MARINE AND FISHERY SCIENCES 35 (2): 299-306 (2022)
https://doi.org/10.47193/mafis.3522022010502
ABSTRACT. Species-level identification of commercially landed fish provides pivotal informa-
tion for stock assessment and fishery management. However, there is a common lack of species
determination in landing records from small-scale fisheries (SSFs) worldwide. Using DNA barcod-
ing analyses, we detected four overlooked bony fish (yellow snapper, union snook, blackspot
wrasse, and steeplined drum) and one shark species (the sicklefin smooth-hound) in official landing
records of SSFs from northern Peru. Of particular concern is the sicklefin smooth-hound shark
Mustelus lunulatus that was found to be overlooked and could mistakenly be landed as the hump-
back smooth-hound M. whitneyi. Increased efforts should be made to improve species identification
capacities in Peruvian fishing landings. There is an urgent need to quantify the catch levels of mem-
bers of the genus Mustelus to species level. This would contribute to a better understanding of the
levels of exploitation in each particular species and to improved management decisions.
Key words: Smooth-hound, mitochondrial DNA, marine diversity, shark conservation, Mustelus.
Código de barras de ADN revela tiburón y peces óseos no identificados en las estadísticas de
desembarque de las pesquerías artesanales del norte del Perú
RESUMEN. La identificación a nivel de especie de los peces desembarcados comercialmente
proporciona información fundamental para la evaluación poblacional y la ordenación pesquera. Sin
embargo, es común tener dificultades para determinar la identidad de algunas de las especies de las
pesquerías artesanales (SSF) en todo el mundo. Utilizando el análisis de códigos de barras de ADN,
se detectaron cuatro especies de peces óseos (pargo amarillo, robalito, doncella y bereche) y una
especie de tiburón (tollo) pasados por alto en los registros oficiales de desembarque de SSF del norte
de Perú. Particularmente preocupante es la ocurrencia del tollo Mustelus lunulatus, que podría estar
desembarcándose por error como tollo común M. whitneyi. Deben realizarse mayores esfuerzos para
mejorar las capacidades de identificación de algunas de las especies de los desembarques pesqueros
de Perú. Existe una urgente necesidad de cuantificar los niveles de captura de miembros del género
Mustelus a nivel de especie. Esto contribuiría a una mejor comprensión de los niveles de explotación
de cada especie en particular y a la mejora de las decisiones de gestión.
Palabras clave: Tollo, ADN mitocondrial, diversidad marina, conservación de tiburones, Mustelus.
299
*Correspondence:
marin@fish.hokudai.ac.jp
Received: 29 September 2021
Accepted: 17 November 2021
ISSN 2683-7595 (print)
ISSN 2683-7951 (online)
https://ojs.inidep.edu.ar
Journal of the Instituto Nacional de
Investigación y Desarrollo Pesquero
(INIDEP)
This work is licensed under a Creative
Commons Attribution-
NonCommercial-ShareAlike 4.0
International License
Marine and
Fishery Sciences
MAFIS
NOTE
DNA barcoding reveals overlooked shark and bony fish species in landing
reports of small-scale fisheries from northern Peru
ALAN MARÍN1, *, RENATO GOZZER WUEST2, JORGE GRILLO-NUÑEZ2, IRINA B. ALVAREZ-JAQUE1and
JUAN CARLOS RIVEROS3
1Laboratorio de Genética, Fisiología y Reproducción, Facultad de Ciencias, Universidad Nacional del Santa, Av. Pacífico 508, Chimbote,
Perú. 2REDES - Sostenibilidad Pesquera, Calle Porta Nº 130, of. 608, 15074 - Lima, Perú. 3OCEANA - Perú, Av. del Ejército 250, of. 302,
Lima, Perú. ORCID Alan Marín https://orcid.org/0000-0001-7842-133X, Renato Gozzer Wuest https://orcid.org/0000-0002-1156-4552,
Jorge Grillo-Nuñez https://orcid.org/0000-0002-7501-6811, Irina Alvarez-Jaque https://orcid.org/0000-0001-5061-6485,
Juan Carlos Riveros https://orcid.org/0000-0003-4051-181X
Accurate species identification at landing
points is pivotal to assessing exploited stocks’
diversity to improve conservation planning and
management. Information about catch estimates
and species composition allows managers to
determine a fishery’s status. Small-scale fisheries
(SSFs) play a crucial role in producing fishing
landing data through fishing logbooks or landing
declarations. However, management in various
SSFs remains underserved and is characterized
by a common lack of species determination in
landing records, affecting the accuracy of official
fish catch and stock estimates (Morgan and
Burgess 2005).
