International Fisheries Threaten Globally Endangered

Home , Galapagos shark, Grey reef shark, Oceanic whitetip shark, Tiger shark

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OPEN International fsheries threaten globally endangered sharks in the Eastern Tropical Pacifc Ocean: the case of the Fu Yuan Yu Leng 999 reefer vessel seized within the Galápagos Marine Reserve Elisa Bonaccorso1,2*, Nicté Ordóñez‑Garza2, Diana A. Pazmiño1,2,3, Alex Hearn1,3, Diego Páez‑Rosas1,3, Sebastián Cruz4, Juan Pablo Muñoz‑Pérez3,5, Eduardo Espinoza6, Jenifer Suárez6, Lauren D. Muñoz‑Rosado7, Andrea Vizuete7, Jaime A. Chaves2,3,8, Maria de Lourde Torres1,3, Walter Bustos6, Danny Rueda6, Maximilian Hirschfeld3,9 & Juan M. Guayasamin1,2,3

Shark fshing, driven by the fn trade, is the primary cause of global shark population declines. Here, we present a case study that exemplifes how industrial fsheries are likely depleting shark populations in the Eastern Tropical Pacifc Ocean. In August 2017, the vessel Fu Yuan Yu Leng 999, of Chinese fag, was detained while crossing through the Galápagos Marine Reserve without authorization. This vessel contained 7639 sharks, representing one of the largest seizures recorded to date. Based on a sample of 929 individuals (12%), we found 12 shark species: 9 considered as Vulnerable or higher risk by the IUCN and 8 listed in CITES. Four species showed a higher proportion of immature than mature individuals, whereas size-distribution hints that at least some of the fshing ships associated with the operation may have been using purse-seine gear fshing equipment, which, for some species, goes against international conventions. Our data expose the magnitude of the threat that fshing industries and illegal trade represent to sharks in the Eastern Tropical Pacifc Ocean.

Amid the current biodiversity crisis, an era of unprecedented species extinction 1, our ability to document species loss and ecosystem collapse in marine environments has lagged behind compared with terrestrial ecosystems2. Tis situation is particularly worrisome in the case of sharks, a group with a high proportion of top predators central to marine food web dynamics 3. Some sharks have a high market value and hence are targeted by fsheries for their fns and other products4. Given the economic incentives for exploitation and the difculties involved in controlling this trade, shark fshing has caused substantial population declines 5. Consequently, at least 74 species of sharks (16% of shark diversity) are threatened worldwide6.

1Universidad San Francisco de Quito USFQ, Colegio de Ciencias Biológicas y Ambientales, Quito, Ecuador. 2Laboratorio de Biología Evolutiva, Instituto Biósfera, Universidad San Francisco de Quito, Quito, Ecuador. 3Universidad San Francisco de Quito (USFQ) and UNC-Chapel Hill Galapagos Science Center (GSC) Av. Alsacio Northia, Isla San Cristóbal, Galápagos, Ecuador. 4Independent Researcher, Puerto Ayora, Santa Cruz, Galápagos, Ecuador. 5Faculty of Science and Engineering, University of the Sunshine Coast, Sunshine Coast, QLD, Australia. 6Dirección del Parque Nacional Galápagos, Puerto Ayora, Galápagos, Ecuador. 7Pontifcia Universidad Católica del Ecuador Sede Manabí, Bahía de Caráquez, Manabí, Ecuador. 8Department of Biology, San Francisco State University, 1600 Holloway Ave, San Francisco, CA, USA. 9James Cook University, Townsville, QLD, Australia. *email: [email protected]

Scientifc Reports | (2021) 11:14959 | https://doi.org/10.1038/s41598-021-94126-3 1 Vol.:(0123456789) www.nature.com/scientificreports/

Figure 1. Sample of shark carcasses seized in vessel Fu Yuan Yu Leng 999. A, inside one of the vessel freezers. B, unborn scalloped hammerhead sharks; C, whale shark. We assembled this fgure using Adobe Creative Suite (https://www. adobe. com ).

