View source for Other Techniques

=Other Techniques=
==Filter Feeding==
[[File:gillrakers.gif|thumb|Gill rakers<ref>Miller, Daniel J., and Robert N. Lea. "Guide to the Coastal Marine Fishes of California." State of California: The Resources Agency Department of Fish and Game Fish Bulletin 157 (1972). Accessed April 19, 2015. http://texts.cdlib.org/view?docId=kt896nb2qd&brand=calisphere&doc.view=entire_text.</ref>]]
[[File:Manta.gif|thumb|Manta Ray Feeding Structures<ref>Martin, R. Aidan. "Manta Ray (Manta Birostris) FAQ." Biology of Sharks and Rays. Accessed April 19, 2015. http://www.elasmo-research.org/education/topics/lh_manta_faq.htm.</ref>]]
[[File:Crossflow.jpg|thumb|Illustration of crossflow filtration<ref>Brainerd, Elizabeth L. "Caught in the Crossflow." Nature 412 (2001): 387-88. Accessed April 19, 2015. http://www.nature.com/nature/journal/v412/n6845/fig_tab/412387a0_F2.html.</ref>]]
Filter feeding is a feeding method observed among some types of fish in Caribbean reefs. Filter feeders filter dissolved and suspended organic matter from the water by forcing water to flow across specialized filtration structures. The most common food source for filter feeding fish is [[plankton]] or other small organisms. In fish, the anatomical structures used to filter out [[plankton]] are most often gill rakers.<ref name="coralscience">Salman, Nadir A., Ghaith J. Al-Mahdawi, and M.A. Hassan. "Gill Rakers Morphometry and Filtering Mechanism in Some Marine Teleosts from Red Sea Coasts of Yemen." Egyptian Journal of Aquatic Research 31 (2005): 286-96.</ref>


Herring are one example of a fish that use filter feeding, although they use it as one of two methods of feeding (the other is particulate-feeding). <ref name="herring">Gibson, R.N., and A. Ezzi. "Effect of Particle Concentration on Filter- and Particulate-feeding in the Herring Clupea Harengus." Marine Biology 88, no. 2 (1985): 109-16.</ref> When filter feeding, herring swim through the water with an open or snapping mouth, allowing water to flow through the oral cavity and out gill openings, using long, thin gill rakers to sift out the small crustaceans and [[plankton]] present in the water column.<ref>Hiatt, Robert W., and Donald W. Strasburg. "Ecological Relationships of the Fish Fauna on Coral Reefs of the Marshall Islands." Ecological Monographs 30, no. 1 (1960): 65-127. Accessed February 20, 2015. http://www.jstor.org/stable/pdf/1942181.pdf?acceptTC=true.
.</ref> The sieve effect of the gill rakers causes food particles to concentrate in the oral cavity, after which they are ingested. <ref name="herring2">Sanderson, S. L., Cheer, A. Y., Goodrich, J. S., Graziano, J. D., & Callan, W. T. (2001). Crossflow filtration in suspension-feeding fishes. Nature, 412(6845), 439-41. doi:http://dx.doi.org/10.1038/35086574</ref>


Another example of filter feeders is the manta ray. 
Manta rays practice a variation called “ram filter-feeding,” in which they swim at high speeds through a concentration of [[plankton]] with their mouth open. At lower concentrations of prey they may continuously filter out [[plankton]] as they swim, while amid higher concentrations of prey they engulf and filter out large concentrations of [[plankton]] at a time.<ref name="ram">Paig-Tran, E. (2012). Filtration at the mega-scale: Exploring the filter morphology and filtration mechanisms in the cartilaginous fishes (Order No. 3552837). Available from ProQuest Dissertations & Theses Full Text; ProQuest Dissertations & Theses Global. (1312511009). Retrieved from http://search.proquest.com/docview/1312511009?accountid=14244</ref> A manta ray expands its flattened body while propelling itself forward through the water with its mouth open, allowing water to flow through the oral cavity and out over five pairs of gill slits that line its throat. It unfurls two horn-like cephalic fins on either side of its mouth to form a funnel to channel more water and [[plankton]] into its oral cavity. Any [[plankton]] larger than a grain of sand is then captured by the feathered gill plates that line its gill slits.<ref name="mantas">"Feeding Frenzy." Manta Trust. January 1, 2015. Accessed April 15, 2015. http://www.mantatrust.org/about-mantas/feeding-frenzy/.</ref>