In Peru, estimated total landings of SSF during
2018 were around 1.1 million tons (De la Puente
et al. 2020). However, despite its social and eco-
nomic importance, most Peruvian SSFs have
weak management systems (De la Puente et al.
2020; Gozzer-Wuest et al. 2021), limited records
and low taxonomic resolution on fisheries land-
ings (Velez-Zuazo et al. 2015; Amorós et al.
2017). Landing data from Peruvian SSFs is gath-
ered by the Instituto del Mar del Perú (IMARPE)
through a network of field observers (Guevara-
Carrasco and Bertrand 2017). They are tasked
with the complex assignment of identifying sev-
eral aquatic species, including highly similar con-
generic species or individuals that have lost their
distinctive morphological characteristics due to
cellular and biochemical changes associated with
postmortem (Ocaño-Higuera et al. 2009). Species
identification of shark landings can be even more
challenging because they are often landed head-
less and finless; albeit shark finning has been
banned since 2016 (Supreme Decree Nº 021-
2016 PRODUCE, El Peruano 2016).
The Peruvian sea is home to at least 67 shark
species (Cornejo et al. 2015; Kelez et al. 2020),
half of which interact with Peruvian SSFs (Gon-
zalez-Pestana et al. 2016). Currently, six
Mustelus species occur in Peruvian waters: M.
californicus, M. dorsalis, M. henlei, M. lunula-
tus, M. mento, and M. whitneyi (Cornejo et al.
2015). In Peru, the humpback smooth-hound M.
whitneyi is listed among the top six most landed
shark species (Gonzalez-Pestana et al. 2016),
representing the only member of Mustelus genus
reported to species-level in recent landing
records from northern Peru (i.e., the regions of
Piura and Tumbes). Landing records from the
same regions reported two other Mustelus
species (M. mento and M. dorsalis) to species
level only during the mid-1990s. On the other
hand, the sicklefin smooth-hound M.lunulatus
has never been reported to species-level in Peru-
vian landing records, despite being listed as a
commercial species by IMARPE in the ’90s
decade (Elliot-Rodriguez et al. 1996). Further-
more, two independent molecular studies found
M. lunulatus in retail markets and supermarkets
from Lima and Tumbes (Marín et al. 2018; Biffi
et al. 2020). Mustelus species are traded in
domestic markets simply as tollos (López de la
Lama et al. 2018). They often look similar and
display partially overlapping characters within
species, making their identification difficult even
for experts (Morgan and Burgess 2005; Pérez-
Jiménez et al. 2005). Aiming to reveal economi-
cally important species that could be overlooked
in official landing reports, a DNA barcoding
approach for the authentication of fish landings
of SSFs from northern Peru was used. The results
of genetic identification were contrasted against
official landing records from Tumbes and Piura
obtained over the past three decades.
A total of 95 fish samples were collected in
twelve landing sites from northern Peru, includ-
ing the regions of Piura (n = 8) and Tumbes (n =
4) (Figure 1; supplementary material, Table S1)
from May to August 2019. Fin clip samples were
preserved in ethanol 96% at the moment of land-
ing and stored at -15 °C until DNA extraction.
Genomic DNA was isolated using the phenol-
chloroform method (Sambrook and Rusell 2001).
We used two mitochondrial markers: cytochrome
c oxidase subunit I (COI) and the control region
(D-Loop); the latter was used only to discriminate
300 MARINE AND FISHERY SCIENCES 35 (2): 299-306 (2022)
tuna species (Pedrosa-Gerasmio et al. 2012)
(primer sequences are shown in supplementary
material, Table S2). PCR products were amplified
in a local private laboratory using the same con-
ditions described in Marín et al. (2018). PCR
products were purified and sequenced at the
Macrogen Inc. sequencing facilities in Korea.
