Week 6: Predation and predatory behavior: Lecture summaryhomepages.wmich.edu › ... › Lectures...
Transcript of Week 6: Predation and predatory behavior: Lecture summaryhomepages.wmich.edu › ... › Lectures...
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BIOS 6150: Ecology - Dr. S. Malcolm. Week 6: Predation and predatory behavior Slide - 1
BIOS 6150: Ecology Dr. Stephen Malcolm, Department of Biological Sciences
• Week 6: Predation and predatory behavior:
• Lecture summary: • Nature of predation. • Diet breadth & choice. • Optimal foraging. • Functional responses • Mutual interference. • Aggregative response. • Marginal value theorem.
J. Kobalenko. 1997. Forest Cats Of North America. Firefly Books
http://www.americazoo.com/goto/index/mammals/134.htm
Snowshoe hare and lynx.
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BIOS 6150: Ecology - Dr. S. Malcolm. Week 6: Predation and predatory behavior Slide - 2
2. Predation:
• Is a description of the interaction between predator foraging behavior and prey defense.
• This includes both behavior and population dynamics.
• Fig. 20.1 from Malcolm (1992) in “Natural Enemies” edited by M.J. Crawley (Blackwell).
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BIOS 6150: Ecology - Dr. S. Malcolm. Week 6: Predation and predatory behavior Slide - 3
3. Predation literature:
• Very strong emphasis on predator foraging behavior and prey-predator dynamics.
• Defense is mostly relegated to the realms of natural history description.
• Predator foraging behavior is a description of:
• where they feed. • what they feed on. • how they are influenced by other predators. • how they are influenced by prey density.
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BIOS 6150: Ecology - Dr. S. Malcolm. Week 6: Predation and predatory behavior Slide - 4
4. Diet composition and food preference:
• Predators can be: • Monophagous:
• single prey type and have a large impact on prey population dynamics
• Oligophagous: • few prey types, or,
• Polyphagous: • many prey types and probably have little impact on
the population dynamics of any one species.
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BIOS 6150: Ecology - Dr. S. Malcolm. Week 6: Predation and predatory behavior Slide - 5
5. Prey choice:
• Within different diet breadths predators choose more profitable prey preferentially (Table 9.1) and so food can also be assessed by predators as either: • Ranked food resources that are most valuable or
“perfectly substitutable” • see Figs. 9.14 and 9.15, or,
• Balanced food resources that are integral or “complementary”
• Usually necessary to balance required nutrients that may be absent from high ranked foods.
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BIOS 6150: Ecology - Dr. S. Malcolm. Week 6: Predation and predatory behavior Slide - 6
6. Switching:
• Predators can also “switch” their food preference as in Fig. 9.15: • Perhaps through learned abilities to handle prey
more profitably: • More efficient balance among search, pursuit, and
handling behaviors before consumption: • This may be facilitated by specific “search images”.
• Such changes in diet may also be seasonal or on shorter time scales that may be associated with the induction of physiologies better suited to exploiting the food resource.
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BIOS 6150: Ecology - Dr. S. Malcolm. Week 6: Predation and predatory behavior Slide - 7
7. “Optimal foraging” and diet width:
• Why are real diets "narrower" than potential diets? • If energy maximization is the primary criterion that correlates
well with fitness then optimal foraging theory is useful. • MacArthur & Pianka (1966) initiated the influential
optimal foraging theory approach for the description of the evolutionary ecology of predatory behavior based on:
• Maximization of the net rate of energy intake: • gross energy intake - energetic costs of obtaining that energy.
• Predators incur energy and time costs of: • Searching for prey • Handling prey:
• Includes: detection, pursuit, acceptance, subjugation & consumption
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BIOS 6150: Ecology - Dr. S. Malcolm. Week 6: Predation and predatory behavior Slide - 8
8. Optimal foraging theory:
• The aim is to predict the expected foraging “strategy” under specified conditions (Fig. 9.17):
• Is it a “tactic” or a “strategy”? • Generalist costs:
• Low time search costs but higher costs of handling both unprofitable and profitable prey.
• Specialist costs: • High time costs but lower costs of handling profitable
prey.
