White Shark or Shortfin Mako? Lamnid sharks identification in an overlapping area of the Equatorial Southwestern Atlantic
DOI:
https://doi.org/10.1590/Keywords:
Isurus oxyrinchus, Carcharodon carcharias, small-scale fisheries, Lamnidae, Misidentificatio, DNA barcodingAbstract
Despite their worldwide distribution, the occurrence of Lamnidae species in tropical zones remain poorly
understood. In the Equatorial Southwestern Atlantic, the Shortfin Mako, Isurus oxyrinchus, has been captured
in commercial fisheries, whereas the White Shark, Carcharodon carcharias, is only rarely recorded. This study
aimed to characterize lamnid records for the Equatorial Southwestern Atlantic, as well as to apply molecular
markers to identify a lamnid carcass from the only area in the region where these two species are known to
overlap (off Ceará State, Northeastern Brazil). The landed carcass, although sold in the market as a White
Shark, in fact was a Shortfin Mako, I. oxyrinchus, with a total of 99.6% and 100% match to its ND2 and D-loop
mitochondrial DNA sequences, respectively. Despite the existence of a record of landings of elasmobranchs by
artisanal and commercial fisheries from Ceará, lamnid landings in this region are still rare and comprise only
four records for I. oxyrinchus, including juvenile specimens. As for the White Shark, Ceará State remains the
only region in the Equatorial Southwestern Atlantic to hold records (three) of C. carcharias (all of them adults).
Oceanographic conditions in this region may contribute to the occasional presence of large lamnid specimens.
Moreover, fishers seem to support the presence of both lamnid species in the region.
References
Abascal, F. J., Quintans, M., Ramos-Cartelle, A. & Mejuto,
J. 2011. Movements and environmental preferences of
the shortfin mako, Isurus oxyrinchus, in the southeastern
Pacific Ocean. Marine Biology, 158(5), 1175–1184.
Almerón-Souza, F., Sperb, C., Castilho, C. L., Figueiredo,
P. I. C. C., Gonçalves, L. T., Machado, R., Oliveira, L.
R., Valiati, V. H. & Fagundes, N. J. R. 2018. Molecular
identification of shark meat from local markets in
Southern Brazil based on DNA barcoding: evidence for
mislabeling and trade of endangered speciesa. Frontiers
in Genetics, 9, 138. DOI: https://doi.org/10.3389/
fgene.2018.00138
Altschul, S. F., Gish, W., Miller, W., Myers, E. W. & Lipman,
D. J. 1990. Basic local alignment search tool. Journal of
Molecular Biology, 215(3), 403–410.
Amorim, A.F., Arfelli, C.A., Bornatowski, H. et al. Rare
giants? A large female great white shark caught in
Brazilian waters. Mar Biodiv 48, 1687–1692 (2018).
DOI: https://doi.org/10.1007/s12526-017-0656-9
Amorim, A. F., Arfelli, C. A. & Fagundes, L. 1998. Pelagic
elasmobranchs caught by longliners off southern Brazil
during 1974–97: an overview. Marine and Freshwater
Research, 49(7), 621–632. DOI: https://doi.org/10.1071/
MF97111
Andreotti, S., von der Heyden, S., Henriques, R., Rutzen, M.,
Meÿer, M., Oosthuizen, H. & Matthee, C. A. 2016. New
insights into the evolutionary history of white sharks;
Carcharodon carcharias. Journal of Biogeography,
(2), 328–339. DOI: https://doi.org/10.1111/jbi.12641
Barcellos, L. R., Barreto, R. & Lessa, R. P. T. 2022.