Sequences were manually edited using MEGA 7
(Kumar et al. 2016). For species identification,
both the Identification Engine at the Barcode of
Life Data System and the Basic Local Alignment
Search Tool (BLAST) at the National Center for
Biotechnology Information were used. Species
identification obtained by DNA barcoding analy-
ses was compared with the corresponding scien-
tific and common/market names included in the
IMARPE reports from Tumbes and Piura landing
sites. These reports were issued from 1986 to
1988 (Wosnitza-Mendo et al. 1988), from 1996 to
2000 (Estrella et al. 1998a, 1998b, 1999 and
2000), and from 2010 to 2020 (Sistema de
Captación de Información de la Pesca Artesanal
del IMARPE) and were kindly provided by
IMARPE (IMARPE 2019, 2020, 2021). DNA
sequences obtained in this work were submitted
to the GenBank database under the accession
numbers MN880503 to MN880608.
DNA barcoding results revealed the presence
of 40 fish species belonging to 15 families (sup-
plementary material, Table S3). Among these
species, five were found to be overlooked in the
IMARPE landing reports by location or at least
were not explicitly mentioned in the data provid-
ed by IMARPE (i.e., some species could be
implicitly included in the statistics as aggregated
records such as ‘others’ or registered to the genus
level only). These five species were M. lunulatus,
Centropomus unionensis, Decodon melasma,
Larimus acclivis, and Lutjanus argentiventris
(Table 1; supplementary material, Figure S1).
301
MARÍN: OVERLOOKED FISH SPECIES IN PERUVIAN SMALL-SCALE FISHERIES
Figure 1. Sampling locations from this study. Fish landing sites from Tumbes region: Puerto Pizarro, La Cruz, Caleta Grau, and
Cancas. Fish landing sites from Piura region: Los Órganos, El Ñuro, Cabo Blanco, Paita, Yacila, La Islilla, La Tortuga,
and Bayóvar. The inset picture depicts the landing of Mustelus lunulatus collected at Cancas fish landing site (Tumbes).
500 100 km25
4°
5°
S
6°
81° 80° 79°
7°
Ecuador
Pacific
Ocean
Peru
Tumbes
Piura
82°83°
Tumbes
Piura
Puerto Pizarro
La Cruz
Caleta Grau
Cancas
Los Órganos
El Ñuro
Cabo Blanco
La Islilla
Paita
Yacila
La Tortuga
Bayóvar
Sampling sites
W
Ecuador
Peru
Brazil
Pacific
Ocean
1000 km5002500
Among the total number of smooth-hound shark
samples collected in this study (n = 8), six samples
(75%) were identified as M. whitneyi (100%
sequence identity in BOLD). In comparison, two
samples (25%) were identified as M.lunulatus.
(99.7-99.8% sequence identity in BOLD, Gen-
Bank access MN880526 and MN880605). The
two M. lunulatus samples were collected at Cancas
landing site (Tumbes, collection date June and
August 2019, supplementary material, Figure S1)
and mislabeled as tollo mama (common name for
M. whitneyi). Since this species is not reported in
statistics, its landings could often be misidentified
as M. whitneyi at landing points, or just listed in
the IMARPE annual statistics landing reports
under the category tollo. The latter may include
different species such as M. whitneyi, M. lunula-
tus, Triakis maculata and Schroederichthys chilen-
sis (Flores-Palomino et al. 1994; Flores 1996; Fer-
nández et al. 2000). Additionally, the Peruvian
Ministry of Production, which compiles official
statistics, considers tollo as one single species
despite being a common name for several species,
as previously mentioned (PRODUCE, 2020).
A previous study by Marín et al. (2018) report-
ed this species’ incidence in a fillet sample bought
in a supermarket in Lima (collection date June
2017). Similarly, using the DNA barcode
approach, Biffi et al. (2020) identified four M.
lunulatus samples (two came from retail markets
in Tumbes and two from supermarkets in Lima)
collected between May and June 2017. Before
our study, there were no records of M. lunulatus
collected from Peruvian landings and authenticat-
ed by DNA analysis. Together, these identifica-
tions support the hypothesis of regular catches of
this species in northern Peru.
Globally, there is a general lack of data report-
ing on sharks’ catch particularly species-specific
data, which makes fisheries conservation and
management challenging (NOAA 2020). Further-
more, most life history patterns of shark species
(including Mustelus species) display slow
growth, long gestation times, low fecundity, and
late sexual maturity (Medina-Morales et al.
2020), which makes them especially vulnerable
to exploitation. Therefore, there is an urgent need
to quantify catch levels of members of the genus
302 MARINE AND FISHERY SCIENCES 35 (2): 299-306 (2022)
Table 1. List of overlooked species from the SSF in northern Peru as identified through DNA barcoding.
