1. 41
Species Interactions

2. Overview: Communities in Motion
A biological community is an assemblage of populations of various species living close enough for potential interaction
For example, the “carrier crab” carries a sea urchin on its back for protection against predators 2

3. Figure 41.1
Figure 41.1 Which species benefits from this interaction? 3

4. Concept 41.1: Interactions within a community may help, harm, or have no effect on the species involved
Ecologists call relationships between species in a community interspecific interactions
Examples are competition, predation, herbivory, symbiosis (parasitism, mutualism, and commensalism), and facilitation
Interspecific interactions can affect the survival and reproduction of each species, and the effects can be summarized as positive (), negative (−), or no effect (0) 4

5. Competition
Interspecific competition (−/− interaction) occurs when species compete for a resource that limits their growth or survival 5

6. Competitive Exclusion
Strong competition can lead to competitive exclusion, local elimination of a competing species
The competitive exclusion principle states that two species competing for the same limiting resources cannot coexist in the same place 6

7. Ecological Niches and Natural Selection
Evolution is evident in the concept of the ecological niche, the specific set of biotic and abiotic resources used by an organism
An ecological niche can also be thought of as an organism’s ecological role
Ecologically similar species can coexist in a community if there are one or more significant differences in their niches 7

8. Resource partitioning is differentiation of ecological niches, enabling similar species to coexist in a community 8

9. A. distichus perches on fence posts and other sunny surfaces.
Figure 41.2
A. distichus perches on
fence posts and other
sunny surfaces.
A. insolitus usually
perches on shady
branches.
A. ricordii
Figure 41.2 Resource partitioning among Dominican Republic lizards
A. insolitus
A. aliniger
A. christophei
A. distichus
A. cybotes
A. etheridgei 9

10. Figure 41.2a
A. distichus
Figure 41.2a Resource partitioning among Dominican Republic lizards (part 1: A. distichus) 10

11. Figure 41.2b
A. insolitus
Figure 41.2b Resource partitioning among Dominican Republic lizards (part 2: A. insolitus) 11

12. A species’ fundamental niche is the niche potentially occupied by that species
A species’ realized niche is the niche actually occupied by that species
As a result of competition, a species’ fundamental niche may differ from its realized niche
For example, the presence of one barnacle species limits the realized niche of another species 12

13. Experiment High tide Chthamalus Balanus Chthamalus realized niche
Figure 41.3
Experiment
High tide
Chthamalus
Balanus
Chthamalus
realized niche
Balanus
realized niche
Ocean
Low tide
Results
High tide
Figure 41.3 Inquiry: Can a species’ niche be influenced by interspecific competition?
Chthamalus
fundamental niche
Ocean
Low tide 13

14. Character Displacement
Character displacement is a tendency for characteristics to be more divergent in sympatric populations of two species than in allopatric populations of the same two species
An example is variation in beak size between populations of two species of Galápagos finches 14

15. Santa María, San Cristóbal Sympatric populations 40
Figure 41.4
G. fuliginosa
G. fortis
Beak
depth 60
Los Hermanos 40
G. fuliginosa,
allopatric 20 60
Daphne
Percentages of individuals in each size class 40
G. fortis,
allopatric 20
Figure 41.4 Character displacement: indirect evidence of past competition 60
Santa María, San Cristóbal
Sympatric
populations 40 20 8 10 12 14 16
Beak depth (mm) 15

16. Predation
Predation (/− interaction) refers to an interaction in which one species, the predator, kills and eats the other, the prey
Some feeding adaptations of predators are claws, teeth, stingers, and poison 16

17. Prey display various defensive adaptations
Behavioral defenses include hiding, fleeing, forming herds or schools, and active self-defense
Animals also have morphological and physiological defense adaptations
Cryptic coloration, or camouflage, makes prey difficult to spot 17

18. Video: Sea Horses

19. (c) Batesian mimicry: A harmless species mimics a harmful one.
Figure 41.5
(a) Cryptic
coloration
(b) Aposematic
coloration
Canyon
tree frog
Poison
dart frog
(c) Batesian mimicry: A harmless species mimics
a harmful one.
Nonvenomous
hawkmoth larva
(d) Müllerian mimicry: Two unpalatable
species mimic each other.
Venomous green
parrot snake
Cuckoo bee
Yellow jacket
Figure 41.5 Examples of defensive coloration in animals 19