Occurrence and population structure of sharks in two
ecologically or biologically significant marine areas off
north‐eastern Brazil. Aquatic Conservation: Marine and
Freshwater Ecosystems, 32(5), 713–726. DOI: https://
doi.org/10.1002/aqc.3787
Barreto, R. R., de Farias, W. K. T., Andrade, H., Santana,
F. M. & Lessa, R. 2016. Age, Growth and Spatial
Distribution of the Life Stages of the Shortfin Mako, Isurus
oxyrinchus (Rafinesque, 1810) Caught in the Western
and Central Atlantic. PloS One, 11(4), e0153062. DOI:
https://doi.org/10.1371/journal.pone.0153062
Barros, A. C., de Oliveira, G. M., Evangelista, J. E. V.,
dos Santos, A. R. B., Haimovici, M. & Fischer, L. G.
Prospecções na região Nordeste. In: Haimovici,
M. (ed.), A prospecção pesqueira e abundância de
estoques marinhos no Brasil nas décadas de 1960
a 1990: Levantamento de dados e Avaliação Crítica
(pp. 89-123). Brasília, DF: Ministério do Meio Ambiente.
Bigelow, H.B. & W.C. Schroeder, 1948. Sharks. p. 59-546.
In: J. Tee-Van, C.M. Breder, S.F. Hildebrand, A.E. Parr
and W.C. Schroeder (eds.) Fishes of the Western North
Atlantic. Part one. Lancelets, cyclostomes, sharks.
Sears Foundation for Marine Research, Yale University,
New Haven. 576 p.
Bonfil, R., Meÿer, M., Scholl, M. C., Johnson, R., O’Brien,
S., Oosthuizen, H., Swanson, S., Kotze, D. & Paterson,
M. 2005. Transoceanic migration, spatial dynamics,
and population linkages of white sharks. Science,
(5745), 100–103.
Boustany, A. M., Davis, S. F., Pyle, P., Anderson, S. D., Le
Boeuf, B. J. & Block, B. A. 2002. Expanded niche for
white sharks. Nature, 415(6867), 35–36.
Brasil. 2022. Portaria GM/MMA n. 300, de 13 de dezembro de
Brasília, DF: Ministério de Meio Ambiente. Available
from: https://www.in.gov.br/en/web/dou/-/portaria-gm/
mma-n-300-de-13-de-dezembro-de-2022-450425464
Access date: 11 jan. 2023
Canani, G. & Oddone, M. C. 2020. Reproductive biology
of Isurus oxyrinchus captured by the south Brazilian
surface longline commercial fleet in the Southwest
Atlantic Ocean, with data on CPUE and size distribution
by sex. Journal of Northwest Atlantic Fishery Science,
DOI: https://doi.org/10.1101/2024.11.04.621869
Chang, C.-H., Chiang, W.-C., Lin, Y.-S., Jang-Liaw, N.-H.
& Shao, K.-T. 2016. Complete mitochondrial genome of
the longfin mako shark, Isurus paucus (Chondrichthyes,
Lamnidae). Mitochondrial DNA. Part A, DNA Mapping,
Sequencing, and Analysis, 27(1), 690–691. DOI: https://
doi.org/10.3109/19401736.2014.913145
Chang, C.-H., Jabado, R. W., Lin, Y.-S. & Shao, K.-T.
The complete mitochondrial genome of the
sand tiger shark, Carcharias taurus (Chondrichthyes,
Odontaspididae). Mitochondrial DNA, 26(5), 728–729.
DOI: https://doi.org/10.3109/19401736.2013.845761
Chang, C.-H., Jang-Liaw, N.-H., Lin, Y.-S., Carlisle, A., Hsu,
H. H., Liao, Y.-C. & Shao, K.-T. 2016. The complete
mitochondrial genome of the salmon shark, Lamna
ditropis (Chondrichthyes, Lamnidae). Mitochondrial
DNA. Part A, DNA Mapping, Sequencing, and Analysis,
(1), 316–317. DOI: https://doi.org/10.3109/19401736
.2014.892095
Chang, C.-H., Shao, K.-T., Lin, Y.-S., Fang, Y.-C. & Ho, H.-C.