In response to global threats, the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES) has identifed the international shark fn trade as the primary driver of population declines, protecting 46 shark species under this convention. Further, 26 shark species are listed in Annex 1 of the Memo- randum of Understanding on the Conservation of Migratory Sharks (www.cms.int/sharks/en/legalinstrument/ sharks-mou), the frst global instrument for the conservation of migratory species of elasmobranchs. Tus, management to prevent population collapses of some of the most heavily traded and vulnerable shark species has become a global priority (CITES, https://www.citessharks.org). Moreover, sharks beneft from marine protected areas that preserve habitats and populations 7, and export biomass to non-protected areas, sustaining the yield of nearby fsheries 8, 9. Unfortunately, fsheries management regulations and benefts derived from marine protected areas become inefective when sharks move into international waters10 or are undermined when international or local feets fsh protected species within countries’ exclusive economic zones (EEZs)11, 12. Here, we present a case study from the Eastern Tropical Pacifc Ocean as an example of how the operation of international feets may hamper national and international eforts to preserve shark populations worldwide. Our case is staged in the Galápagos Islands, an archipelago world-renowned for its iconic species and marine environments. Established in 1959, the Galápagos National Park (GNP) expanded to the Galápagos Marine

Scientifc Reports | (2021) 11:14959 | https://doi.org/10.1038/s41598-021-94126-3 2 Vol:.(1234567890) www.nature.com/scientificreports/

Family/species Common name Spanish name Present in GMR Total Frequency Conservation status IUCN CITES Family Alopidae Alopias pelagicus Pelagic thresher Tiburón rabón bueno Yes 122 13.1% Endangered Appendix II Tiburón rabón amargo/ Alopias superciliosus Bigeye thresher Yes 54 5.8% Vulnerable Appendix II Tiburón zorro ojón Family Charcharhinidae Carcharhinus amblyrhynchos Grey reef shark Tiburón gris de arrecife Yes 2 0.2% Endangered No data Carcharhinus falciformis Silky shark Tiburón mico/sedoso Yes 257 27.7% Vulnerable Appendix II Tiburón aletón/ puntas blan- Carcharhinus longimanus Oceanic whitetip shark Yes 188 20.2% Critically endangered Appendix II cas oceánico Galeocerdo cuvier Tiger shark Tiburón tigre Yes 2 0.2% Near threatened No data Prionace glauca Blue shark Tiburón azul o Aguado Yes 109 11.7% Near threatened No data Family Lamnidae Isurus oxyrinchus Shortfn mako Tiburón tinto Yes 4 0.4% Endangered Appendix II Family Rhincodontidae Rhincodon typus Whale shark Tiburón Ballena Yes 1 0.1 Endangered Appendix II Family Sphyrnidae Sphyrna lewini Scalloped hammerhead Cachuda roja Yes 122 13.1% Critically endangered Appendix II Sphyrna zygaena Smooth hammerhead Cachuda blanca Yes 67 7.2% Vulnerable Appendix II Family Triakidae Mustelus mustelus Common smooth-hound Tollo No 1 0.1% Vulnerable No data 929 100%

Table 1. Conservation status of species of sharks from vessel Fu Yuan Yu Leng 999 intercepted within the limits of the Galápagos Marine Reserve. Data from www.iucn.org and www.cites.org, updated to December 2020. All species except Mustelus mustelus have been reported for the Galápagos Marine Reserve (GMR) (Supplementary Information Table S1).

Reserve in 1998 13, a protected area of about 133,000 km2. Tis reserve has aforded over 20 years of protection for endemic and native marine species and holds at least 36 species of sharks (Supplementary Information Table S1). On August 13th, 2017, an alert was issued to the Navy of Ecuador from the Monitoring Center of the GNP Directorate regarding a sizeable vessel crossing the Galápagos Marine Reserve. Te reefer vessel Fu Yuan Yu Leng 999, of Chinese fag, was detained 12 miles north of Punta Pitt, on San Cristóbal Island. Te vessel was escorted to Puerto Baquerizo Moreno, the Galápagos Islands’ political and judicial capital, where an inquiry and subsequent trial began. Here, we combined morphological identifcation with molecular barcoding to assess the identity of 12% of the carcasses contained in the vessel Fu Yuan Yu Leng 999 and estimated the size distribution of sharks targeted. We argue that the high shark biomass and the high proportion of endangered species transported by the reefer vessel, as well as the recent operation of international feets around the Ecuadorian Exclusive Economic Zones, reveal a pattern of sustained exploitation of sharks in this region and, more generally, the Eastern Tropical Pacifc Ocean. We discuss our results in terms of their implications for the impact of international fshing feets on the conservation of shark species. Results Te vessel Fu Yuan Yu Leng 999 was carrying 572 tons of fsh. Te ship contained six freezer holds, and only one and ~ 1/3 were full. Te cargo included 7639 sharks (7207 juveniles or adults, 432 unborn; see examples in Fig. 1A,B), 2114 bony fsh, and 537 bags of shark fns. Out of the 7639 sharks found, 929 juveniles or adults (12% of the cargo) were morphologically identifed and molecularly barcoded (GenBank accession numbers listed in Supplementary Data S1). Tis sample consisted of 12 species of sharks (Table 1), including a single specimen of the whale shark, Rhincodon typus (Fig. 1C).