20. (a) Cryptic coloration Canyon tree frog Figure 41.5a
Figure 41.5a Examples of defensive coloration in animals (part 1: cryptic coloration) 20

21. Animals with effective chemical defenses often exhibit bright warning coloration, called aposematic coloration
Predators are particularly cautious in dealing with prey that display such coloration 21

22. (b) Aposematic coloration Poison dart frog Figure 41.5b
Figure 41.5b Examples of defensive coloration in animals (part 2: aposematic coloration) 22

23. In some cases, a prey species may gain significant protection by mimicking the appearance of another species
In Batesian mimicry, a palatable or harmless species mimics an unpalatable or harmful model 23

24. (c) Batesian mimicry: A harmless species mimics a harmful one.
Figure 41.5c
(c) Batesian mimicry: A harmless species mimics
a harmful one.
Nonvenomous
hawkmoth larva
Venomous green
parrot snake
Figure 41.5c Examples of defensive coloration in animals (part 3: Batesian mimicry) 24

25. Nonvenomous hawkmoth larva Figure 41.5ca
Figure 41.5ca Examples of defensive coloration in animals (part 3a: Batesian mimicry, hawkmoth)
Nonvenomous hawkmoth
larva 25

26. Venomous green parrot snake
Figure 41.5cb
Figure 41.5cb Examples of defensive coloration in animals (part 3b: Batesian mimicry, snake)
Venomous green parrot snake 26

27. In Müllerian mimicry, two or more unpalatable species resemble each other27

28. (d) Müllerian mimicry: Two unpalatable species mimic each other.
Figure 41.5d
(d) Müllerian mimicry: Two unpalatable
species mimic each other.
Cuckoo bee
Yellow jacket
Figure 41.5d Examples of defensive coloration in animals (part 4: Müllerian mimicry) 28

29. Figure 41.5da
Figure 41.5da Examples of defensive coloration in animals (part 4a: Müllerian mimicry, cuckoo bee)
Cuckoo bee 29

30. Yellow jacket Figure 41.5db
Figure 41.5db Examples of defensive coloration in animals (part 4b: Müllerian mimicry, yellow jacket)
Yellow jacket 30

31. Herbivory
Herbivory (/− interaction) refers to an interaction in which an herbivore eats parts of a plant or alga
In addition to behavioral adaptations, some herbivores may have chemical sensors or specialized teeth or digestive systems
Plant defenses include chemical toxins and protective structures 31

32. Figure 41.6
Figure 41.6 A marine herbivore 32

33. Symbiosis
Symbiosis is a relationship where two or more species live in direct and intimate contact with one another 33

34. Parasitism
In parasitism (/− interaction), one organism, the parasite, derives nourishment from another organism, its host, which is harmed in the process
Parasites that live within the body of their host are called endoparasites
Parasites that live on the external surface of a host are ectoparasites 34

35. Many parasites have a complex life cycle involving multiple hosts
Some parasites change the behavior of the host in a way that increases the parasites’ fitness
Parasites can significantly affect survival, reproduction, and density of host populations 35

36. Mutualism
Mutualistic symbiosis, or mutualism (/ interaction), is an interspecific interaction that benefits both species
In some mutualisms, one species cannot survive without the other
In other mutualisms, both species can survive alone
Mutualisms sometimes involve coevolution of related adaptations in both species 36

37. Video: Clownfish and Anemone

38. (a) Ants (genus Pseudomyrmex) in acacia tree
Figure 41.7
Figure 41.7 Mutualism between acacia trees and ants
(a) Ants (genus Pseudomyrmex) in
acacia tree
(b) Area cleared by ants around an
acacia tree 38

39. (a) Ants (genus Pseudomyrmex) in acacia tree
Figure 41.7a
Figure 41.7a Mutualism between acacia trees and ants (part 1: ants)
(a) Ants (genus Pseudomyrmex) in acacia tree 39