Species match Common Family Label Sampling Sampling n GenBank ID
(BOLD/NCBI) name site date accession
Mustelus Sicklefin Triakidae Tollo mama Cancas, Tumbes June 25, 2019 2 MN880526
lunulatus smooth-hound
August 1, 2019 MN880605
Centropomus Union snook Centropomidae Robalo Cancas, Tumbes June 27, 2019 1 MN880528
unionensis plateado
Decodon Blackspot Labridae San pedrano La Islilla, Piura May 23, 2019 1 MN880543
melasma wrasse
Lutjanus Yellow Lutjanidae Paramo rojo Cancas, Tumbes June 25, 2019 2 MN880525
argentiventris snapper
Paramo muelon July 30, 2019 MN880602
Larimus Steeplined Sciaenidae Bereche La Tortuga, Piura June 11, 2019 1 MN880556
acclivis drum
Mustelus to species level. This would contribute
to a better understanding of the exploitation level
in each particular species, so as to improve man-
agement decisions (Pérez-Jiménez et al. 2016).
Misidentifications of other Mustelus species,
such as M. whitneyi, also raise concerns over its
catch landing estimates, which may be biased by
the landings of M. dorsalis and M. mento (report-
ed to species level only during the years 1996,
1997, and 1999).
Further studies are needed to determine to
what extent Peruvian landings of M. lunulatus
(or other Mustelus species) are misidentified as
M. whitneyi. Additionally, the IUCN Red List of
Threatened Species listed M. whitneyi and M.
lunulatus as critically endangered and least con-
cern, respectively. Inaccurate species identifica-
tions may also have an impact on global IUCN
listings. In this regard, a molecular survey of
Mustelus landings could be a good starting point
to obtain accurate baseline data. However, more
substantial efforts must also be made to find
more practical and cheaper species identifica-
tion solutions. For instance, the possibility of
developing field identification techniques based
on diagnostic morphometric measurements is
required, which should be developed alongside
DNA barcoding to obtain more voucher speci-
mens of Mustelus species. Additionally, multi-
variate and machine learning methods success-
fully applied in Carcharhinidae shark species
should also be evaluated, as this approach could
be applied to other Chondrichthyan species
(Johnson et al. 2017). It is also highly recom-
mended that field observers receive more
intense training by recognized experts on identi-
fication of the Mustelus species. There are two
identification guides for commercially impor-
tant Peruvian shark species (IMARPE 2015;
Romero et al. 2015), but none of these include
M. lunulatus.
Our results also detected four landed Oste-
ichthyes species that were not included in land-
ing reports. Two yellow snappers Lutjanus
argentiventris (supplementary material, Figure
S2) were sampled at Cancas landing site
(Tumbes, collection date June and July 2019),
and one was misidentified as spotted rose snap-
per L. guttatus by fishermen. Official landing
records only include two species of genus Lut-
janus: L. guttatus and L. jordani. In another case,
a sample taken in Cancas (Tumbes, collection
date June 2019) landed as white snook Centropo-
mus viridis was actually the union snook Cen-
tropomus unionensis (supplementary material,
Figure S3). Official landing records include only
C. nigrescens and aggregated records to genus
level of Centropomus spp. One individual identi-
fied as the blackspot wrasse Decodon melasma
(supplementary material, Figure S4) and one
sample of the steeplined drum Larimus acclivis
(supplementary material, Figure S5) collected in
La Islilla and La Tortuga landing sites (Piura),
respectively, were found to be overlooked in the
official landing reports. Three other overlooked
species detected in this study (C. unionensis, L.
acclivis, and L. argentiventris) have been
described as part of the marine diversity of
Tumbes (Luque 2007), whereas the blackspot
wrasse D. melasma has been reported in a survey
cruise by IMARPE (Pastor et al. 2018). Never-
theless, there is still a lack of knowledge about
the population status and landing estimates of
these species. Further studies are needed to
determine whether catches are significant or neg-
ligible from the point of view of the impacts of
fishing on the health of the stocks.
Finally, this note emphasizes the usefulness
and importance of DNA barcoding analyses in
detecting species that may have been overlooked
in official statistics. Accordingly, we recommend
that proper attention should be paid to these
events. More specialized training on how to iden-
tify morphologically similar species should be
given to field observers and fishermen, which
would significantly improve landing data and,
consequently, could trigger future management
actions to preserve Peruvian marine diversity.