Shark identifcation and geographic afnities. Out of 929 samples analyzed, 800 (86%) matched the morphological and molecular identifcations (Supplementary Information Table S2). An overview of the phylogenetic tree is shown in Fig. 2. Te complete maximum likelihood tree, with species names and bootstrap supports is available in the Supplementary Information Figure S1, where all samples were located in the same clade as samples of the species assigned by BLAST (bootstrap support ≥ 70%). We could not determine the geographic afnities between our samples and those retrieved from GenBank because of the limited number of sequences per species from few geographic locations on this database. Also, at least 20% of GenBank sequences were not associated with specifc localities and the overall level of individual variation in ND2 was low (see tree in Supplementary Information Figure S1). Still, the 929 sequences generated during this study, 86% of which are backed up by morphological diagnosis, will expand the availability of comparative shark data on GenBank, enhancing species identifcations in future studies.

Scientifc Reports | (2021) 11:14959 | https://doi.org/10.1038/s41598-021-94126-3 3 Vol.:(0123456789) www.nature.com/scientificreports/

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Sharks in the Fu Yuan Yu Leng 999 Species followed by number of individuals in the sample

A.Whale shark (1) B.Pelagic thresher (122) C.Bigeye thresher (54) D. (4) E.Common smooth-hound(1) F. Tiger shark (2) G.Scalloped hammerhead (122) H. Smooth hammerhead (67) I. Oceanic whitetip shark (188) J. Grey reef shark (2 ) K. Blue shark (109) L.Silky shark (257)

257 0

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◂Figure 2. Phylogenetic tree obtained from 929 samples from 12 species of sharks found in vessel Fu Yuan Yu Leng 999, comparative sequences from GenBank (1275 individuals from 229 shark species distributed in the Pacifc Ocean), and 9 Galapagos sharks from the Galápagos Islands. Te fgure also shows the number of identifed carcasses per species found in the vessel (a total of 929 individuals); note that these numbers represent only a fraction of the 7639 sharks (7207 juveniles or adults, 432 unborn) found in the Fu Yuan Yu Leng 999. Photo credits: A, Whale shark (Rhincodon typus), Alex Hearn; B, Pelagic thresher (Alopias pelagicus), licenced by CSIRO; C, Bigeye thresher (Alopias superciliosus), licenced by CSIRO; D, Shortfn mako (Isurus oxyrinchus), licenced by CSIRO; E, Common smooth-hound (Mustelus mustelus), Henri Gervais (1877); F, Tiger shark (Galeocerdo cuvier), licenced by CSIRO; G, Scalloped hammerhead (Sphyrna lewini), Henri Gervais (1877); H, Smooth hammerhead (Sphyrna zygaena), Francys Day (1878); I, Oceanic whitetip shark (Carcharhinus longimanus), licenced by CSIRO; J, Grey reef shark (Carcharhinus amblyrhynchos), ReefLifeApps.com, licensed by Creative Commons Attribution-Share Alike 3.0; K, Blue shark (Prionace glauca), licenced by CSIRO; L, Silky shark (Carcharhinus falciformis), CSIRO. We assembled this fgure using FigTree 1.4.4 (http://tree. bio. ed. ac. uk/ sofware/ fgtr ee/ ) and Adobe Creative Suite (https://www. adobe. com ).

Conservation status of seized sharks. From the 12 species of sharks found in the vessel Fu Yuan Yu Leng 999, 11 are found within the Galápagos Marine Reserve, 9 are listed as Vulnerable or higher risk by the IUCN (https://www.iucnredlist.org), and 8 are listed as CITES species (Appendix II; http://checklist.cites.org) (Table 1). Te species most represented in the cargo were silky shark (Vulnerable; 28%), oceanic whitetip shark (Critically Endangered; 20%), scalloped hammerhead (Sphyrna lewini; Critically Endangered; 13%), pelagic thresher (Endangered; 13%), and blue shark (Near Treatened; 12%).

Size and estimation of size of sexual maturity. We obtained pre-caudal length measurements for 745 individuals of 11 species (Supplementary Data S2). Both sexually mature and immature individuals were present for all species except for tiger shark, grey reef shark Carcharhinus amblyrhynchos, and common smooth-hound (Table 2). Out of the seven species that had sufcient sampling, four—pelagic thresher, silky shark, oceanic whitetip shark, and blue shark—showed a higher proportion of immature than mature individuals. For oceanic whitetip shark (Critically Endangered), nearly all individuals were immature (96%; 157 out of 164), whereas for silky shark (Vulnerable), immature individuals comprised 86% of the sample (195 out of 228). Size structure for silky and oceanic whitetip sharks is presented in Fig. 3.

