Trends in elasmobranch feeding ecology studies
DOI:
https://doi.org/10.1590/Keywords:
Chondrichthyes, Conservation, Diet, Stomach Contents, Trophic EcologyAbstract
Reviews can be useful to provide an overview of specific knowledge areas to facilitate research guidelines and
the comparison of study results. Therefore, we aimed to detect regions, environments, and taxonomic groups of
elasmobranch that lack information on their feeding ecology and consequently need further attention. Using specific
keywords on diet and feeding studies, we searched for articles on the trophic and feeding ecology of elasmobranchs.
We found an increase in studies over the last 24 years, with an emphasis on stomach contents and stable isotopes.
The USA, Australia, Mexico, Argentina, and Brazil were the countries with the highest number of published articles.
Australia, the USA, Argentina, Brazil, Mexico, South Africa, Spain, Uruguay and India were the countries with the
highest number of species studied. Estuarine and freshwater environments seem to need more attention, since
studies on species in these environments were scarce (~3%). The total number of shark and ray species studied
showed no significant differences (164 and 186, respectively), but most of the articles reviewed concerned sharks
(67.5%). The larger elasmobranch families also need more attention, especially demersal species such as Rajidae,
Dasyatidae, Arhynchobatidae, and Sycliorhinidae. Feeding studies only analyze some of the data deficient (
References
Amundsen, P. A. & Sánchez‐Hernández, J. 2019. Feeding
studies take guts–critical review and recommendations
of methods for stomach contents analysis in fish.
Journal of Fish Biology, 95, 1364–1373.
Akhilesh, K. V., Bineesh, K. K., Gopalakrishnan, A., Jena,
J. K., Basheer, V. S. & Pillai, N. G. K. 2015. Checklist of
Chondrichthyans in Indian water. Journal of the Marine
Biological Association of India, 56, 109-120.
Atwood, T. B., Connolly, R. M., Ritchie, E. G., Lovelock, C.
E., Heithaus, M. R., Hays, G. C, Fourqurean, J. W. &
Macreadie, P. I. 2015. Predators help protect carbon
stocks in blue carbon ecosystems. Nature Climate
Change, 5, 1038-1045.
Ballantyne, J. S. & Robinson, J. W. 2010. Freshwater
elasmobranchs: a review of their physiology and
biochemistry. Journal of Comparative Physiology B,
, 475-493.
Barbini, S. A. & Lucifora, L. O. 2011. Feeding habits of the
Rio skate, Rioraja agassizi (Chondrichthyes: Rajidae),
from off Uruguay and north Argentina. Journal of the
Marine Biological Association of the United Kingdom,
, 1175-1184.
Barbini, S. A., Sabadin, D. E. & Lucifora, L. O. 2018.
Comparative analysis of feeding habits and dietary
niche breadth in skates: the importance of body size,
snout length, and depth. Reviews in Fish Biology and
Fisheries, 28, 625-636.
Barreto, R., Ferretti, F., Flemming, J. M., Amorim, A.,
Andrade, H., Worm, B. & Lessa, R. 2016. Trends in
the exploitation of South Atlantic shark populations.
Conservation Biology, 30, 792-804.
Baum, J. K., Myers, R. A., Kehler, D. G., Worm, B.,
HARLEY, S. J. & DOHERTY, P. A. 2003. Collapse and
conservation of shark populations in the Northwest
Atlantic. Science, 299, 389-392.
Belicka, L. L., Matich, P., Jaffé, R. & Heithaus, M. R. 2012.
Fatty acids and stable isotopes as indicators of early-life
feeding and potential maternal resource dependency
in the bull shark Carcharhinus leucas. Marine Ecology
Progress Series, 455, 245-256.
Bornatowski, H., Wosnick, N., Do Carmo, W. P. D., Corrêa,
M. F. M. & Abilhoa, V. 2014a. Feeding comparisons
of four batoids (Elasmobranchii) in coastal waters
of southern Brazil. Journal of the Marine Biological
Association of the United Kingdom, 94, 1491-1499.
Bornatowski, H., Braga, R. R., Abilhoa, V. & Corrêa, M. F.
M. 2014b. Feeding ecology and trophic comparisons of
six shark species in a coastal ecosystem off southern
Brazil. Journal of Fish Biology, 85, 246-263.