Manta rays use a variation of filter feeding called “cross-flow filtration” in which water flows parallel to its gill plate surfaces, only deviating slightly to dip across the filter surface and siphon out of the gills. This causes denser food particles with more momentum than the surrounding water medium to travel straight to the back of the mouth and form a concentrated food ball for the manta ray to eat.  <ref name="crossflow">Brainerd, E. L. (2001). Caught in the crossflow. Nature, 412(6845), 387-8. doi:http://dx.doi.org/10.1038/35086666</ref> This filtration method can act as a self-cleaning process that pushes food particles collecting on the filters back to the opening of the esophagus without needing mucus to capture or carry them. This means that manta rays have less filter-clogging problems and can swim and feed for longer periods of time without needing to stop and close their mouths to clear and eat the food off their filters.<ref name="nomucus">Paig-Tran, Misty. "Secrets of Filter-Feeding Sharks and Rays." College of the Environment at the University of Washington. January 1, 2013. http://depts.washington.edu/fhl/enews/winter2013/misty.html.</ref>

Manta rays employ many different strategies while filter feeding. One such strategy is called "barrel rolling", involves the ray rolling itself backwards as they feed on dense patches of [[plankton]]. Another strategy employed while filter feeding is simply bottom feeding, where the manta ray will scoop up [[plankton]] from the seabed. One strategy called "cyclone feeding" is observed only in the Hanifaru Bay in the Maldives, and only occurs about a dozen times each year. In cyclone feeding, the mantas feed in a chain that loops around to form a spiraling column of up to 150 individuals. This spiraling column acts as a cyclone that creates a vortex where filtered water is pulled towards the surface. [[Plankton]] on the outside of the cyclone get pulled in to the cyclone and to the open mouths of the manta rays.<ref name="mantas">"Feeding Frenzy." Manta Trust. January 1, 2015. Accessed April 15, 2015. http://www.mantatrust.org/about-mantas/feeding-frenzy/.</ref>


Whale Sharks are also filter-feeders that use the ram method, swimming around while continuously ramming water and [[plankton]] through their mouths and catching the food with filtering pads covering their throat entrances while letting the water escape through their gills. <ref name="usf">Rutger, Hayley. "USF study examines whale shark eating habits." University of South Florida. October 29, 2010. Accessed April 14, 2015. http://www.cas.usf.edu/news/s/89.</ref> Whale sharks also use suction feeding, where negative pressure is created in their oral cavity which pulls nearby water and [[plankton]] inside when the mouth opens.<ref name="ram">Paig-Tran, E. (2012). Filtration at the mega-scale: Exploring the filter morphology and filtration mechanisms in the cartilaginous fishes (Order No. 3552837). Available from ProQuest Dissertations & Theses Full Text; ProQuest Dissertations & Theses Global. (1312511009). Retrieved from http://search.proquest.com/docview/1312511009?accountid=14244</ref>

==Detritivores==
Detritivores consume dead organic material and return nutrients to the sediment.<ref>Zuhlke, Samantha. "Coral Reef Food Web." National Geographic. January 1, 2015. Accessed April 15, 2015. http://education.nationalgeographic.com/education/media/coral-reef-food-web/?ar_a=1.</ref> Such dead organic material consists of animal and plant remains, biological waste products, and the bacteria and microorganisms associated with biological waste.  <ref name="fishchannel">Scott, Michael. "Marine Fish Feeding Guilds." FishChannel.com. Accessed April 15, 2015. http://www.fishchannel.com/fish-health/saltwater-conditions/marine-fish-feeding-guilds.aspx</ref> At least three reef fish families, surgeonfishes, damselfishes, and blennies, have been identified as feeding heavily on detritus in what is referred to as the “epilithic algal matrix”. <ref name="fishchannel">Scott, Michael. "Marine Fish Feeding Guilds." FishChannel.com. Accessed April 15, 2015. http://www.fishchannel.com/fish-health/saltwater-conditions/marine-fish-feeding-guilds.aspx</ref> This matrix consists of a mixed layer of algae, detritus, invertebrates, and sediment covering dead coral surfaces, which makes up 30-80% of total surface area on most coral reefs. <ref>Kramer, M. J., D. R. Bellwood, and O. Bellwood. "Cryptofauna of the Epilithic Algal Matrix on an Inshore Coral Reef, Great Barrier Reef." Coral Reefs 31 (2012): 1007-015. Accessed April 14, 2015. http://link.springer.com/article/10.1007/s00338-012-0924-x#page-1. 
</ref>