40. (b) Area cleared by ants around an acacia tree
Figure 41.7b
Figure 41.7b Mutualism between acacia trees and ants (part 2: acacia tree)
(b) Area cleared by ants around an acacia tree 40

41. Commensalism
In commensalism (/0 interaction), one species benefits and the other is neither harmed nor helped
Commensal interactions are hard to document in nature because any close association likely affects both species 41

42. Figure 41.8
Figure 41.8 A possible example of commensalism between cattle egrets and African buffalo 42

43. Facilitation
Facilitation (/ or 0/) is an interaction in which one species has positive effects on another species without direct and intimate contact
For example, the black rush makes the soil more hospitable for other plant species 43

44. With Juncus Without Juncus
Figure 41.9 8 6
Number of plant species 4 2
Figure 41.9 Facilitation by black rush (Juncus gerardi) in New England salt marshes
(a) Salt marsh with Juncus
(foreground)
With
Juncus
Without
Juncus
(b) 44

45. (a) Salt marsh with Juncus (foreground)
Figure 41.9a
Figure 41.9a Facilitation by black rush (Juncus gerardi) in New England salt marshes (photo)
(a) Salt marsh with Juncus
(foreground) 45

46. Concept 41.2: Diversity and trophic structure characterize biological communities
Two fundamental features of community structure are species diversity and feeding relationships
Sometimes a few species in a community exert strong control on that community’s structure 46

47. Species Diversity
Species diversity of a community is the variety of organisms that make up the community
It has two components: species richness and relative abundance
Species richness is the number of different species in the community
Relative abundance is the proportion each species represents of all individuals in the community 47

48. A B C D Community 1 Community 2 A: 25% B: 25% C: 25% D: 25% A: 80%
Figure 41.10 A B C D
Community 1
Community 2
Figure Which forest is more diverse?
A: 25%
B: 25%
C: 25%
D: 25%
A: 80%
B: 5%
C: 5%
D: 10% 48

49. H  −(pA ln pA  pB ln pB  pC ln pC  …)
Two communities can have the same species richness but a different relative abundance
Diversity can be compared using a diversity index
Widely used is the Shannon diversity index (H)
H  −(pA ln pA  pB ln pB  pC ln pC  …)
where A, B, C are the species, p is the relative abundance of each species, and ln is the natural logarithm 49

50. Molecular tools can be used to help determine microbial diversity
Determining the number and relative abundance of species in a community is challenging, especially for small organisms
Molecular tools can be used to help determine microbial diversity 50

51. Figure 41.11
Results
3.6
3.4
3.2
Shannon diversity (H)
3.0
2.8
Figure Research method: determining microbial diversity using molecular tools
2.6
2.4
2.2 3 4 5 6 7 8 9
Soil pH 51

52. Diversity and Community Stability
Ecologists manipulate diversity in experimental communities to study the potential benefits of diversity
For example, plant diversity has been manipulated at Cedar Creek Natural History Area in Minnesota for two decades 52

53. Figure 41.12
Figure Study plots at the Cedar Creek Natural History Area, site of long-term experiments in which researchers have manipulated plant diversity 53

54. Communities with higher diversity are
More productive and more stable in their productivity
Able to produce biomass (the total mass of all individuals in a population) more consistently than single species plots
Better able to withstand and recover from environmental stresses
More resistant to invasive species, organisms that become established outside their native range 54

55. Trophic Structure
Trophic structure is the feeding relationships between organisms in a community
It is a key factor in community dynamics
Food chains link trophic levels from producers to top carnivores 55

56. Quaternary consumers: carnivores
Figure 41.13
Quaternary consumers:
carnivores
Tertiary consumers:
carnivores
Secondary consumers:
carnivores
Figure Examples of terrestrial and marine food chains
Primary consumers:
herbivores and zooplankton
Primary producers:
plants and phytoplankton 56

57. A food web is a branching food chain with complex trophic interactions
Species may play a role at more than one trophic level 57

