMOVE STARFISH tool and the STEP 200 button to
repeat the experiment for Starfish.
[ 9.1 ] In the data table, record the population size of each species in the AFTER STARFISH REMOVAL
column.

[ 10 ] When your data table is complete, answer the following questions. Try to be as quantitative as
possible with your answers, indicating by approximately how much each species increased or
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decreased in size (e.g., The Starfish population more than doubled; The population of Coral
Weed decreased to about half its original size.).
[ 10.1 ] What were the most dramatic changes to the community after Whelk were removed?

[ 10.2 ] What were the most dramatic changes to the community after Chiton were removed?

[ 10.3 ] What were the most dramatic changes to the community after Starfish were removed?

[ 10.4 ] Which removal had the greatest impact upon the rest of the community?

[ 10.5 ] Referring back to your competitive dominance hierarchy and food web diagrams, try to
explain what happened in the removal experiment that had the greatest impact on community
structure. Why was the effect so pronounced?

[ 10.6 ] Look back at what you predicted would happen when you removed each of the three mobile
consumer species in Step 1 above. Were you correct for all three? If not, describe what you think you
missed in each case.

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The Keystone Species Concept

As described in the Introduction, a keystone is the stone in the middle of the top of an arch that
supports all the other stones. If you remove the keystone, the whole arch falls down. In many ecological
communities, one species can play a particularly important role in determining and supporting
community structure. Remove this species and the community structure changes radically. When you
removed one mobile consumer from the intertidal simulation, it had a much larger impact on the rest of
the community compared to the removal of the other two mobile consumers. This species is an example
of a Keystone Species, which has a disproportionately large impact on its ecological community
compared to its relatively low abundance.
The keystone predator in the intertidal zone you studied occupies a position at the top of the food
chain, which is common for keystone species. They also tend to be susceptible to both natural and
human disturbance. Other examples of the dramatic effects of keystone species removals in the real
world include deer populations rapidly increasing with the local extinction of predatory wolves,
kangaroo rats dominating rodent communities with the removal of coyotes, and sea urchins decimating
kelp forests when predatory sea otters go into decline.

Species Reintroduction
[ 11 ] Conservationists sometimes advocate reintroducing native species that have gone extinct locally. A
well-known recent example is the reintroduction of wolves into Yellowstone National Park.
[ 11.1 ] If you were to reintroduce Starfish into the Intertidal Zone, what do you think would
happen to the intertidal community? Write your prediction in the space provided
below:

[ 12 ] RESET the simulation, REMOVE STARFISH, and STEP 200 weeks forward (this returns the
simulation to its state at the end of the previous exercise). Click the ADD MOBILE CONSUMER tool and
select Starfish.
[ 13 ] Move your mouse into the Intertidal Zone and click six or seven times to add some Starfish back
into the community.
[ 14 ] Use the STEP 200 button to advance the simulation another 200 weeks and watch what happens
to the population sizes of the different species.
[ 14.1 ] Briefly describe the changes you observed in abundance of different species.

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[ 14.2 ] Look at the population sizes and describe how they compare to your BEFORE REMOVAL
population data when you first ran the simulation before removing any of the mobile consumers.

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Biocontrol

Organisms that are introduced to eliminate or otherwise limit population growth for unwanted pest
species are known as biocontrol agents. Biocontrol agents are often parasites, predators, or
pathogens. Ideally, when biocontrol agents are introduced into an ecological community, the unwanted
species will be controlled without further intervention and the system will be self-sustaining. Use of
biocontrol agents is not without risk however, and biologists must be very careful that the introduced
biocontrol organism will not further negatively impact the native species and become a pest itself. There
are plenty of nightmare examples from our past where introduced biocontrol agents have become a
bigger problem than the species they were introduced to control. Famous examples include the
mongoose in Hawaii, which was introduced to eat rats but preferred eating the native bird fauna; and
cane toads in Australia, which were introduced to control a sugar cane beetle, but instead had
devastating impacts on indigenous amphibian and reptile communities and did almost nothing to
control the insect pests.
There is a parasitic barnacle, Sacculina, which research has suggested might be useful as a biocontrol
agent for invasive European green crabs. The larvae of Sacculina settle on crabs, piercing the
exoskeleton. This type of infestation slows down the crabs ability to reproduce and therefore, over time
should lead to a decrease in the crab population size.
[ 2.2 ] Having investigated the impacts of the European Green Crab on the intertidal zone community,
the Department of Fish and Wildlife is considering using parasitic barnacles (Sacculina) to control the
crab. Given what you know about the importance of species interactions to community structure,
what would you suggest should be learned about this parasitic barnacle species before it is introduced
into the intertidal community you have been studying?

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Key Publications
This laboratory was inspired by the following classic papers by R.T. Paine, whose work on intertidal
communities originated the idea of keystone predation:
Paine, R.T. 1966. Food Web Complexity and Species Diversity. The American Naturalist 100: 65-75
Paine, R.T. 1969. The Pisaster-Tegula Interaction: Prey Patches, Predator Food Preference, and
Intertidal Community Structure. Ecology 50: 950-961.
The following review article addresses the idea of keystone species from a more modern perspective:
Power, M.E., D. Tilman, J.A. Estes, B.A. Menge, W.J. Bond, L.S. Mills, G. Daily, J.C. Castilla, J. Lubchenco,
R.T. Paine. 1996. Challenges in the Quest For Keystones: Identifying Keystone Species is Difficult But
Essential To Understanding How Loss of Species Will Affect Ecosystems. BioScience 46: 609-620.

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Week 14 Thanksgiving

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Week 15 Review

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