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BIOS 6150: Ecology - Dr. S. Malcolm. Week 6: Predation and predatory behavior Slide - 9
9. Diet profitability:
• MacArthur & Pianka argued that a prey item (i) should be included (and diet width expanded) if it is equal to, or more profitable than, the average profitability of the present diet, thus if: • Ei /hi ≥ E/(s + h) • where i is the next most profitable prey item • E = energy content • h = handling time (therefore E/h = profitability) • s = search time
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BIOS 6150: Ecology - Dr. S. Malcolm. Week 6: Predation and predatory behavior Slide - 10
10. Foraging guilds:
• Handling time < search time = generalists: • e.g. foliage gleaning bird guild:
• A guild is a group of individuals that exploit the same resource in the same way (after Root).
• Handling time > search time = specialists: • e.g. lions living near prey:
• Note: handling time includes pursuit time! • See text – these don’t make sense to me, despite
discussing this with Mike Begon!
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BIOS 6150: Ecology - Dr. S. Malcolm. Week 6: Predation and predatory behavior Slide - 11
11. Foraging constraints:
• Abiotic and Biotic: • More dimensions of “realized” niches! • Biotic:
• see Figures 9.18 and 9.19 • Abiotic:
• e.g. the interaction between temperature and oxygen constrains Notonecta foraging for submerged or floating prey according to dissolved oxygen levels (see Figs 2 & 4 from Cockrell (1984) Journal of Animal Ecology 53(2): 519-532.)
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BIOS 6150: Ecology - Dr. S. Malcolm. Week 6: Predation and predatory behavior Slide - 12
12. Functional Responses:
• Describe the relationship between an individual predator’s consumption rate and prey density:
• After Solomon (1949) but developed by Holling (1959).
• 3 kinds recognized by Holling: • Type 1 (linear). • Type 2 (asymptotic). • Type 3 (sigmoid).
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BIOS 6150: Ecology - Dr. S. Malcolm. Week 6: Predation and predatory behavior Slide - 13
13. Type 2 Functional Response:
• Type 2 functional response is most frequently observed (see Figs 10.9 & 9.7):
• Handling t stays constant but search t decreases with increasing prey density.
• Thus total handling time increases. • Handling time Th determines the height of the
curve plateau. • Attack rate a determines rate that plateau is
reached.
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BIOS 6150: Ecology - Dr. S. Malcolm. Week 6: Predation and predatory behavior Slide - 14
14. Types 1 & 3 Functional Responses:
• Type 1 functional response (slope = a) as in filter-feeding Daphnia (Fig. 10.8).
• Type 3 functional responses as in vertebrate predators capable of learning (Fig. 10.10) and showing “switching” behaviors:
• Increased attack rate and increased searching time or decreased handling time.
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BIOS 6150: Ecology - Dr. S. Malcolm. Week 6: Predation and predatory behavior Slide - 15
15. Holling's “disc” equation:
• The functional response relationship is described by Holling's “disc” equation in which: • Prey eaten, Pe = a TsN
• where Ts is the period of searching time during which Pe prey are eaten, and
• N = prey density • but, Ts = T - ThPe • where T = total time
• and so, Pe = a (T - ThPe)N • Y = a(T - bY)X, in Tostowaryk (1972)
• or, rearranging, Pe = aNT/1 + aThN
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BIOS 6150: Ecology - Dr. S. Malcolm. Week 6: Predation and predatory behavior Slide - 16
16. Michaelis-Menten-Holling equation:
• Holling’s disc equation is the same as the continuous form: • b(N) = mN/(w + N),
• where m is the maximum predator attack rate, • b = rate of change of N due to the interaction, and, • w is prey density where attack rate is half saturated.
• This is also the same as the Michaelis-Menten equation that describes the kinetics of enzyme catalyzed reactions:
• vo = VmaxS/Km + S, where, • Km = w, S = N, Vmax = m
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BIOS 6150: Ecology - Dr. S. Malcolm. Week 6: Predation and predatory behavior Slide - 17
17. Effects of functional responses on population dynamics:
• 1) Decelerating consumption rate results in destabilization because it is inversely density dependent:
• All 3 functional responses at high density.
• 2) Accelerating consumption rate results in density-dependent stabilization:
• Type 3 functional response at low density.
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BIOS 6150: Ecology - Dr. S. Malcolm. Week 6: Predation and predatory behavior Slide - 18
18. Predator density - mutual interference:
• Mutual interference - effects of competition: • Effects of territoriality, or resource defense, or direct
interference competition, or indirect exploitative competition, can all increase with increased predator density:
• Figure 9.10 shows density dependent changes when searching efficiency a (=attack rate) is plotted against predator density:
• The slope of this relationship m is the coefficient of interference. This negative slope tends to stabilize predator-prey dynamics.