303
MARÍN: OVERLOOKED FISH SPECIES IN PERUVIAN SMALL-SCALE FISHERIES
ACKNOWLEDGEMENTS
We thank Oceana-Peru for funding this
research. We also thank Carlos Gutierrez, Alejan-
dra Mendoza and Frank Altamirano for fish sam-
pling and data collection and Biodes Laboratorios
Soluciones Integrales for their support during the
molecular analyses.
REFERENCES
AMORÓS S, GOZZER R, MELGAR, V, ROVEGNO N.
2017. Peruvian mahi mahi fishery (Coryphae-
na hippurus): characterization and analysis of
the supply chain. WWF (Marine Program of
WWF) Peru. http://d2ouvy59p0dg6k.cloud-
front.net/downloads/mahi_mahi_value_chai_
en.pdf.
BIFFI D, LÓPEZ-MOBILIA A, KELEZ S, WILLIAMS
DA, CHUMCHAL MM, WEINBURGH M. 2020.
Mislabelling and high mercury content ham-
pers the efforts of market-based seafood initia-
tives in Peru. Sci Rep. 10 (1): 1-12.
CORNEJO R, VÉLEZ-ZUAZO X, GONZÁLEZ-PESTA-
NA A, KOURI C, MUCIENTES, G. 2015. An
updated checklist of Chondrichthyes from the
southeast Pacific off Peru. Check List. 11 (6):
1809.
DELAPUENTE S, LÓPEZ DELALAMA R, BENA-
VENTE S, SUEIRO JC, PAULY D. 2020. Growing
into poverty: Reconstructing Peruvian small-
scale fishing effort between 1950 and
2018. Front Mar Sci. 7, 681.
ELPERUANO. 2016. Decreto supremo que estable-
ce medidas de ordenamiento para la pesquería
del recurso tiburón. 2 de noviembre del 2016.
Decreto Supremo Nº 021-2016-PRODUCE;
[accessed 2021 September 16]. https://
elperuano.pe/normaselperuano/2016/11/02/14
48564-3.html.
ELLIOTT-RODRÍGUEZ W, PAREDES-BULNES, F,
ZEBALLOS-FLOR J, JUÁREZ L, BARRETO-MEZA
J. 1996. Nomenclatura actualizada de peces
comerciales del Perú. Informe Progresivo Nº
37. Inf Inst Mar Perú. 12 p.
ESTRELLA-ARELLANO C, GUEVARA-CARRASCO R.
1998a. Informe Nº 131. Informe estadístico
anual de los recursos hidrobiológicos de la
pesca artesanal por especies, artes, caletas y
meses durante 1996. Inf Inst Mar Perú. p. 215.
ESTRELLA-ARELLANO C, GUEVARA-CARRASCO R.
1998b. Informe Nº 132. Informe estadístico
anual de los recursos hidrobiológicos de la
pesca artesanal por especies, artes, caletas y
meses durante 1997. Inf Inst Mar Perú. p. 412.
ESTRELLA-ARELLANO C, GUEVARA-CARRASCO R,
AVILA-PÉREZ W, PALACIOS-LEÓN J, MEDINA-
CRUZ A. 2000. Informe Nº 151. Informe esta-
dístico de los recursos hidrobiológicos de la
pesca artesanal por especies, artes, meses y
caletas durante el segundo semestre de 1999.
Inf Inst Mar Perú. 151: 1-194.
ESTRELLA-ARELLANO C, GUEVARA-CARRASCO R,
PALACIOS-LEÓN J, AVILA-PÉREZ W, MEDINA-
CRUZ A. 1999. Informe Nº 148. Informe esta-
dístico de los recursos hidrobiológicos de la
pesca artesanal por especies, artes, meses y
caletas durante el primer semestre de 1999. Inf
Inst Mar Perú. p. 214.
[FAO] FOOD AND AGRICULTURE ORGANIZATION OF
THE UNITED NATIONS. 2020. Illuminating Hid-
den harvests study in the spotlight at COFI
virtual dialogues. Voluntary guidelines for
securing sustainable small-scale fisheries in
the context of food security and poverty erad-
ication online 17/07/2020; [accessed 2021
September 16]. http://www.fao.org/voluntary-
guidelines-small-scale-fisheries/news-and-
events/detail/en/c/1300712/
FERNÁNDEZ-RAMÍREZ F, VERA-DÍAZ S, MARCELO-
PADILLA R, CHIRINOS E. 2000. Estadísticas de
los desembarques de la pesquería marina
peruana 1999. Inf Inst Mar Perú. 115.
