Background to Foraging

C.S. Buzz Holling

aka "the man"
Functional response was developed
based on a 1959 paper

These eating These

Sometimes called the disk equation cause this is
how he originally developed the model
Holling Processes of Predation

– Search
– Encounter
– Pursuit
– Capture
– Handling
 Holling’s Disk Equation

Rate of Energy Gained = (λe – s)/(1 + λh)

λ = rate of encounter with diet item
e = energy gained per encounter
s = cost of search per unit time
h = average handling time

Energy gained per time invested per item

 Functional Response

Energy gained per time invested per item

 Functional Response

Type I passive predators (spiders)

Type II most typical: predator saturation

(Daphnia).

Type III Learning – occurs in predators which

increase their search activity with increasing prey
density (bass/goby)
 Numerical Response
Predators increase in abundance as prey density
increases.

Two Potential Mechanisms

1. Increased rate of predator reproduction

when prey are abundant

2. Attraction of predators to prey aggregations

("aggregational response")
 Diets
Goals – Explained in lecture, applied in lab
• Why are diets important?
• How do we get diet data?
• What do we do with diet data?
 Why are diets important?
What information do they tell us?
What questions do they answer?
• Fish’s view
– Source of energy (bioen)
• Managers view
– Stocking
• Aquaculture
– Nutrients
• As an indicator of environmental change
– Nutrient loading, change in pop density
 How do we get diets?
• Fish collection:
– what should we be cautious of? Sources of
error/ bias due to methods selection?
• Regurgitation
• Post capture digestion
• Alteration in behavior due to capture
– Traps
• Did you use bait?
 Sampling strategies
Things to consider/keep in mind
• Diel effects
• Seasonal effects
– Both of the above affect predator and prey
behavior
• Fish size
– Ontogenetic shifts? What age/size class is
your diet representing?
 Sampling strategies cont…
Do stomach contents accurately depict the fish’s diet?

• Digestion rates
– Too fast - protozoans
• Under representation in diets
• Could be determined by watching behavior in
a tank
– Too slow
• Over representation in diets
• Could be solved by determining gut passage
times
 Removal of gut contents
• Puking
• Dissection
 Identification of Diet
Components
• Crushed/digested organism
– Find characteristic structure for each
organism
• Level of identification depends on the
research question
– Higher resolution=greater time investment
Quantitative diet description
• Frequency of Occurrence
• Percent composition by number
• Percent composition by weight
 Frequency occurrence
• What proportion of the diets contained one or
more of a given food type

• Describes presence absence

– Example: 18/22 bluegill contain chironomid,
frequency of occurrence = .82 or 82%
• lower frequency occurrence = selective or
opportunistic feeding
 Frequency of occurrence
Drawbacks
• High frequency of occurrence may not
mean this diet item is of nutritional
importance, only that it is consumed
with some regularity

• Example: benthic fish and algae

 Percent Composition by
number
• Number of food items in a group relative to
total number of diet items consumed (%)

• Potential for fragmentation, count a

characteristic part of prey item
– Dragonflies have four wings, so 4 wings = 1
dragonfly

• Percent composition by number + estimates of

feeding rates = effects of predators on prey
Percent Composition by Weight
• Weight of each type as a percentage
of the total weight of the diet
– Wet and dry weights can be used
• Dry are more precise and offer more
information about nutritional value

• Remember some component of the

diets have already been digested!!!!

• begins to identify food importance in

fish nutrition
Further Analysis and Interpretation
• Selectivity indices: Do fish feed at random
or preferentially?

• Diet overlap indices

 Laboratory

• Groups of two/three
• “work up” 1-2 diets
– Fill out data sheet including weight of
each diet item
• Combine data with other groups
• Analyze the diets
– Freq occ, % comp by number, % comp by
weight
Amphipods
Isopod