• In contrast to social facilitation at low predator density: • e.g. foraging dolphins.
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BIOS 6150: Ecology - Dr. S. Malcolm. Week 6: Predation and predatory behavior Slide - 19
19. Prey density - aggregative responses by predators to prey “patches”:
• Aggregative response • Predators spend more time in high density prey
patches than low density patches (where spatial distribution varies) (Fig. 9.11).
• Combined functional and aggregative responses (Fig. 9.22).
• Impact on population dynamics: • Partial prey refuges at both high and low prey
density: • Lowered probability of attack tends to stabilize
predator-prey population dynamics (Fig. 5.19)
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BIOS 6150: Ecology - Dr. S. Malcolm. Week 6: Predation and predatory behavior Slide - 20
20. The “ideal free distribution” of predators and prey:
• Aggregation + interference may combine to generate: • An ideal free distribution (Fig. 9.27), or, • Patchiness in time and space can generate
stability: • As in Huffaker's orange+mites experiment:
• Through equal (“ideal”) patch profitabilities after (“free”) redistribution.
• Interaction between competition and predation!
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BIOS 6150: Ecology - Dr. S. Malcolm. Week 6: Predation and predatory behavior Slide - 21
21. The Marginal Value Theorem:
• Based on the work of Charnov (1976) and Parker & Stuart (1976) to predict the behavior of an optimal forager in patches of food of different profitabilities:
• The forager should maximize its overall intake of a resource (energy) per time spent foraging in habitats with food distributed patchily: • How long should the forager spend in patches of
varying profitability? • Fig. 9.22 illustrates the model and Fig. 9.23 is a test
of the model (Cowie).
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Table 9.1 (3rd ed.):
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Figure 9.14: Selection of the most profitable prey by crabs and wagtails
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Figure 9.15: Preference (a & c), switching (b) and switching + learning (d & e)
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Figure 9.17: Predictions and observations of diet choice in great tits and bluegill sunfish
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Figure 9.18: Seasonal variation in predicted and observed habitat profitabilities for bluegill sunfish.
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BIOS 6150: Ecology - Dr. S. Malcolm. Week 6: Predation and predatory behavior Slide - 27
Figure 9.19: Effect of largemouth bass on sunfish feeding distribution.
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BIOS 6150: Ecology - Dr. S. Malcolm. Week 6: Predation and predatory behavior Slide - 28
Figure 2: Effect of water temperature on time spent submerged by foraging Notonecta (Cockrell, 1984):
Journal of Animal Ecology 53(2): 519-532.)
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Figure 4:
Cockrell, B.J. 1984. Journal of Animal Ecology 53(2): 519-532.)
Mean length of time spent submerged by Notonecta and number of attacks on flies at the surface and Asellus on the bottom of water tanks at 3 dissolved oxygen concentrations.
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Figure 10.9: Type 2 functional responses of (a) damselfly nymphs and (b) bank voles.
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Figure 9.7 (3rd ed.): Type 2 functional responses in a parasitoid and effect of experience
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Figure 10.8: Type 1 functional response in Daphnia filter-feeding yeast.
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Figure 10.10:
Type 3 functional response in: (a) shrews & mice, (b,d) flies, (c,e) wasp parasitoid
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Figure 9.10 (3rd ed.): Negative impact of mutual interference increases with forager density
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Figure 9.11 (3rd ed.): Aggregative responses of foragers to host or prey density
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Figure 9.22 (3rd ed.): Interaction between aggregative and functional responses
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Figure 5.19: Effect of tide fluctuations on the distributions of predatory whelks and their barnacle prey.
Begon, Mortimer & Thompson (1996)
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Figure 9.27: Ideal-free distribution in foraging ducks
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BIOS 6150: Ecology - Dr. S. Malcolm. Week 6: Predation and predatory behavior Slide - 39
Figure 9.22:
The marginal value theorem
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BIOS 6150: Ecology - Dr. S. Malcolm. Week 6: Predation and predatory behavior Slide - 40
Figure 9.23: Predicted and observed foraging times spent by great tits in prey patches with different traveling times.