58. Video: Shark Eating a Seal

59. Humans Smaller toothed whales Baleen whales Sperm whales Elephant
Figure 41.14
Humans
Smaller
toothed
whales
Baleen
whales
Sperm
whales
Elephant
seals
Crab-
eater
seals
Leopard
seals
Birds
Fishes
Squids
Figure An Antarctic marine food web
Carnivorous
plankton
Copepods
Krill
Phyto-
plankton 59

60. Species with a Large Impact
Certain species have a very large impact on community structure
Such species are highly abundant or play a pivotal role in community dynamics 60

61. Dominant species are those that are most abundant or have the highest biomass
One hypothesis suggests that dominant species are most competitive in exploiting resources
Another hypothesis is that they are most successful at avoiding predators and disease 61

62. Keystone species exert strong control on a community by their ecological roles, or niches
In contrast to dominant species, they are not necessarily abundant in a community
Field studies of sea stars illustrate their role as a keystone species in intertidal communities 62

63. Number of species present
Figure 41.15
Experiment
Results 20 15
With Pisaster (control)
Number of species
present
Figure Inquiry: Is Pisaster ochraceus a keystone predator? 10
Without Pisaster (experimental) 5
1963’64
’65
’66
’67
’68
’69
’70
’71
’72
’73
Year 63

64. Figure 41.15a
Experiment
Figure 41.15a Inquiry: Is Pisaster ochraceus a keystone predator? (part 1: experiment) 64

65. Number of species present
Figure 41.15b
Results 20 15
With Pisaster (control)
Number of species
present 10
Without Pisaster (experimental) 5
Figure 41.15b Inquiry: Is Pisaster ochraceus a keystone predator? (part 2: results)
1963’64
’65
’66
’67
’68
’69
’70
’71
’72
’73
Year 65

66. Ecosystem engineers (or “foundation species”) cause physical changes in the environment that affect community structure
For example, beaver dams can transform landscapes on a very large scale 66

67. Figure 41.16
Figure Beavers as ecosystem engineers 67

68. Bottom-Up and Top-Down Controls
The bottom-up model of community organization proposes a unidirectional influence from lower to higher trophic levels
In this case, the presence or absence of mineral nutrients determines community structure, including the abundance of primary producers 68

69. The bottom-up model can be represented by the equation
where
N  mineral nutrients
V  plants
H  herbivores
P  predators
N V H P 69

70. The top-down model, also called the trophic cascade model, proposes that control comes from the trophic level above
In this case, predators control herbivores, which in turn control primary producers
N V H P 70

71. Biomanipulation is used to improve water quality in polluted lakes
In a Finnish lake, blooms of cyanobacteria (primary producers) occurred when zooplankton (primary consumers) were eaten by large populations of roach fish (secondary consumers)
Removal of roach fish and addition of pike perch (tertiary consumers) controlled roach populations, allowing zooplankton populations to increase and ending cyanobacterial blooms 71

72. Polluted State Restored State Fish Abundant Rare Zooplankton Rare
Figure 41.UN02
Polluted State
Restored State
Fish
Abundant
Rare
Zooplankton
Rare
Abundant
Figure 41.UN02 In-text figure, biomanipulation, p. 855
Algae
Abundant
Rare 72

73. Concept 41.3: Disturbance influences species diversity and composition
Decades ago, most ecologists favored the view that communities are in a state of equilibrium
This view was supported by F. E. Clements, who suggested that species in a climax community function as an integrated unit 73

74. Other ecologists, including A. G. Tansley and H. A
Other ecologists, including A. G. Tansley and H. A. Gleason, challenged whether communities were at equilibrium
Recent evidence of change has led to a nonequilibrium model, which describes communities as constantly changing after being buffeted by disturbances
A disturbance is an event that changes a community, removes organisms from it, and alters resource availability 74

75. Characterizing Disturbance
Fire is a significant disturbance in most terrestrial ecosystems
A high level of disturbance is the result of a high intensity and high frequency of disturbance
Low disturbance levels result from either low intensity or low frequency of disturbance 75

76. High levels of disturbance exclude many slow-growing species
The intermediate disturbance hypothesis suggests that moderate levels of disturbance can foster greater diversity than either high or low levels of disturbance
High levels of disturbance exclude many slow-growing species
Low levels of disturbance allow dominant species to exclude less competitive species 76