FLORES M, VERA S, MARCELO R, CHIRINOS E.
304 MARINE AND FISHERY SCIENCES 35 (2): 299-306 (2022)
1996. Estadísticas de los desembarques de la
pesquería marina peruana durante 1995. Inf
Prog Inst Mar Perú. 45. 35 p.
FLORES-PALOMINO M, VERA-DÍAZ S, MARCELO-
PADILLA R, CHIRINOS E. 1994. Estadísticas de
los desembarques de la pesquería marina
Peruana 1983-1992. Inf Inst Mar Perú. 105.
202 p.
GONZALEZ-PESTANA A, KOURI C, VELEZ-ZUAZO X.
2016. Shark fisheries in the Southeast Pacific:
a 61-year analysis from Peru. F1000Res. 3.
GOZZER-WUEST R, ALONSO-POBLACIÓN E, TIN-
GLEY G. 2021. Identifying priority areas for
improvement in Peruvian Fisheries. Mar
Policy. 129, 104545.
GUEVARA-CARRASCO R, BERTRAND A, editors.
2017. Atlas de la pesca artesanal del mar del
Perú. Lima: Edición IMARPE-IRD. 183 p.
[IMARPE] INSTITUTO DEL MAR DEL PERÚ. 2015.
Guía para la determinación de tiburones de
importancia comercial en el Perú. Serie de
divulgación científica. 1 (2). 80 p.
[IMARPE] INSTITUTO DEL MAR DEL PERÚ. 2019.
Respuesta a la solicitud de acceso a la infor-
mación pública: información sobre estimados
de desembarque de los principales recursos
hidrobiológicos registrados por la pesquería
artesanal en las caletas de Tumbes y Piura,
durante los años 2014-2018. Carta Nº 073
2019-IMARPE/OGA. Inst Mar Perú. 18 p.
[IMARPE] INSTITUTO DEL MAR DEL PERÚ. 2020.
Respuesta a la solicitud de acceso a la informa-
ción pública: información sobre estimados de
desembarque de los principales recursos hidro-
biológicos registrados por la pesquería artesa-
nal en las caletas de Tumbes y Piura, durante
los años 2010-2020. Oficios Nº 017, 018 y 019-
2020-IMARPE/OGA. Inst. Mar Perú. 18 p.
[IMARPE] INSTITUTO DEL MAR DEL PERÚ. 2021.
Respuesta a la solicitud de acceso a la infor-
mación pública: información sobre estimados
de desembarque mensuales 2010-2020 en
Piura y Tumbes de: Centropomus unionensis,
Decodon melasma, Lutjanus argentiventris,
Caulolatilus princeps, Pareques lanfeari,
Larimus acclivis, Umbrina santi, Prionace
glauca, Epinephelus analogus, Brotula ord-
wayi y Mustelus lunulatus. Oficios Nº 23 y 24-
2021-IMARPE/OGA. Inst Mar Perú. 8 p.
JOHNSON GJ, BUCKWORTH RC, LEE H, MORGAN
JAT, OVENDEN JR, MCMAHON CR. 2017. A
novel field method to distinguish between
cryptic carcharhinid sharks, Australian black-
tip shark Carcharhinus tilstoni and common
blacktip shark C. limbatus, despite the pres-
ence of hybrids. J Fish Biol. 90 (1): 39-60.
KELEZ S, NAPURI RM, PFENNIG AM, MARTINEZ
OC, CARRASCO AT. 2020. First reports of
Megamouth Shark, Megachasma pelagios
Taylor, Compagno & Struhsaker, 1983 (Lam-
niformes, Megachasmidae), in Peru. Check
List. 16: 1361.
KUMAR S, STECHER G, TAMURA K. 2016.
MEGA7: molecular evolutionary genetics
analysis version 7.0 for bigger datasets. Mol
Biol Evol. 33 (7): 1870-1874.
LÓPEZ DE LA LAMA R, DELAPUENTE S, RIVEROS
JC. 2018. Attitudes and misconceptions
towards sharks and shark meat consumption
along the Peruvian coast. PLoS ONE. 13 (8):
e0202971.