The complete mitochondrial genome of the great
white shark, Carcharodon carcharias (Chondrichthyes,
Lamnidae). Mitochondrial DNA, 25(5), 357–358. DOI:
https://doi.org/10.3109/19401736.2013.803092
Christiansen, H. M., Lin, V., Tanaka, S., Velikanov, A.,
Mollet, H. F., Wintner, S. P., Fordham, S. V., Fisk, A. T.
& Hussey, N. E. 2014. The last frontier: catch records of
white sharks (Carcharodon carcharias) in the Northwest
Pacific Ocean. PloS One, 9(4), e94407. DOI: https://doi.
org/10.1371/journal.pone.0094407
Clement, M., Snell, Q., Walke, P., Posada, D. & Crandall, K.
TCS: estimating gene genealogies. In: Proceedings
th International Parallel and Distributed Processing
Symposium Ft. Lauderdale, FL, USA, 2002, pp. 7 pp-,
DOI: https://doi.org/10.1109/IPDPS.2002.1016585.
Lamnids in the Equatorial Southwestern Atlantic
Ocean and Coastal Research 2024, v72:e24075 9
Gonçalves-Neto et al.
Colefax, A. P., Kelaher, B. P., Pagendam, D. E. & Butcher,
P. A. 2020. Assessing White Shark (Carcharodon
carcharias) Behavior along coastal beaches for
conservation-focused shark mitigation. Frontiers
in Marine Science, 7. DOI: https://doi.org/10.3389/
fmars.2020.00268
Comassetto, L., Hazin, F. H. V., Hazin, H. G., Sant’Ana, R.,
Mourato, B. & Carvalho, F. 2017. Standardized catch
rates of shortfin mako caught by the brazilian fleet
(1978-2012) using a generalized linear mixed model
(glmm), with a delta log approach. Collective Volume of
Scientific Papers, 73(8), 2930–2944.
Compagno, L. J. V. 2002. Sharks of the World. An
annotated and illustrated catalogue of Shark species
know to date (Vol. 2). Bullhead, mackerel and
carpet sharks (Heterodontiformes, Lamniformes and
Orectolobiformes). FAO Species Catalogue for Fishery
Purposes. No. 1, Vol. 2. Rome, FAO. 2001. 269p.
Corrigan, S., Lowther, A. D., Beheregaray, L. B., Bruce,
B. D., Cliff, G., Duffy, C. A., Foulis, A., Francis, M. P.,
Goldsworthy, S. D., Hyde, J. R., Jabado, R. W., Kacev,
D., Marshall, L., Mucientes, G. R., Naylor, G. J. P.,
Pepperell, J. G., Queiroz, N., White, W. T., Wintner, S. P.
& Rogers, P. J. 2018. Population connectivity of the highly
migratory shortfin Mako (Isurus oxyrinchus Rafesquine
and implications for management in the southern
hemisphere. Frontiers in Ecology and Evolution, 6. DOI:
https://doi.org/10.3389/fevo.2018.00187
Curtis, T. H., McCandless, C. T., Carlson, J. K., Skomal,
G. B., Kohler, N. E., Natanson, L. J., Burgess, G. H.,
Hoey, J. J. & Pratt, H. L., Jr. 2014. Seasonal distribution
and historic trends in abundance of white sharks,
Carcharodon carcharias, in the western North Atlantic
Ocean. PloS One, 9(6), e99240. DOI: https://doi.
org/10.1371/journal.pone.0099240
Dewar, H., Domeier, M. & Nasby-Lucas, N. 2004. Insights
into Young of the Year White Shark, Carcharodon
carcharias, Behavior in the Southern California Bight.
Environmental Biology of Fishes, 70(2), 133–143. DOI:
https://doi.org/10.1023/B:EBFI.0000029343.54027.6a
Díaz-Jaimes, P., Uribe-Alcocer, M., Adams, D. H., RangelMorales, J. M. & Bayona-Vásquez, N. J. 2016. Complete
mitochondrial genome of the porbeagle shark, Lamna
nasus (Chondrichthyes, Lamnidae). Mitochondrial DNA
Part B, 1(1), 730–731. DOI: https://doi.org/10.1080/238
2016.1233465
Dudgeon, C. L., Blower, D. C., Broderick, D., Giles, J. L.,
Holmes, B. J., Kashiwagi, T., Krück, N. C., Morgan,
J. A. T., Tillett, B. J. & Ovenden, J. R. 2012. A review
of the application of molecular genetics for fisheries
management and conservation of sharks and rays.