Bornatowski, H., Navia, A. F., Braga, R. R., Abilhoa, V. &
Corrêa, M. F. M. 2014c. Ecological importance of sharks
and rays in a structural foodweb analysis in southern
Brazil. ICES Journal of Marine Science, 71, 1586-1592.
Bornatowski, H., Angelini, R., Coll, M., Barreto, R. R. &
Amorim, A. F. 2018a. Ecological role and historical
trends of large pelagic predators in a subtropical marine
ecosystem of the South Atlantic. Reviews in Fish Biology
and Fisheries, 28, 241-259.
Bornatowski, H., Braga, R. R. & Barreto, R. P. 2018b.
Elasmobranchs consumption in Brazil: impacts and
consequences. In: ROSSI-SANTOS, M. R. & FINKL,
C. W. (eds.), Advances in marine vertebrate research in
Latin America (pp. 251-262). Berlin: Springer.
Braga, R. R., Bornatowski, H, & Vitule, J. R. S. 2012.
Feeding ecology of fishes: an overview of worldwide
publications. Reviews in Fish Biology and Fisheries, 22,
-929,
Camhi, M. D., Valenti, S. V., Fordham, S. V., Fowler, S. L.
& Gibson, C. (eds.). 2009. The Conservation Status of
Pelagic Sharks and Rays: Report of the IUCN Shark
Specialist Group Pelagic Shark Red List Workshop.
Newbury: IUCN Species Survival Commission Shark
Specialist Group.
Cliff, G. 1995. Sharks caught in the protective gill nets off
KwaZulu-Natal, South Africa. 8. The great hammerhead
shark Sphyrna mokarran (Rüppell). South African
Journal of Marine Science, 15, 105-114.
Cliff, G. & Dudley, S. F. J. 1991. Sharks caught in the
protective gill nets off Natal, South Africa. 4. The bull
shark Carcharhinus leucas Valenciennes. South African
Journal of Marine Science, 10, 253-270.
Coll, M., Navarro, J. & Palomera, I. 2013. Ecological role,
fishing impact, and management options for the recovery
Elasmobranch feeding ecology studies
Ocean and Coastal Research 2023, v71:e23060 11
Bornatowski et al.
of a Mediterranean endemic skate by means of food web
models. Biological Conservation, 157, 108-120.
Coll, M., Akoglu, E. & Arreguin-Sanchez, F. 2015. Modelling
dynamic ecosystems: venturing beyond boundaries with
the Ecopath approach. Reviews in Fish Biology and
Fisheries, 25, 413-424.
Compagno, L. J. V., Dando, M. & Fowler, S. 2005. A field
guide to the sharks of the world. London: Collins.
Compagno, L. J. V. 2013. National Plan of Action for the
Conservation and Management of Sharks (NPOASharks). South Africa: Department of Agriculture,
Forestry and Fisheries.
Consalvo, I., Iraci Sareri, D., Bottaro, M., Tudisco, A.,
Cantone, G. & Vacchi, M. 2010. Diet composition of
juveniles of rough ray Raja radula (Chondrichthyes:
Rajidae) from the Ionian Sea. Italian Journal of Zoology
, 438-442.
Cortés, E. 1997. A critical review of methods of studying
fish feeding based on analysis of stomach contents:
application to elasmobranch fishes. Canadian Journal
of Fisheries and Aquatic Sciences 54, 726-738.
Cortés, E. 1999. Standardized diet compositions and trophic
levels of sharks. ICES Journal of Marine Science, 56,
-717.
Cortés, E., Papastamatiou, Y. P., Carlson, J. K., FerryGraham, L. & Wetherbee, B. M. 2007. An overview of
the feeding ecology and physiology of elasmobranch
fishes. In: CYRINO, J. E. P., BUREAU, D. & KAPOOR,
B. G. (eds) Feeding and digestive functions in fishes
(pp. 393-443). Rawalpindi: Science Publishers.
Couturier, L. I. E., Rohner, C. A. & Richardson, A. J. 2013.
Unusually high levels of n-6 polyunsaturated fatty acids
in whale sharks and reef manta rays. Lipids, 48(10),
-1034.
Davidson, L. N., Krawchuk, M. A. & Dulvy, N. K. 2016. Why
have global shark and ray landings declined: improved
management or overfishing? Fish and Fisheries, 17,
-458.
Del Moral–Flores, L. F., Morrone, J. J., Alcocer Durand, J.,
Espinosa–Pérez, H. & Pérez–Ponce De León, G. 2015.