The legal case against vessel Fu Yuan Yu Leng 999. Te provincial court of Galápagos, on San Cris- tobal Island, ruled in favor of the Galápagos National Park Directorate, imposing legal sanctions on the Fu Yuan Yu Leng 999 vessels’ owner and crew. Te process’ legal base rested on the Ecuadorian law prohibiting the unau- thorized possession and transport of protected species and on trespassing into the Galápagos Marine Reserve without authorization 14. Te fnal indictment resulted in the incarceration for 1–3 years of 20 crew members, confscation of the vessel, and a fne of US$ 6.1 million for damages inficted to protected species 15. Discussion Te case of vessel Fu Yuan Yu Leng 999 represents the largest reported shark seizure in Ecuadorian waters to date. It reveals an unprecedented magnitude of targeted shark fsheries and highlights the vulnerability of crew members employed by international fshing feets. Although based on a single case, the fsheries catch reported here exemplifes ongoing industrial operations that are likely depleting shark populations in the region on a vast scale. Tis situation might have profound implications for the management and conservation of shark populations in the Eastern Tropical Pacifc. Tere is no certainty about the origin of sharks found in the vessel Fu Yuan Yu Leng 999. However, machine- learning analysis of the data from the Automatic Identifcation System (AIS) of the vessel detected an anomaly in its trajectory 2735 km northwest of the Galápagos Marine Reserve 16 (Fig. 4). Estimated locations of vessel rendezvous coincide with the movements of four Chinese fagged tuna long-liners, suggesting that the Fu Yuan Yu Leng 999 loaded cargo in the area without authorization, making the operation illegal 17. Tese inferences are in line with the statements of crew members during legal procedures. Also, it is possible that some of the longliners trespassed other countries’ economic exclusive zones (EEZ) or received catch from coastal fshing ships. Our fnding of two grey reef sharks, a non-pelagic species found in the western Pacifc and the Central Pacifc Islands 18, supports this scenario. Tese detection anomalies highlight the importance of satellite technologies, such as AIS and vessel monitoring systems (VMS), in inferring true vessel trajectories, enforcing approved routes, and diminishing the probability of vessels trespassing into EEZs and marine protected areas19. Whether the Fu Yuan Yu Leng 999 vessel would engage in potentially illegal operations around the Galápagos EEZ, remains unclear. Still, in its empty freezer holdings, the vessel had a gross capacity for at least 4.5 times the number of sharks found in this study (more than 30,000 additional individuals), which may have been received by the vessel along its south-bound trajectory if not been seized. Te number of individuals analyzed in this study constitutes a representative sample of the Fu Yuan Yu Leng 999 vessel’s total cargo, which was highly biased towards sharks. Te most common species found in the sample, the silky shark (28%), has been reported as the second most abundant species in the Hong Kong and Guangzhou markets in China, the largest shark fn retail centers in the world20. Even more worrisome is the situation of the oceanic whitetip shark (20% of the sample), which is Critically Endangered. In 2011, in response to concerns about declining trends in catches of oceanic whitetip sharks, a ban by the Inter-American Tropical Tuna Commis- sion (IATTC) was placed on the retention, transshipping, landing, storing, or selling of this species (Resolution

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Proportion of # Sexually # Sexually mature Size range SM Size of sexual Species Common name immatures immature (SI) Size range SI (cm) (SM) (cm) Length used maturity* Alopias pelagicus Pelagic thresher 55% 52 63.0–137.8 42 139.0–174.0 PCL 138 cm51 Alopias supercil- Bigeye thresher 48% 21 160.2–249.5 23 254.4–302.3 TL 253 cm52 iosus Carcharhinus Grey reef shark – 0 NA 2 132.9–143.6 TL 116.753 amblyrhynchos Carcharhinus Silky shark 86% 195 37.0–133.0 33 136.0–195.0 PCL 135–140 cm38 falciformis Carcharhinus longi- Oceanic whitetip 96% 157 76.7–169.9 7 189.1–257.6 TL 172 cm39 manus shark Galeocerdo cuvier Tiger shark – 2 145.1–212.3 0 NA TL 276 cm54 Prionacea glauca Blue shark 86% 58 96.8–180.8 9 182.1–199.2 TL 182 cm55 Isurus oxyrinchus Shortfn mako – 2 120.4–171.2 1 203.4 TL 180 cm56 Scalloped ham- Sphyrna lewini 27% 25 117.0–162.0 69 163.3–295.3 TL 162–181.2 cm57 merhead Smooth ham- Sphyrna zygaena 7% 3 118.4–172.8 43 179.7–289.9 TL 178.158 merhead Common smooth- Mustelus mustelus – 0 NA 1 161.9 TL 88 cm59 hound Total 515 230

Table 2. Proportion of sexually mature individuals inferred from pre-caudal length. References are for studies reporting size for sexual maturity for males. PCL precaudal length, TL total length.