Journal of Fish Biology, 80(5), 1789–1843. DOI: https://
doi.org/10.1111/j.1095-8649.2012.03265.x
Ebert, D. A., Dando, M. & Fowler, S. 2021. Sharks of
the World: A Complete Guide. Princeton, Princeton
University Press.
Evans, S. 2015. Shark Week and the rise of infotainment
in science documentaries. Communication Research
Reports: CRR, 32(3), 265–271. DOI: https://doi.org/10
.1080/08824096.2015.1052903
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(4), e60797.
DOI: https://doi.org/10.1371/journal.pone.0060797
Faria, V. V., McDavitt, M. T., Charvet, P., Wiley, T. R.,
Simpfendorfer, C. A. & Naylor, G. J. P. 2013. Species
delineation and global population structure of Critically
Endangered sawfishes (Pristidae). Zoological Journal of
the Linnean Society, 167(1), 136–164. DOI: https://doi.
org/10.1111/j.1096-3642.2012.00872.x
Faria, V. V., Rolim, L. S., Vaz, L. A. L. & Furtado-Neto, M. A.
A. 2012. Reevaluation of RAPD markers involved in a
case of stingray misidentification (Dasyatidae: Dasyatis).
Genetics and Molecular Research, 11(4), 3835–3845.
DOI: https://doi.org/10.4238/2012.October.25.2
Freitas, I. A. 1976. Pescarias experimentais de tubarões,
com espinhéis de anzóis, em águas costeiras do estado
do Ceará (Brasil) (Trabalho de Conclusão de Curso).
Fortaleza: Universidade Federal do Ceará.
Gadig, O. B. F. 2001. Tubarões da costa brasileira (tese de
doutorado). Rio Claro: Universidade Estadual Paulista.
Gadig, O. B. & Rosa, R. 1996. Occurrence of the white
shark along the Brazilian coast. In: Klimley, A. P. &
Ainley, D. G. (eds.), Great White Sharks: The biology
of Carcharodon carcharias (p. 347–350). Cambridge:
Academic Press.
Giareta, E. P., Prado, A. C., Leite, R. D., Padilha, É.,
Santos, I. H., Wosiak, C. C. D. L. & Wosnick, N. 2021.
Fishermen’s participation in research and conservation
of coastal elasmobranchs. Ocean & Coastal
Management, 199, 105421.
Gomes, U. L., Signori, C. N., Gadig, O. B. F., Santos, H.
R. & Vicente, M. M. 2019. Guia para identificação
dos tubarões, raias e quimeras do Rio de Janeiro
(Chondrichthyes: Elasmobranchii e Holocephali).
Revista Nordestina de Biologia, 27(1). DOI: https://doi.
org/10.22478/ufpb.2236-1480.2019v27n1.47122
Gonçalves Neto, J. B., Lima, C. de F., Jucá-Queiroz, B.,
Freitas, J. E. P., Charvet, P. & Faria, V. V. 2022. White
sharks in the equatorial southwest Atlantic. In: Sharks
International 2022 conference, Valencia.
Gorman, J., Marra, N., Shivji, M. S. & Stanhope, M. J.
The complete mitochondrial genome of an
Atlantic Ocean Shortfin Mako Shark, Isurus oxyrinchus.
Mitochondrial DNA Part B, 4(2), 3642–3643.