Hard substrate detritivores press their jaw against the substrate and throw their lower jaw upward to brush particulate matter off of rocks, dead coral, and out of turf algae. This feeding technique includes most members of the genus Ctenochaetus.<ref name="fishchannel">Scott, Michael. "Marine Fish Feeding Guilds." FishChannel.com. Accessed April 15, 2015. http://www.fishchannel.com/fish-health/saltwater-conditions/marine-fish-feeding-guilds.aspx</ref> Sediment detritivores eat mouthfuls of sand and associated debris to consume the detritus mixed in with the sediment.<ref name="fishchannel">Scott, Michael. "Marine Fish Feeding Guilds." FishChannel.com. Accessed April 15, 2015. http://www.fishchannel.com/fish-health/saltwater-conditions/marine-fish-feeding-guilds.aspx</ref> Spotted surgeonfish are one such fish that use this technique, and they feed by whisking comb-like teeth over the sandy bottom as they open and close their mouths. <ref>Capuli, Estelita Emily. "Ctenochaetus Strigosus." Fish Base. January 1, 2001. Accessed April 15, 2015. http://www.fishbase.org/summary/6015.</ref> Algal detritivores, such as combtooth blennies, scrape algae off of hard  surfaces  along with associated filamentous algae, diatoms, detritus, and sand. While such feeders do consume some algae, their primary source of nutrition is the detritus particles within the algae.<ref name="tfh">Michael, Scott. "Über Algae Eaters: The Lawnmower Blennies (Full Article)." Tropical Fish Hobbyist Magazine. April 1, 2013. Accessed April 15, 2015. http://www.tfhmagazine.com/details/articles/ber-algae-eaters-the-lawnmower-blennies-full-article.htm.</ref>


Mucus feeders eat only the mucus covering the outer layer of coral tissues, and do so without consuming any underlying live coral tissue or skeleton. <ref name="Rotjan">Rotjan, Randi D. "Impact of Coral Predators on Tropical Reefs." Marine Ecology Progress Series 367 (2008): 73-91.</ref> Corals produce lots of mucus to coat their outer tissue layer that contains energy-rich wax esters, fatty acids, and triglycerides, which provide a valuable source of energy for many fish. <ref name="fishchannel">Scott, Michael. "Marine Fish Feeding Guilds." FishChannel.com. Accessed April 15, 2015. http://www.fishchannel.com/fish-health/saltwater-conditions/marine-fish-feeding-guilds.aspx</ref><ref name="Rotjan">Rotjan and Lewis 2008</ref><ref name="Coffroth">Coffroth, Mary Alice. "Ingestion and Incorporation of Coral Mucus Aggregates by a Gorgonian Soft Coral." Marine Ecology - Progress Series 17 (1984): 193-99. http://www.nsm.buffalo.edu/Bio/burr/Publications/4%20Coffroth1984.pdf</ref> For example, ornate butterflyfish are known to consume large amounts of coral mucus when feeding.  <ref name="fishchannel">Scott, Michael. "Marine Fish Feeding Guilds." FishChannel.com. Accessed April 15, 2015. http://www.fishchannel.com/fish-health/saltwater-conditions/marine-fish-feeding-guilds.aspx</ref><ref name="cole">Cole, Andrew J., Morgan S. Pratchett, and Geoffrey P. Jones. "Diversity and Functional Importance of Coral-feeding Fishes on Tropical Coral Reefs." Fish and Fisheries 9 (2008): 1-22.</ref> One study showed that when coral mucus was artificially dispersed, fish assembled and avidly consumed it; researchers concluded that coral mucus is an important food source for reef inhabitants and could be an energy source linking the coral producers and small fish consumers in reef communities. <ref name="Benson"> Benson, A.A., and L. Muscatine. "Wax in Coral Mucus: Energy Transfer from Corals to Reef Fishes." Limnology and Oceanography 19, no. 5 (1974): 810-14. http://aslo.net/lo/toc/vol_19/issue_5/0810.pdf</ref>


Scavenging is another means of detritus feeding used by a variety of different fish species. Carrion is not very common on coral reefs, not because reef inhabitants don’t die often but because it is consumed very quickly by reef inhabitants. <ref name="rassweiler">Rassweiler, Andrew, and Thomas Rassweiler. "Does Rapid Scavenging Hide Non-predation Mortality in Coral-reef Communities?" Marine and Freshwater Research 62 (2011): 510-15.</ref>
One study found that all carrion left on a reef was consumed at very rapid rates within 24 hours, indicating that scavenged biomass is tightly recycled within the reef community. <ref name="rassweiler">Rassweiler, Andrew, and Thomas Rassweiler. "Does Rapid Scavenging Hide Non-predation Mortality in Coral-reef Communities?" Marine and Freshwater Research 62 (2011): 510-15.</ref> One example of a scavenger is the tiger shark. The stomach contents of the tiger shark have been found to contain a variety of unusual items, including license plates and old tires.<ref>"Tiger Shark." National Geographic. January 1, 2015. Accessed April 14, 2015. http://animals.nationalgeographic.com/animals/fish/tiger-shark/</ref>

==Notes==
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