Listado anotado de los tiburones, rayas y quimeras
(Chondrichthyes, Elasmobranchii, Holocephali) de
México. Arxius de Miscellània Zoològica, 13, 47-163.
Dulvy, N. K., Baum, J. K., Clarke, S., Compagno, L. J. V.,
Cortés, E., Domingo, A., Fordham, S., Fowler, S., Francis,
M. P., Gibson, C., Martínez, J., Musick, J. A., Soldo, A.,
Stevens, J. D. & VALENTI, S. 2008. You can swim but
you can’t hide: the global status and conservation of
oceanic pelagic sharks and rays. Aquatic Conservation,
, 459-482.
Dulvy, N. K., Fowler, S. L., Musick, J. A., Cavanagh, R. D.,
Kyne, P. M., Harrison, L. R., Carlson, J. K., Davidson,
L. N. K., Fordham, S. V., Francis, M. P., Pollock, C. M.,
Simpfendorfer, C. A., Burgess, G. H., Carpenter, K. E.,
Compagno, L. J. V., Ebert, D. A., Gibson, C., Heupel,
M. R., Livingstone, S. R., Sanciangco, J. C., Stevens,
J. D., Valenti, S. & White, W. T. 2014. Extinction risk
and conservation of the world’s sharks and rays. Elife,
, e00590.
Dulvy, N. K., Pacoureau, N., Rigby, C. L., Pollom, R. A.,
Jabado, R. W., Ebert, D. A., Finucci, B., Pollock, C. M.,
Cheok, J., Derrick, D. H., Herman, K. B., Sherman, C.
S., Vanderwright, W. J., Lawson, J. M., Walls, R. H. L.,
Carlson, J. K., Charvet, P., Bineesh, K. K., Fernando,
D., Ralph, G. M., Matsushiba, J. H., Hilton-Taylor, C.,
Fordham, S. V. & Simpfendorfer, C. A. 2021. Overfishing
drives over one-third of all sharks and rays toward a
global extinction crisis. Current Biology, 31, 4773–4787.
Ebert, D. A. 1994. Diet of the sixgill shark Hexanchus
griseus off southern. South African Journal of Marine
Science, 14, 213-218.
Ebert, D. A. 2002. Ontogenetic changes in the diet of the
sevengill shark (Notorynchus cepedianus). Marine and
Freshwater Research, 53, 517-523.
Ebert, D. A. & Bizzarro, J. J. 2007. Standardized
diet compositions and trophic levels of skates
(Chondrichthyes: Rajiformes: Rajoidei). Environmental
Biology of Fishes, 80, 221–237.
Ebert, D. A. & Stehmann, M. F. W. 2013. Sharks, batoids
and chimaeras of the North Atlantic. Rome: Food and
Agricultural Organization of the United Nations.
Ebert, D. A., Ho, H. C., White, W. T. & Carvalho, M. R. 2013.
Introduction to the systematic and biodiversity of sharks,
rays, and chimaeras (Chondrichthyes) of Taiwan.
Zootaxa, 3752, 5-19.
Ebert, D. A. & Van Hees, K. E. 2015. Beyond Jaws:
rediscovering the ‘lost sharks’ of southern Africa. African
Journal of Marine Science, 37, 141–156.
Ebert, D. A., Bigman, J. S. & Lawson, J. M. 2017. Biodiversity,
life history, and conservation of Northeastern Pacific
Chondrichthyans. Advances in Marine Biology, 77, 9-78.
Ellis, J. R., Pawson, M. G. & Shackley, S. E. 1996. The
comparative feeding ecology of six species of shark and
four species of ray (Elasmobranchii) in the north-east
Atlantic. Journal of the Marine Biological Association of
the United Kingdom, 76, 89-106.
Fallows, C., Gallagher, A. J. & Hammerschlag, N. 2013.
White sharks (Carcharodon carcharias) scavenging
on whales and its potential role in further shaping the
ecology of an apex predator. PLoS One, 8, e60797.
Fahmi. 2010. Sharks and rays in Indonesia. Marine
Research in Indonesia, 35, 43-54.
Freitas, R. H., Aguiar, A. A., Freitas, A. K. C., Lima, S.