77. In a New Zealand study, the richness of invertebrate taxa was highest in streams with an intermediate intensity of flooding 77

78. Index of disturbance intensity (log scale)
Figure 41.17 35 30 25
Number of taxa 20 15
Figure Testing the intermediate disturbance hypothesis 10
0.9
1.0
1.1
1.2
1.3
1.4
1.5
1.6
1.7
1.8
1.9
2.0
Index of disturbance intensity (log scale) 78

79. The Yellowstone forest is an example of a nonequilibrium community
The large-scale fire in Yellowstone National Park in 1988 demonstrated that communities can often respond very rapidly to a massive disturbance
The Yellowstone forest is an example of a nonequilibrium community 79

80. (a) Soon after fire (b) One year after fire Figure 41.18
Figure Recovery following a large-scale disturbance
(a) Soon after fire
(b) One year after fire 80

81. (a) Soon after fire Figure 41.18a
Figure 41.18a Recovery following a large-scale disturbance (part 1: soon after)
(a) Soon after fire 81

82. (b) One year after fire Figure 41.18b
Figure 41.18b Recovery following a large-scale disturbance (part 2: one year after)
(b) One year after fire 82

83. Ecological Succession
Ecological succession is the sequence of community and ecosystem changes after a disturbance
Primary succession occurs where no soil exists when succession begins
Secondary succession begins in an area where soil remains after a disturbance 83

84. Early-arriving species and later-arriving species may be linked in one of three processes
Early arrivals may facilitate the appearance of later species by making the environment favorable
Early species may inhibit the establishment of later species
Later species may tolerate conditions created by early species, but are neither helped nor hindered by them 84

85. Retreating glaciers provide a valuable field research opportunity for observing succession
Succession on the moraines in Glacier Bay, Alaska, follows a predictable pattern of change in vegetation and soil characteristics
The exposed moraine is colonized by pioneering plants, including liverworts, mosses, fireweed, Dryas, and willows 85

86. Glacier Bay Alaska 0 5 10 15 Kilometers Figure 41.19-1
Figure Glacial retreat and primary succession at Glacier Bay, Alaska (step 1)
Kilometers 86

87. 1941 1 Pioneer stage Glacier Bay Alaska 0 5 10 15 Kilometers
Figure
1941 1
Pioneer stage
Glacier
Bay
Alaska
Figure Glacial retreat and primary succession at Glacier Bay, Alaska (step 2)
Kilometers 87

88. 1941 1907 1 Pioneer stage 2 Dryas stage Glacier Bay Alaska 0 5 10 15
Figure
1941
1907 1
Pioneer stage 2
Dryas stage
Glacier
Bay
Alaska
Figure Glacial retreat and primary succession at Glacier Bay, Alaska (step 3)
Kilometers 88

89. 1941 1907 1 Pioneer stage 1860 2 Dryas stage Glacier Bay Alaska 3
Figure
1941
1907 1
Pioneer stage
1860 2
Dryas stage
Glacier
Bay
Alaska
Figure Glacial retreat and primary succession at Glacier Bay, Alaska (step 4)
Kilometers 3
Alder stage 89

90. 1941 1907 1 Pioneer stage 1860 2 Dryas stage Glacier Bay Alaska 1760 4
Figure
1941
1907 1
Pioneer stage
1860 2
Dryas stage
Glacier
Bay
Alaska
Figure Glacial retreat and primary succession at Glacier Bay, Alaska (step 5)
1760
Kilometers 4
Spruce stage 3
Alder stage 90

91. Figure 41.19a
Figure 41.19a Glacial retreat and primary succession at Glacier Bay, Alaska (part 1: pioneer) 1
Pioneer stage 91

92. After about three decades, Dryas dominates the plant community92

93. Figure 41.19b
Figure 41.19b Glacial retreat and primary succession at Glacier Bay, Alaska (part 2: Dryas) 2
Dryas stage 93

94. A few decades later, alder invades and forms dense thickets94

95. Figure 41.19c
Figure 41.19c Glacial retreat and primary succession at Glacier Bay, Alaska (part 3: alder) 3
Alder stage 95