LUQUE C. 2007. Estudio de la diversidad hidro-
biologica en Tumbes. Informe Anual 2007.
Tumbes: Inst Mar Perú. 36 p.
MARÍN A, SERNA J, ROBLES C, RAMÍREZ B,
REYES-FLORES LE, ZELADA-MÁZMELA E,
SOTIL G, ALFARO R. 2018. A glimpse into the
genetic diversity of the Peruvian seafood sec-
tor: Unveiling species substitution, mislabel-
ing and trade of threatened species. PLoS
ONE. 13 (11): e0206596.
MEDINA-MORALES SA, CORRO-ESPINOSA D,
ESCOBAR-SÁNCHEZ O, DELGADO-ALVAREZ CG,
RUELAS-INZUNZA J, FRÍAS-ESPERICUETA MG,
et al. 2020. Mercury (Hg) and selenium (Se)
content in the shark Mustelus henlei (Triaki-
dae) in the northern Mexican Pacific. Environ
Sci Pollut Res. 27 (14): 16774-16783.
305
MARÍN: OVERLOOKED FISH SPECIES IN PERUVIAN SMALL-SCALE FISHERIES
MORGAN A, BURGESS GH. 2005. Fishery depend-
ent sampling: total catch, effort and catch
composition. In: MUSICK JA, BONFIL R, edi-
tors. Elasmobranch fisheries management
techniques. FAO Tech Rep. 454. p. 182-200.
[NOAA] NATIONAL OCEANIC AND ATMOSPHERIC
ADMINISTRATION. 2020. International affairs.
Shark conservation. https://www.fisheries.
noaa.gov/national/international-affairs/shark-
conservation.
OCAÑO-HIGUERA VM, MARQUEZ-RÍOS E, CANIZA-
LES-DÁVILA M, CASTILLO-YÁÑEZ FJ, PACHE-
CO-AGUILAR R, LUGO-SÁNCHEZ ME, et al.
2009. Postmortem changes in cazon fish mus-
cle stored on ice. Food Chem. 116 (4): 933-
938.
PASTOR R, ZAVALAGA F, PALACIOS J. 2018. Carac-
terización de la comunidad bentodemersal que
cohabita con la merluza Merluccius gayi
peruanus Ginsburg entre la frontera norte del
dominio marítimo peruano y Huarmey. 2015.
Inf Inst Mar Perú. 45 (1): 104-121.
PEDROSA-GERASMIO IR, BABARAN RP, SANTOS
MD. 2012. Discrimination of juvenile yel-
lowfin (Thunnus albacares) and bigeye (T.
obesus) tunas using mitochondrial DNA con-
trol region and liver morphology. PLoS
ONE. 7 (4): e35604.
PÉREZ-JIMÉNEZ JC, NISHIZAKI OS, CASTILLO-
GENIZ JL. 2005. A new eastern North Pacific
smooth-hound shark (genus Mustelus, family
Triakidae) from the Gulf of California.
Copeia. (4): 834-845.
PÉREZ-JIMÉNEZ J, WHITE CF, RUIZ C, CARLISLE
AB, LOWE CG. 2016. Mustelus lunulatus. The
IUCN Red List of Threatened Species 2016:
e.T161640A80672480.
[PRODUCE] MINISTERIO DE LA PRODUCCIÓN.
2020. Anuario estadístico pesquero y acuícola
2019. Lima: Ministerio de la Producción. 181
p.
ROMERO MA, ALCÁNTARA PF, VERDE K, editors.
2015. Guía de campo para la determinación de
tiburones en la pesca artesanal del Perú. Lima:
Inst Mar Perú. 15 p.
SAMBROOK J, RUSELL D. 2001. Molecular
cloning: a laboratory manual. 3rd ed. New
York: Cold Spring Harbor.
VELEZ-ZUAZO X, ALFARO-SHIGUETO J, MANGEL J,
PAPA R, AGNARSSON I. 2015. What barcode
sequencing reveals about the shark fishery in
Peru. Fish Res. 161: 34-41.
WOSNITZA-MENDO C, ESPINO M, VÉLIZ M. 1988.
La pesquería artesanal en el Perú durante junio
de 1986 a junio de 1988. Inf Inst Mar Perú. 93.
142 p.
306 MARINE AND FISHERY SCIENCES 35 (2): 299-306 (2022)