M. & Valentin, J. L. 2019. Unravelling the foraging
behavior of the southern stingray, Hypanus americanus
(Myliobatiformes: Dasyatidae) in a Southwestern
Atlantic MPA. Neotropical Ichthyology, 17, e180131.
Froese, R. & Pauly, D. (eds.). 2023. FishBase. World Wide
Web electronic publication. www.fishbase.org, version
(02/2023)
Garrone-Neto, D., Haddad Jr, V. & Gadig, O. B. F. 2014.
Record of ascending passage of potamotrygonid
stingrays through navigation locks: implications for
the management of nonnative species in the Upper
Parana´ River basin, southeastern Brazil. Management
of Biological Invasions, 5, 113–119.
Guallart, J., García‐Salinas, P., Ahuir‐Baraja, A. E., Guimerans,
M., Ellis, J. R. & Roche, M. 2015. Angular roughshark
Oxynotus centrina (Squaliformes: Oxynotidae) in captivity
feeding exclusively on elasmobranch eggs: an overlooked
feeding niche or a matter of individual taste?. Journal of
Fish Biology, 87, 1072-1079.
Elasmobranch feeding ecology studies
Ocean and Coastal Research 2023, v71:e23060 12
Bornatowski et al.
Heithaus, M. R. & Vaudo, J. J. 2012. Predator-Prey
Interactions. In: Carrier, J. C., Musick, J. A. & Heithaus,
M. R. (eds) Biology of Sharks and their Relatives. Boca
Raton: CRC Press.
Heupel, M. R. & Simpfendorfer, C. A. 2010. Science
or slaughter: need for lethal sampling of sharks.
Conservation Biology, 24, 1212-1218.
Hussey, N. E., Macneil, M. A. & Fisk, A. T. 2010. The
requirement for accurate diet-tissue discrimination
factors for interpreting stable isotopes in sharks.
Hydrobiologia, 654, 1-5.
Hussey, N. E., Macneil, M. A., Olin, J. A., Mcmeans, B. C.,
Kinney, M. J., Chapman, D. D. & Fisk, A. T. 2012. Stable
isotopes and elasmobranchs: tissue types, methods,
applications and assumptions. Journal of Fish Biology,
, 1449-1484.
Hussey, N. E., Macneil, M. A., Mcmeans, B. C., Olin, J. A.,
Dudley, S. F. J., Cliff, G., Wintner, S. P., Fennessy, S.
T. & Fisk, A. T. 2014. Rescaling the trophic structure of
marine food webs. Ecology Letters, 17, 239-250.
IUCN. 2023. The IUCN Red List of Threatened Species.
www.iucnredlist.org/
IUCN Standards and Petitions Committee. 2019. Guidelines
for Using the IUCN Red List Categories and Criteria.
Version 14. http://www.iucnredlist.org/documents/
RedListGuidelines.pdf.
Jarman, S. N. & Wilson, S. G. 2004. DNA‐based species
identification of krill consumed by whale sharks. Journal
of Fish Biology, 65, 586-591.
Lasso, C. A., Rosa, R. S., Morales-Betancourt, M. A.,
Garrone-Neto, D. & Carvalho, M. R. 2016. XV. Rayas
de agua dulce (Potamotrygonidae) de Suramerica. Parte
II. Colombia, Brasil, Perú, Bolivia, Paraguay, Uruguay y
Argentina. Bogotá: Instituto de Investigación de Recursos
Biológicos Alexander von Humboldt.
Last, P. R. & Stevens, J. D. 2009. Sharks and Rays of
Australia. London: Harvard University Press.
Last, P. R. & White, W. T. 2011. Biogeographic patterns in
the Australian chondrichthyan fauna. Journal of Fish
Biology, 79, 1193-1213.
Last, P. R., White, W. T., Carvalho, M. R., Séret, B.,
Stehmann, M. F. W. & Naylor, G. J. P. 2016. Rays of the
World. Ithaca: Cornell University Press.
Lea, J. S. E., Daly, R., Leon, C., Daly C. A. K. & Clarke, C.
R. 2019. Life after death: behaviour of multiple shark
species scavenging a whale carcass. Marine and
Freshwater Research, 70, 302-306.
Leigh, S. C., Papastamatiou, Y. P. & German, D. P.
Seagrass digestion by a notorious ‘carnivore’.
Proceedings of the Royal Society B, 285, 20181583.
Lowe, C. G., Wetherbee, B. M., Crow, G. L. & Tester, A. L.
