Award Abstract # 1061475
Effects of Predator Diversity on the Strength of Trophic Cascades in an Oceanic Benthic Ecosystem

NSF Org: OCE
Division Of Ocean Sciences
Recipient: BROWN UNIVERSITY
Initial Amendment Date: March 8, 2011
Latest Amendment Date: March 8, 2011
Award Number: 1061475
Award Instrument: Standard Grant
Program Manager: David Garrison
OCE
 Division Of Ocean Sciences
GEO
 Directorate For Geosciences
Start Date: March 15, 2011
End Date: February 28, 2015 (Estimated)
Total Intended Award Amount: $628,896.00
Total Awarded Amount to Date: $628,896.00
Funds Obligated to Date: FY 2011 = $628,896.00
History of Investigator:
  • Jon Witman (Principal Investigator)
    jon_witman@brown.edu
Recipient Sponsored Research Office: Brown University
1 PROSPECT ST
PROVIDENCE
RI  US  02912-9100
(401)863-2777
Sponsor Congressional District: 01
Primary Place of Performance: Brown University
1 PROSPECT ST
PROVIDENCE
RI  US  02912-9100
Primary Place of Performance
Congressional District:
01
Unique Entity Identifier (UEI): E3FDXZ6TBHW3
Parent UEI:
NSF Program(s): BIOLOGICAL OCEANOGRAPHY
Primary Program Source: 01001112DB NSF RESEARCH & RELATED ACTIVIT
Program Reference Code(s): 9169, EGCH, OTHR
Program Element Code(s): 165000
Award Agency Code: 4900
Fund Agency Code: 4900
Assistance Listing Number(s): 47.050

ABSTRACT

Intellectual Merit.
Predator diversity has decreased dramatically in the world's oceans due to overfishing, anthropogenic habitat destruction and possibly climate change. Yet, still unknown for most ecosystems is the importance of predator diversity and abundance for ecosystem functioning. One of the most pervasive community-wide consequences of top predators is the Trophic Cascade (TC), where herbivores are suppressed, which releases plants from consumption, thus increasing plant productivity. Recent studies have shown that the diversity of predators may reduce, increase, or have no effect on the strength of trophic cascades. The small number, to date, of experimental tests of predator diversity effects on cascade strength precludes broad generalizations vital to the development of predictive theory. Such research is limited by the lack of experimental realism due to the small number of predator species that can be manipulated in simplified mesocosms. Without more realistic species numbers, it is impossible to extrapolate results to natural ecosystems that experience losses of predator diversity.
To meet these challenges, and to better understand the consequences of present and changing levels of predator diversity in marine ecosystems, a series of experimental manipulations will be conducted on natural levels of predator diversity and their herbivorous sea urchin prey. The hypotheses test the ultimate effects on benthic algae, as a measure of cascade strength in oceanic benthic ecosystems of the Galapagos Marine Reserve (GMR). Because of years of protection from industrial fishing as a UNESCO World Heritage Site, and of local conservation protection as the GMR, there are diverse guilds of higher trophic level predators, such as large fish and sharks. Likewise, there is high diversity of intermediate-level fish and invertebrates that prey on sea urchins, creating an unusual opportunity for testing and developing predator diversity and Biodiversity Ecosystem Functioning theory. The overarching questions addressed in this project are: How do naturally occurring large ranges of oceanic predator diversity influence the strength of trophic cascades? and How does environmental variation and conservation protection influence these processes? The first question will be addressed in experiments manipulating both horizontal (within trophic level; urchin herbivores) and vertical (across trophic level; predators) consumer diversity and in another experiment manipulating the diversity of predatory fish and invertebrates guilds. The experiments employ open fenced treatments containing urchins but allowing access by fish and invertebrate predators of the urchins. To record natural levels of fish and invertebrate predator richness encountering the treatments, consuming the urchins and interacting with each other, the entire experimental layout will be video-recorded for up to several weeks at a time. The time-lapse cameras/lighting system is capable of day and night imaging without affecting predator behavior. A simplified manipulation to measure the influence of predator diversity on cascade strength will be replicated and video-recorded at 16 sites -- representing different levels of upwelling and conservation protection -- to place the mechanistic understanding gleaned from detailed experiments at local sites into a broader (mesoscale) context.