Hazin, F. H. V., Zagaglia, J. R., Broadhurst, M. K.,
Travassos, P. E. P. & Bezerra, T. R. Q. 1998. Review of
a small-scale pelagic longline fishery off northeastern
Brazil. Marine Fisheries Review, 60(3). Available from:
https://aquadocs.org/handle/1834/26422. Access date:
apr. 01
Hazin, F, Hazin, H. & Travassos, P. 2007. CPUE and catch
trends of shark species caught by Brazilian longliners in
the Southwestern Atlantic Ocean. Collective Volume of
Scientific Papers, 60(2), 636–647.
Heist, E. J. 2000. Genetics: stock identification. In: Musick,
J. A. & Bonfil, R. (eds.). Management techniques for
elasmobranch fisheries. Roma: FAO. https://www.fao.
org/4/a0212e/A0212E09.htm#ch5
Hester, J., Atwater, K., Bernard, A., Francis, M. & Shivji, M.
S. 2015. The complete mitochondrial genome of the
basking shark Cetorhinus maximus (Chondrichthyes,
Lamnids in the Equatorial Southwestern Atlantic
Ocean and Coastal Research 2024, v72:e24075 10
Gonçalves-Neto et al.
Cetorhinidae). Mitochondrial DNA, 26(5), 730–731.
DOI: https://doi.org/10.3109/19401736.2013.845762
Jucá-Queiroz, B., Silva, A. C. da, Nascimento, F. C.
P. & Furtado-Neto, M. A. A. 2011. Composição e
abundância de elasmobrânquios capturados com
espinhel de fundo no litoral oeste do estado do Ceará.
Arquivos de Ciências do Mar, 44(2), 5–11. DOI: https://
doi.org/10.32360/acmar.v44i2.157
Jucá-Queiroz, B. 2002. Personal communication.
Klimley, A. P., Le Boeuf, B. J., Cantara, K. M., Richert, J.
E., Davis, S. F., Van Sommeran, S. & Kelly, J. T. 2001.
The hunting strategy of white sharks (Carcharodon
carcharias) near a seal colony. Marine Biology, 138(3),
–636. DOI: https://doi.org/10.1007/s002270000489
Klimley, P. A. & Ainley, D. G. 1996. Great White Sharks:
The Biology of Carcharodon Carcharias. Cambridge:
Academic Press.
Kotas, J. E., Barreto, R., Santos, R. A., Lessa, R., Rosa,
R. S., Vizuete E. P., Baggio M., Salge, P. G., Tavares,
F. E. & Gadig, O. B. F. 2023. Capítulo 3. Plano de
Ação Nacional para Conservação dos Tubarões e
Raias Marinhos Ameaçados de Extinção. In: Kotas, J.
E., Barreto, R., Santos, R. A., Lessa, R., Rosa, R. S.,
Vizuete Eloisa P Baggio Maya, Salge, P. G., Tavares,
F. E. & Gadig, O. B. F. (Org.). PAN Tubarões: Primeiro
Ciclo do Plano de Ação Nacional para a Conservação
dos Tubarões e Raias Marinhos Ameaçados de
Extinção (pp. 88-118). Brasília, DF: ICMBio.
Lanfear, R., Calcott, B., Ho, S. Y. W. & Guindon, S. 2012.
Partitionfinder: combined selection of partitioning
schemes and substitution models for phylogenetic
analyses. Molecular Biology and Evolution, 29(6), 1695–
DOI: https://doi.org/10.1093/molbev/mss020
Leduc, A. O. H. C., De Carvalho, F. H. D., Hussey, N. E.,
Reis-Filho, J. A., Longo, G. O. & Lopes, P. F. M. 2021.
Local ecological knowledge to assist conservation
status assessments in data poor contexts: a case study
with the threatened sharks of the Brazilian Northeast.
Biodiversity and Conservation, 30(3), 819–845. DOI:
https://doi.org/10.1007/s10531-021-02119-5
Leigh, J. W. & Bryant, D. 2015. POPART: Full‐feature
software for haplotype network construction. Methods in
Ecology and Evolution, 6(9), 1110–1116. DOI: https://
doi.org/10.1111/2041-210X.12410
Marra, N. J., Wang, M., Sun, Q., Pavinski Bitar, P. D.,
Stanhope, M. J. & Shivji, M. S. 2016. Mitochondrial
genome of an Atlantic white shark (Carcharodon
carcharias). Mitochondrial DNA Part B, 1(1), 717–719.