Ontogenetic dietary shifts and feeding behavior of
the tiger shark, Galeocerdo cuvier, in Hawaiian waters.
Environmental Biology of Fishes, 47, 203-211.
Macneil, M. A., Skomal, G. B. & Fisk, A. T. 2005. Stable
isotopes from multiple tissues reveal diet switching in
sharks. Marine Ecology Progress Series, 302, 199–206.
Martin, R. A. 2005. Conservation of freshwater and euryhaline
elasmobranchs: a review. Journal of the Marine Biological
Association of the United Kingdom, 85, 1049–1073.
Martinho, F., Sá, C., Falcão, J., Cabral, H. N. & Pardal, M. Â.
Comparative feeding ecology of two elasmobranch
species, Squalus blainville and Scyliorhinus canicula,
off the coast of Portugal. Fishery Bulletin, 110, 71-84.
Menni, R. C. & Lucífora, L. O. 2007. Condrictios de la
Argentina y Uruguay: Lista de Trabajo. Buenos Aires:
FCNyM.
Metcalfe, J. D. & Craig, J. F. 2011. Ethical justification for
the use and treatment of fishes in research: an update.
Journal of Fish Biology, 78, 393-394.
Meyer, C. G. & Holland, K. N. 2012. Autonomous
measurement of ingestion and digestion processes in
free-swimming sharks. Journal of Experimental Biology,
, 3681-3684.
Molina, J. M. & Cooke, S. J. 2012. Trends in shark bycatch
research: current status and research needs. Reviews
in Fish Biology and Fisheries, 22, 719-737.
Motta, P. J. 2004. Prey capture behavior and feeding
mechanics of elasmobranchs. In: Carrier, J. C., Musick,
J. A. & Heithaus, M. R. (eds) Biology of Sharks and their
Relatives. Boca Raton: CRC Press.
Mourier, J., Maynard, J., Parravicini, V., Ballesta, L., Clua,
E., Domeier, M. L. & Planes, S. 2016. Extreme inverted
trophic pyramid of reef sharks supported by spawning
groupers. Current Biology, 26, 2011-2016.
Mulas, A., Bellodi, A., Cannas, R., Carbonara, P., Cau, A.,
Marongiu, M. F., Pesci, P., Porcu, C. & Follesa, M. C. 2019.
Resource partitioning among sympatric elasmobranchs
in the central-western Mediterranean continental shelf.
Marine Biology, 166, 153.
Navia, A. F., Mejía-Falla, P. A. & Giraldo, A. 2007. Feeding
ecology of elasmobranch fishes in coastal waters of the
Colombian Eastern Tropical Pacific. BMC Ecology, 7, 8.
Navia, A. F., Mejía-Falla, P. A., López-García, J., Giraldo, A.
& Cruz-Escalona, V. H. 2017. How many trophic roles
can elasmobranchs play in a marine tropical network?.
Marine and Freshwater Research, 68, 1342-1353.
Pethybridge, H. R., Parrish, C. C., Bruce, B. D., Young, J.
W. & Nichols, P. D. 2014. Lipid, fatty acid and energy
density profiles of white sharks: insights into the feeding
ecology and ecophysiology of a complex top predator.
PloS One, 9, e97877.
Queiroz, N., Humphries, N. E., Couto, A. Et al. 2019. Global
spatial risk assessment of sharks under the footprint of
fisheries. Nature, 572, 461–466.
R CORE TEAM. 2018. R: A language and environment
for statistical computing. Vienna: R Foundation for
Statistical Computing. https://www.Rproject.org/
Riccioni, G., Stagioni, M., Piccinetti, C. & Libralato, S.
A metabarcoding approach for the feeding habits
of European hake in the Adriatic Sea. Ecology and
Evolution, 8, 10435–10447.
Roff, G., Brown, C. J., Priest, M. A. & Mumby, P. J. 2018.
Decline of coastal apex shark populations over the past
half century. Communications Biology, 1, 223.
Rosa, R. S. & Gadig, O. B. F. 2014. Conhecimento da
diversidade dos Chondrichthyes marinhos no Brasil: a
contribuição de José Lima de Figueiredo. Arquivos de
Zoologia, 45, 89-104.
Elasmobranch feeding ecology studies
Ocean and Coastal Research 2023, v71:e23060 13
Bornatowski et al.