Broader Impacts.
Several major impacts of the proposed research on education are expected beyond the implications for conservation and ecosystem-based management. For instance, the program will provide hands-on training for one graduate student and two undergraduates in experimental ecology, sub-tidal community ecology, statistics, marine reserve science, scientific writing and oral presentation skills. The PI is committed to educational and public outreach activities beyond the typical use of the information gained on predators and food webs by using it to enhance lectures in his large undergraduate Ecology class at Brown University. One of these outreach activities is to help train the next generation of teacher leaders by collaborating with the Director of the Brown University's Masters in Teaching (MAT) program to offer a science internship for one MAT student per year. The general information on food webs, ecology of charismatic Galapagos megafuana and images of marine predators in action obtained from the time-lapse video would be shared with the MAT interns to inspire their lesson plans for elementary and high school students. In summary, the broader educational impacts of the proposed research will be to enhance the workforce of globally trained doctoral students in the life sciences, help prepare undergraduate students for graduate school in marine ecology, augment the training of the next generation of K-12 teachers and enhance the understanding of one of the most pressing environmental issues -- the ecosystem consequences of over-harvesting top predators and concomitant loss of predator diversity in the sea.

PUBLICATIONS PRODUCED AS A RESULT OF THIS RESEARCH

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Altieri, A.H. and J.D. Witman "Modular mobile foundation species as reservoirs of biodiversity" Ecosphere , v.5 , 2014 , p.124 http://dx.doi.or
Dee, L. E, J.D. Witman and M. Brandt. "Refugia and top-down control of the pencil urchin Eucidaris galapagensis in the Galapagos Marine Reserve" Journal of Experimental Marine Biology and Ecology , v.416 , 2012 , p.135

PROJECT OUTCOMES REPORT

Disclaimer

This Project Outcomes Report for the General Public is displayed verbatim as submitted by the Principal Investigator (PI) for this award. Any opinions, findings, and conclusions or recommendations expressed in this Report are those of the PI and do not necessarily reflect the views of the National Science Foundation; NSF has not approved or endorsed its content.

Sound management of the oceans requires understanding the roles that key species and biodiversity plays in ecosystem functioning. Since we cannot investigate all marine ecosystems in enough detail to extract this information, we rely on theories to predict when and where strong ecological interactions from key species will control ecosystem functioning, such as primary production of algae on the sea floor. One of these theories predicts that strong controlling interactions from top predators will be less common in diverse food chains (webs) containing large numbers of species than in food webs with few species. The basis for this is the notion that the many interactions among species that arise in diverse food webs, such as competition for food or aggressive behaviors, could reduce the ability of top predators to consume species on lower levels. However, realistic field experiments are seldom applied to test this theory in diverse marine ecosystems, such as the tropics.  

Consequently, we used the diverse food webs of the Galapagos Islands to test the theory, predicting that by feeding on herbivorous sea urchins, predatory fish could indirectly increase the abundance of primary producers (bottom dwelling algae) that the sea urchins typically control. To mimic conditions of the natural subtidal ecosystem as much as possible, we developed a novel “open” experimental design where unconfined predatory fish could move in and out of the treatments to attack the sea urchins, and interact with each other and top predators (sharks, sea lions) while foraging.  Since the week- long experiments were monitored by high frequency (1 second) time-lapse underwater photography, it was possible to identify both feeding and non-feeding (behavioral) effects of predators, as well as to document the species involved in predation on sea urchins.

The results were striking; pencil urchins (Eucidaris galapgensis) were eaten by blunthead triggerfish (Pseudobalistes naufragium) within 21 hours, releasing benthic algae from urchin grazing, which establishes a strong ecological interaction called a trophic cascade, as triggerfish reduce sea urchin abundance, which indirectly increases algae. Finescale trigger fish (Balistes polylepis) were also identified as major predators of pencil urchins. When unchecked by predators in predator exclusion cages, the pencil urchins removed 10% of algae in a week. Spanish hogfish, previously considered a top predator of sea urchins, were instead revealed as “hitchhikers” on the triggerfish, often circling them closely and reducing their feeding efficiency on sea urchins. Sea lions swam rapidly over the experiments in search of prey, disturbing feeding triggerfish and causing them to drop urchin prey on occasion. The high frequency database on species interactions enabled us to construct a mathematical model of triggerfish foraging on sea urchins. The model revealed that the presence of both top predators (sharks, sea lions) and hogfish reduce triggerfish feeding rates. Surprisingly, these results were highly contingent on the species of sea urchins involved. For example, predators (triggerfish, hogfish) virtually ignored green sea urchins (Lytechinus semituberculatus) when they were used in the experiments instead of pencil urchins. The green urchins consumed 3 x more algae than the pencil urchins. Counter to theory, our results indicate that that strong predatory interactions do occur in diverse food webs, and can cascade down to transform the abundance of bottom dwelling algae, and indicator of ecosystem functioning. The Galapagos marine food web can be characterized as one in which strong predatory interactions are embedded in a diffuse web of behavioral interactions. Biodiversity matters, as certain species such as two species of triggerfish and sea urchins play unique ...

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