Summary
Foraging is one of the most important endeavors undertaken by animals, and it has been studied intensively from both mechanistic-empirical and optimal foraging perspectives. Planktivorous fish make excellent study organisms for foraging studies because they feed frequently and in a relatively simple environment. Most optimal foraging studies of planktivorous fish have focused, either on diet choice or habitat selection and have assumed that these animals used a cruise search foraging strategy. We have recently recognized that white crappie do not use a cruise search strategy (swimming continuously and searching constantly) while foraging on zooplankton but move in a stop and go pattern, searching only while paused. We have termed thissaltatory search. Many other animals move in a stop and go pattern while foraging, but none have been shown to search only while paused. Not only do white crappie search in a saltatory manner but the components of the search cycle change when feeding on prey of different size. When feeding on large prey these fish move further and faster after an unsuccessful search than when feeding on small prey. The fish also pause for a shorter period to search when feeding on large prey. To evaluate the efficiency of these alterations in the search cycle, a net energy gain simulation model was developed. The model computes the likelihood of locating 1 or 2 different size classes of zooplankton prey as a function of the volume of water scanned. The volume of new water searched is dependent upon the dimensions of the search volume and the length of the run. Energy costs for each component of the search cycle, and energy gained from the different sized prey, were assessed. The model predicts that short runs produce maximum net energy gains when crappie feed on small prey but predicts net energy gains will be maximized with longer runs when crappie feed on large prey or a mixed assemblage of large and small prey. There is an optimal run length due to high energy costs of unsuccessful search when runs are short and reveal little new water, and high energy costs of long runs when runs are lengthy. The model predicts that if the greater search times observed when crappie feed on small prey are assessed when they feed on a mixed diet of small and large prey, net energy gained is less than if small prey are deleted from the diet. We believe the model has considerable generality. Many animals are observed to move in a saltatory manner while foraging and some are thought to search only while stationary. Some birds and lizards are, known to modify the search cycle in a manner similar to white crappie.
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Abbreviations
- SST (sec):
-
Successful search time — the average time stationary prior to a pursuit
- USST (sec):
-
Unsuccessful search time — the average time stationary prior to a run
- PT (sec):
-
Pursuit time-PL/SS — the time to pursue prey at a given distance away. It is calculated by dividing the pursuit distance by swim speed
- RT (sec):
-
Run time-RL/SS — the time to complete a run of a given length. It is calculated by dividing the run length, by swim speed
- PL (cm):
-
Pursuit length-distance moved to attack prey
- RL (cm):
-
Run length-distance moved between consecutive searches
- SS (cm/sec):
-
Swim speed — the speed of movement during a pursuit or run
- LS (l):
-
Location space — the area or volume within which prey are located. In the case of white crappie the search space is shaped like a pie wedge with the fish positioned at the apex of the wedge
- LA (o):
-
Location angle—the angle of the wedge-shaped search space
- LH (cm):
-
Location height—the height of the wedge-shaped search space
- LD (cm):
-
Location distance—the length of long axis of the wedge-shaped search space.
- RND:
-
Random number-random number generated through BASICA
- SV (l):
-
Search volume—the volume of water actually searched after one run of given length
- SVMAX (l):
-
Maximum search volume—the greatest search volume that can be based upon LA, LH, LD and unaffected by the previous search
- SVR (l):
-
Search volume researched—that volume of SVMAX that is researched where RL<LD (see Appendix A)
- SVo :
-
Search volume unsearched—that volume of SVMAX not previously searched
- AD (#/1):
-
Absolute density—the density of zooplankton prey in numbers per liter
- VD (#):
-
Visual density—the number of zooplankton prey in the search volume
- LP (%):
-
Location probability—the probability that one or more prey are in the search volume
- NEG (cal/sec):
-
Net energy gain-total calories ingested, less total calories used, divided by total time.
- E e (cal):
-
Energy expended on the search cycle
- E i (cal):
-
Energy intake
- e p (cal):
-
Energy content of a given individual prey
- P i :
-
Total number of prey ingested
- e r (cal):
-
Energy expended while searching
- e s (cal):
-
Energy expended while swimming
- T t (sec):
-
Total time-time expended to eat a given number of prey
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John O'Brien, W., Evans, B.I. & Browman, H.I. Flexible search tactics and efficient foraging in saltatory searching animals. Oecologia 80, 100–110 (1989). https://doi.org/10.1007/BF00789938
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DOI: https://doi.org/10.1007/BF00789938