Rosel, P. E. & Kocher, T. D. 2002. DNA‐based identification
of larval cod in stomach contents of predatory fishes.
Journal of Experimental Marine Biology and Ecology,
, 75-88.
Sandin, S.A., Smith, J.E., Demartini, E.E., Dinsdale, E.A.,
Donner, S.D., Friedlander, A.M., Konotchick, T., Malay,
M., Maragos, J.E., Obura, D., Pantos, O., Paulay, G.,
RICHIE, M., Rohwer, F., Schroeder, R. E., Walsh,
S., Jackson, J. B. C., Knowlton, N. & Sala, E. 2008.
Baselines and degradation of coral reefs in the Northern
Line Islands. PloS One, 3, e1548.
Santos, D. A., De Paiva Affonso, I., Okada, E. K., Gomes,
L. C., Bornatowski, H. & Vitule, J. R. S. 2019. Societal
perception, impacts and judgment values about invasive
freshwater stingrays. Biological Invasions, 21, 3593-3606.
Shibuya, a., zuanon, j. & tanaka, S. 2012. Feeding behavior
of the Neotropical freshwater stingray Potamotrygon
motoro (Elasmobranchii: Potamotrygonidae).
Neotropical Ichthyology, 10, 189-196.
Simpfendorfer, C. A., Heupel, M. R., White, W. T., & Dulvy,
N. K. 2011. The importance of research and public
opinion to conservation management of sharks and
rays: a synthesis. Marine and Freshwater Research,
(6), 518-527.
Stein, R. W., Mull, C. G., Kuhn, T. S., Aschliman, N. C.,
Davidson, L. N. K., Joy, J. B., Smith, G. J., Dulvy, N. K.
& Mooers, A. O. 2018. Global priorities for conserving
the evolutionary history of sharks, rays and chimaeras.
Nature Ecology & Evolution, 2, 288-298.
Stevens, J. D., Bonfil, R., Dulvy, N. K. & Walker, P. A. 2000.
The effects of fishing on sharks, rays, and chimaeras
(chondrichthyans), and the implications for marine
ecosystems. ICES Journal of Marine Science, 57, 476-494.
Strong, W. R., Snelson, F. F. & Gruber, S. H. 1990.
Hammerhead shark predation on stingrays: an
observation of prey handling by Sphyrna mokarran.
Copeia, 1990, 836-840.
Valls, M., Rueda, L. & Quetglas, A. 2017. Feeding strategies
and resource partitioning among elasmobranchs and
cephalopods in Mediterranean deep-sea ecosystems.
Deep Sea Research Part I: Oceanographic Research
Papers, 128, 28-41.
Vitule, J. R. S., Skóra, F. & Abilhoa, V. 2012. Homogenization
of freshwater fish faunas after the elimination of a
natural barrier by a dam in Neotropics. Diversity and
Distributions, 18, 111-120.
Wanchana, W., Ali, A. & Putsa, S. 2016. Recording sharks
and rays statistics from Southeast Asia at species level.
Fish for the People, 14, 2-6.
Weigmann, S. 2017. Reply to BORSA 2017: Comment on
‘Annotated checklist of the living sharks, batoids and
chimaeras (Chondrichthyes) of the world, with a focus
on biogeographical diversity by Weigmann (2016)’.
Journal of Fish Biology, 90, 1176-1181.
Werry, J. M., Lee, S. Y., Otway, N. M., Hu, Y. & Sumpton,
W. 2011. A multi-faceted approach for quantifying the
estuarine–nearshore transition in the life cycle of the
bull shark, Carcharhinus leucas. Marine and Freshwater
Research, 62, 1421-1431.
Wetherbee, B. M. & Cortés, E. 2004. Food consumption
and feeding habits. In: CARRIER, J. C., MUSICK, J. A.
& HEITHAUS, M. R. (eds) Biology of Sharks and their
Relatives. Boca Raton: CRC Press.
Wetherbee, B., Cortés, E. & Bizzarro, J. J. 2012. Food
consumption and feedings habits. In: CARRIER, J. C.,
MUSICK, J. A. & HEITHAUS, M. R. (eds.). Biology of
Sharks and their Relatives. Boca Raton: CRC Press.
Wilga, C. D., Motta, P. J. & Sanford, C. P. 2007. Evolution
and ecology of feeding in elasmobranchs. Integrative
and Comparative Biology, 47, 55-69.
