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Altered behavior and cost of manipulation: the acanthocephalan Leptorhyncoides thecatus in its amphipod host Hyalella azteca

Date

2014

Authors

Stone, Charles F., author
Moore, Janice, advisor
Kondratieff, Boris, committee member
Ghalambor, Cameron, committee member
Naug, Dhruba, committee member

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Abstract

Behavioral manipulation occurs when a parasite causes changes in its host's behavior to the parasite's benefit. The parasite benefits from these behavioral changes by increased survival or transmission. It has been hypothesized that such manipulation carries a cost for the parasite because energy allocated to manipulation does not contribute to growth or reproduction. The acanthocephalan parasite Leptorhynchoides thecatus provides a system in which to test this concept. This parasite uses the amphipod Hyalella azteca as an intermediate host and fish as definitive hosts; it has not been previously shown to alter host behavior. This system is advantageous for testing costs of manipulation: the size of the larval cystacanth stage in the intermediate host provides an easily quantified measure of fitness. Larger cystacanths establish in the fish host more frequently than smaller cystacanths. If manipulation is costly, I predict that there should be a negative relationship between the strength of behavioral change and fitness measures (larval size). I compared geotaxis, phototaxis, photophilia, and activity responses of infected and uninfected H. azteca to determine whether L. thecatus modified behavior. I also measured the responses of infected and uninfected amphipods to alarm pheromones and predator kairomones. I then investigated whether these behavioral changes were correlated with larval size. I found that L. thecatus does indeed alter host behavior. Compared to uninfected amphipods, infected amphipods were found higher in the water column, spent greater time in lighted areas, and were more active. There was no difference in phototaxis; both groups of amphipods swam away from a direct light source. Infected amphipods also reduced anti-predator responses to alarm pheromones and predator kairomones. This is the first example of altered alarm pheromones behavior in parasitized amphipods. These findings strongly suggest that L. thecatus increases encounters between its intermediate host and definitive host predators and that the parasite increases its transmission rate through behavioral manipulation. None of these behavioral changes were correlated with a decrease in larval size as predicted by the manipulation cost hypothesis. In fact, larger cystacanths altered geotaxis and photophilia more than smaller cystacanths did. Finally, I compare L. thecatus host use data collected from Atkinson Reservoir, Nebraska, between 2008 and 2011 to published data from 1979-1980. Both data sets show that the Green Sunfish (Lepomis cyanellus) and Pumpkinseed Sunfish (Lepomis gibbosus) are the highest quality hosts for this population. However, the current data suggest a possible shift in secondary hosts from Largemouth Bass (Micropterus salmoides) to Bluegill Sunfish (Lepomis macrochirus). Understanding the cost associated with any trait sheds light on the evolution and maintenance of that trait. This dissertation uses a unique population of L. thecatus to add this parasite to the growing list of those that behaviorally manipulate their hosts, and to demonstrate that, contrary to predictions from the theoretical literature, behavioral manipulation is not necessarily costly.

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Subject

Hyalella azteca
Acanthocephala
Leptorhynchoides thecatus
parasitic manipulation
amphipoda
cost

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