96. In the next two centuries, alder are overgrown by Sitka spruce, western hemlock, and mountain hemlock 96

97. Figure 41.19d
Figure 41.19d Glacial retreat and primary succession at Glacier Bay, Alaska (part 4: spruce) 4
Spruce stage 97

98. Succession is the result of changes induced by the vegetation itself
On the glacial moraines, vegetation increases soil nitrogen content, facilitating colonization by later plant species 98

99. Human Disturbance
Humans have the greatest impact on biological communities worldwide
Human disturbance to communities usually reduces species diversity
Trawling is a major human disturbance in marine ecosystems 99

100. Figure 41.20
Figure Disturbance of the ocean floor by trawling
100

101. Figure 41.20a
Figure 41.20a Disturbance of the ocean floor by trawling (part 1: before)
101

102. Figure 41.20b
Figure 41.20b Disturbance of the ocean floor by trawling (part 2: after)
102

103. Concept 41.4: Biogeographic factors affect community diversity
Latitude and area are two key factors that affect a community’s species diversity
103

104. Latitudinal Gradients
Species richness is especially great in the tropics and generally declines along an equatorial-polar gradient
Two key factors in equatorial-polar gradients of species richness are probably evolutionary history and climate
104

105. Temperate and polar communities have started over repeatedly following glaciations
The greater age of tropical environments may account for their greater species richness
In the tropics, the growing season is longer, so biological time runs faster
105

106. Climate is likely the primary cause of the latitudinal gradient in biodiversity
Two main climatic factors correlated with biodiversity are sunlight and precipitation
They can be considered together by measuring a community’s rate of evapotranspiration
Evapotranspiration is evaporation of water from soil plus transpiration of water from plants
106

107. Vertebrate species richness
Figure 41.21
200
100
Vertebrate species richness
(log scale) 50
Figure Energy, water, and species richness 10
500
1,000
1,500
2,000
Potential evapotranspiration (mm/yr)
107

108. Area Effects
The species-area curve quantifies the idea that, all other factors being equal, a larger geographic area has more species
108

109. Species richness on islands depends on island size, distance from the mainland, immigration, and extinction
The equilibrium model of island biogeography maintains that species richness on an ecological island levels off at a dynamic equilibrium point
109

110. Studies of species richness on the Galápagos Islands support the prediction that species richness increases with island size
110

111. Area of island (hectares)
Figure 41.22
Results
400
200
100
Number of plant species (log scale) 50 25 10
Figure Inquiry: How does species richness relate to area? 5 10
100
103
104
105
106
Area of island (hectares)
(log scale)
111

112. Concept 41.5: Pathogens alter community structure locally and globally
Ecological communities are universally affected by pathogens, disease-causing organisms and viruses
112

113. Effects on Community Structure
Pathogens can have dramatic effects on community structure when they are introduced into new habitats
For example, coral reef communities are being decimated by white-band disease
Sudden oak death has killed millions of oaks that support many bird species
113

114. Community Ecology and Zoonotic Diseases
Zoonotic pathogens have been transferred from other animals to humans
The transfer of pathogens can be direct or through an intermediate species called a vector
Many of today’s emerging human diseases are zoonotic
114

115. Identifying the community of hosts and vectors for a pathogen can help prevent disease
For example, recent studies identified two species of shrew as the primary hosts of the pathogen for Lyme disease
115

116. Figure 41.23
Figure Identifying Lyme disease host species
116

117. Avian flu is a highly contagious virus of birds
Ecologists are studying the potential spread of the virus from Asia to North America through migrating birds
117

118. Figure 41.24
Figure Tracking avian flu
118

119. Community ecology is needed to help study and combat pathogens
Human activities are transporting pathogens around the world at unprecedented rates
Community ecology is needed to help study and combat pathogens
119

120. Figure 41.UN01a
Figure 41.UN01a Skills exercise: using bar graphs and scatter plots to present and interpret data (part 1)
120

121. Figure 41.UN01b
Figure 41.UN01b Skills exercise: using bar graphs and scatter plots to present and interpret data (part 2)
121

122. Figure 41.UN03
Figure 41.UN03 Summary of key concepts: species interactions
122