Professional Documents
Culture Documents
ECOLOGICAL
METHODS
HANDBOOK
by
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
Description
Objectives
1.
2.
3.
4.
5.
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
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:
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.
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
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
Green Ash
Fraxinus pennsylvanica
13
14
Physicochemical Analysis
Dissolved Oxygen (YSI Model 55 DO Meter)
1.
2.
3.
4.
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.
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
Sample 1
Sample 2
Sample 3
Sample 4
Sample 5
Depth (m)
Water
Temperature
(oC)
Dissolved
Oxygen (mg/l)
pH
Conductivity
16
Method
17
pi*ln(pi)
-0.298626578
-0.160576209
-0.244136064
-0.244136064
-0.367290626
-0.160576209
-0.244136064
-1.719477814
1.719477814
(1)
(1)
and where n = the total number of organisms of a species while N = the total
84
21(20)
= 0.2
18
19
Conductivity meter
Dip net
Mechanical flowmeter
Seine
20
Magnifying loupe
pH meter
Ponar grab
Spectrometer
Plankton net
Kick net
21
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
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
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
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)
2)
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)
C) We will pool the data from all quadrats, calculate the following for each tree species, and record those
values on Table 3.
1)
2)
3)
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)
8)
Total number of trees of all species combined per hectare (a hectare is 10,000m2)
9)
32
Location ________________________________________
Date _______________________________
Number
Species
DBH
33
34
Location ________________________________________
Date _______________________________
Number
Species
DBH
35
36
Location ________________________________________
Date _______________________________
Number
Species
DBH
37
38
Table 2 - Summary of Community Analysis of Single Quadrat by the Quadrat Sampling Technique (#1)
Location ________________________________________
Date _______________________________
Species
Density
Total Basal
Area Covered
Importance
(D + Do) / 2
39
Species
Density
Total Basal
Area Covered
Importance
(D + Do) / 2
Totals
40
Table 2 - Summary of Community Analysis of Single Quadrat by the Quadrat Sampling Technique (#2)
Location ________________________________________
Date _______________________________
Species
Density
Total Basal
Area Covered
Species
Density
Total Basal
Importance
(D + Do) / 2
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 ________________________________________
Date _______________________________
Species
Density
Total Basal
Area Covered
Species
Density
Total Basal
Importance
(D + Do) / 2
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 _____________________________
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
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 do relative dominance and relative density values compare for different species?
Are there differences between the seedling or sapling numbers in different quadrats?
48
Compass
Dbh tape
Tape measure
Note that dbh (diameter at breast height) is sampled 1.4 m above the ground.
49
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.
50
Location ________________________________________
Date _______________________________
Number
Species
DBH
51
52
Location ________________________________________
Date _______________________________
Number
Species
DBH
53
54
Location ________________________________________
Date _______________________________
Number
Species
DBH
55
56
Table 2 - Summary of Community Analysis of Single Quadrat by the Quadrat Sampling Technique (#1)
Location ________________________________________
Date _______________________________
Species
Density
Total Basal
Area Covered
Importance
(D + Do) / 2
57
Species
Density
Total Basal
Area Covered
Importance
(D + Do) / 2
Totals
58
Table 2 - Summary of Community Analysis of Single Quadrat by the Quadrat Sampling Technique (#2)
Location ________________________________________
Date _______________________________
Species
Density
Total Basal
Area Covered
Importance
(D + Do) / 2
59
Species
Density
Total Basal
Area Covered
Importance
(D + Do) / 2
Totals
60
Table 2 - Summary of Community Analysis of Single Quadrat by the Quadrat Sampling Technique (#3)
Location ________________________________________
Date _______________________________
Species
Density
Total Basal
Area Covered
Importance
(D + Do) / 2
61
Species
Density
Total Basal
Area Covered
Importance
(D + Do) / 2
Totals
62
Table 3 - Summary of Community Analysis of All Quadrats by the Quadrat Sampling Technique
Location ___________________________
Date _____________________________
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
64
Questions
What species had the highest relative frequency, density and dominance in each habitat (riparian vs. crosstimbers forest)?
65
Create a Shannon-weiner and Simpson diversity index for each habitat. Which habitat (riparian or cross-timbers)
had the highest diversity?
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 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
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
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.) =
40.9
45
40.9
20
= 2.05
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)
69
2)
3)
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)
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
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.
79
How do relative dominance and relative density values compare for different species?
80
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
Vegetation lab
81
82
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
Taxa
1A
3A
5A
7A
9A
No. of individuals
No. of species
Species diversity
84
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
Taxa
1C
3C
5C
7C
9C
No. of individuals
No. of species
Species diversity
88
Taxa
2C
4C
6C
8C
10C
No. of individuals
No. of species
Species diversity
89
Questions
1.
2.
3.
Which area (prairie or forest) had the highest species diversity value?
4.
5.
6.
7.
90
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
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
Nine-banded Amradillo
Dasypus novemcinctus
Virginia Oppossum
Didelphis virginiana
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
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
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European Starling
Sturnus vulgaris
Red-tailed Hawk
Buteo jamaicensis
Eastern Screech-Owl
Megascops asio
House Sparrow
Passer domesticus
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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
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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.
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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).
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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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[ 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):
[ 6.2 ] 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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[ 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?
[ 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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[ 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?
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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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[ 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:
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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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Introduction
ecology.
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.
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.
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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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[ 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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[ 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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[ 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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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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