DOI: https://doi.org/10.1080/23802359.2016.1222248
Mehlrose, M. R., Bernard, A. M., Finnegan, K. A., Krausfeldt,
L. E., Lopez, J. V. & Shivji, M. S. 2022. Three complete
mitochondrial genomes of shortfin mako sharks, Isurus
oxyrinchus, from the Atlantic and Pacific Oceans.
Mitochondrial DNA Part B, 7(4), 652–654.
O’Leary, S. J., Feldheim, K. A., Fields, A. T., Natanson, L. J.,
Wintner, S., Hussey, N., Shivji, M. S. & Chapman, D. D.
Genetic Diversity of White Sharks, Carcharodon
carcharias, in the Northwest Atlantic and Southern
Africa. The Journal of Heredity, 106(3), 258–265. DOI:
https://doi.org/10.1093/jhered/esv001
Petean, F. F., Naylor, G. J. P. & Lima, S. M. Q. 2020.
Integrative taxonomy identifies a new stingray species
of the genus Hypanus Rafinesque, 1818 (Dasyatidae,
Myliobatiformes), from the Tropical Southwestern
Atlantic. Journal of Fish Biology, 97(4), 1120–1142.
DOI: https://doi.org/10.1111/jfb.14483
Rigby, C. L., Barreto, R., Carlson, J., Fernando, D.,
Fordham, S., Francis, M. P., Herman, K., Jabado, R.
W., Jones, G. C. A., Liu, K. M., Lowe, C. G., Marshall,
A., Pacoureau, N., Romanov, E., Sherley, R. B. &
Winker, H. 2022. Carcharodon carcharias. IUCN Red
List of Threatened Species. IUCN. DOI: https://doi.
org/10.2305/iucn.uk.2022-1.rlts.t3855a212629880.en
Rigby, C. L., Barreto, R., Carlson, J., Fernando, D.,
Fordham, S., Francis, M. P., Jabado, R. W., Liu, K. M.,
Marshall, A., Pacoureau, N., Romanov, E., Sherley, R.
B. & Winker, H. 2018. Isurus oxyrinchus. IUCN Red
List of Threatened Species. IUCN. DOI: https://doi.
org/10.2305/iucn.uk.2019-1.rlts.t39341a2903170.en
Rodrigues Filho, L. F. S., Feitosa, L. M., Silva Nunes, J. L.,
Onodera Palmeira, A. R., Martins, A. P. B., Giarrizzo,
T., Carvalho-Costa, L. F., Monteiro, I. L. P., Gemaque,
R., Gomes, F., Souza, R. F. C., Sampaio, I. & Sales,
J. B. de L. 2020. Molecular identification of ray species
traded along the Brazilian Amazon coast. Fisheries
Research, 223, 105407. DOI: https://doi.org/10.1016/j.
fishres.2019.105407
Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D. L.,
Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M.
A. & Huelsenbeck, J. P. 2012. MrBayes 3.2: efficient
Bayesian phylogenetic inference and model choice
across a large model space. Systematic Biology, 61(3),
–542. DOI: https://doi.org/10.1093/sysbio/sys029
Sales, M. A. N., Freitas, J. E. P. de, Cavalcante, C. C.,
Santander-Neto, J., Charvet, P. & Faria, V. V. 2020. The
southernmost record and an update of the geographical
range of the Atlantic chupare, Styracura schmardae
(Chondrichthyes: Myliobatiformes). Journal of Fish
Biology, 97(1), 302–308. DOI: https://doi.org/10.1111/
jfb.14361
Santander-Neto, J. & Faria, V. V. 2020. Sharks and rays
caught by a small‐scale fisheries in the western
equatorial Atlantic. Journal of Applied Ichthyology,
(6), 830–833. DOI: https://doi.org/10.1111/jai.14103
Santander-Neto, J., Faria, V. V., Castro, A. L. F. & Burgess,
G. H. 2011. New record of the rare ragged-tooth shark,
Odontaspis ferox (Chondrichthyes: Odontaspidae) from
the south-west Atlantic identified using DNA bar coding.
Marine Biodiversity Records, 4, e75.
Santos, C. C., Domingo, A., Carlson, J., Natanson, L. J.,
Travassos, P., Macías, D., Cortés, E., Miller, P., Hazin, F.,
Mas, F., Ortiz de Urbina, J., Lino, P. G. & Coelho, R. 2021.
Movements, habitat use, and diving behavior of shortfin
Mako in the Atlantic Ocean. Frontiers in Marine Science,
DOI: https://doi.org/10.3389/fmars.2021.686343
Sellheim, N. 2020. The CITES appendix II-Listing of
mako sharks — Revisiting counter arguments. Marine
Policy, 115, 103887. DOI: https://doi.org/10.1016/j.
marpol.2020.103887
Seminara, C. I., Souza, G. B. G., Zapelini, C., BarbosaFilho, M. L. V. & Profice, C. C. 2024. Opportunities and
Lamnids in the Equatorial Southwestern Atlantic
Ocean and Coastal Research 2024, v72:e24075 11
Gonçalves-Neto et al.
challenges for small-scale fisheries co-management: A
case study of the unofficial landing records system in a
Brazilian fishing community. Marine Policy, 160, 105991.
DOI: https://doi.org/10.1016/j.marpol.2023.105991
Silva, M., Araujo, M., Geber, F., Medeiros, C., Araujo, J.,
Noriega, C. & Costa da Silva, A. 2021. Ocean dynamics
and topographic upwelling around the Aracati seamount
- north Brazilian chain from in situ observations and
modeling results. Frontiers in Marine Science, 8. DOI:
https://doi.org/10.3389/fmars.2021.609113
Soto, J. M. R. 2001. Annotated systematic checklist and
bibliography of the coastal and oceanic fauna of Brazil.
I. Sharks. Mare Magnum, 1(1), 51–120.
Spaet, J. L. Y., Manica, A., Brand, C. P., Gallen, C. &
Butcher, P. A. 2020. Environmental conditions are
poor predictors of immature white shark Carcharodon
carcharias occurrences on coastal beaches of eastern
Australia. Marine Ecology Progress Series, 653, 167–
DOI: https://doi.org/10.3354/meps13488
Stamatakis, A. 2006. RAxML-VI-HPC: maximum likelihoodbased phylogenetic analyses with thousands of taxa
and mixed models. Bioinformatics, 22(21), 2688–2690.
DOI: https://doi.org/10.1093/bioinformatics/btl446
Stoner, D. S., Grady, J. M., Priede, K. A. & Quattro, J.
M. 2003. Amplification primers for the mitochondrial
control region and sixth intron of the nuclear-encoded
lactate dehydrogenase A gene in elasmobranch fishes.
Conservation Genetics , 4, 805–808.
Westbrook, V., Collin, S., Crawford, D. & Nicholls, M. 2019.
Sharks in the Arts: From Feared to Revered. London,
Routledge.
Whitenack, L. B., Mickley, B. L., Saltzman, J., Kajiura, S.
M., Macdonald, C. C. & Shiffman, D. S. 2022. A content
analysis of 32 years of Shark Week documentaries.
PLoS ONE, 17(11): e0256842. DOI: https://doi.
org/10.1371/journal.pone.0256842
Wosnick, N., Awruch, C. A., Adams, K. R., Gutierre, S. M.
M., Bornatowski, H., Prado, A. C. & Freire, C. A. 2019.
Impacts of fisheries on elasmobranch reproduction: high
rates of abortion and subsequent maternal mortality in
the shortnose guitarfish. Animal Conservation, 22(2),
–206.