C-11-1021). Tus, our fnding of 188 oceanic whitetip sharks implies that the Fu Yuan Yu Leng 999 crew was violating the ban of the IATTC, of which China is a signatory. Another common species in the sample, the blue shark (12%), has been reported as the most abundant species in Asian markets 20. Tis species is not yet been listed by CITES because of its Near Treatened status (i.e., close to qualify for a threatened category in the near future; https://www.iucnredlist.org/). However, regionally, the blue shark represents the most common shark species landed in Peru22, and has been reported as by-catch in fsheries from Ecuador 23 and Chile24. Other endangered shark species, such as hammerheads (collectively 20% of the sample), are highly vulnerable to non-target fsheries in the Eastern Pacifc Ocean23, 25, 26. Additional threads may derive from the high proportion of immature individuals for some species in the sample. A comparison of fork length distribution of the seized silky and oceanic whitetip sharks (Fig. 3) with size-structured catch data from industrial feets in the region27, suggests that these species may have been caught using purse-seine gear. Consistent with these data, the semi-industrial longline feet from mainland Ecuador caught mostly adults of silky shark and only a small proportion of oceanic whitetip shark in 2008–201223, 28. Sam- ples of both species of hammerhead shark in the vessel were comprised mostly of adults, which is also consistent with the use of purse-seine gear 29. In 2016, the IATTC placed a precautionary ban for 2017–2019 on retention, transshipment, landing, or storing of silky sharks caught by purse seines in the convention area (Resolution C-16-0630). Tus, our data, which suggest fshing a high proportion of silky sharks with purse seines, points to yet another violation of the IATTC by the Fu Yuan Yu Leng 999 crew. Tese data exemplify how the operation of international fshing feets may be hampering the eforts of other IATTC members to protect shark species. In recent years, international feets comprising hundreds of vessels have been reported repeatedly near the Ecuadorian EEZs (Galápagos and Continental EEZs). In July 2017 and August 2017 (a few days before the seizure of vessel Fu Yuan Yu Leng 999), international feets of ~ 300 ships (mostly Chinese fagged) were detected in the international waters bordering the Galápagos’ EEZ 31 and a similar event happened in March 2019 32. More recently, on 16 July 2020, a feet of 260 ships (mostly Chinese fagged) was reported along the southern border of Galápagos’ EEZ33. By 6 August 2020, the feet had grown to 340 ships34, and at least 6 were transmitting false coordinates that located them within New Zealand’s waters35. Tese events suggest that the presence of international fshing feets around the Ecuadorian EEZs may be systematic. Given the highly mobile nature of the targeted species identifed here, international feets are likely catching sharks that are demographically linked to populations protected under national legislation, undermining the eforts Ecuador has adopted for the conservation of sharks. Current proposals to enhance the protection of migratory marine life in the region include expanding Ecuador’s EEZ along the Carnegie and Cocos ridges (Sup- plementary Information Figure S2). However, the seasonal return of large industrial feets to the exact borders of the Ecuadorian EEZs implies that the resources within these zones are of particular interest. Te high seas in the Eastern Tropical Pacifc, including those surrounding the Galápagos EEZ, harbor marine resources (in this case, sharks), which are targeted by international fshing feets, with limited governance from Regional Fisheries Management Organizations (RFMOs). A better understanding of the demographic and genetic connectivity of targeted shark populations across jurisdictional boundaries is urgently needed to inform management and improve the conservation of the biological marine resources of the Eastern Tropical Pacifc both within EEZs and in Areas Beyond National Jurisdiction (ABNJs). Here, we provide the largest data set on shark species caught in the region and contribute to a growing data base of genetic information needed to assess the identity of fshed sharks in the future. Improved governance of

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Oceanic Whitetip Shark n = 188 60 Size at maturity 50 Median FL of captures with purse 40 seines 90% Quantiles of 30 Median FL of captures with purse seines Frequency 20

Fork Length (cm)

Silky Shark n = 257

45 Size at maturity 40 35 Median FL of captures with purse 30 seines 25 90% Quantiles of 20 Median FL of captures Frequenc y with purse seines 15 10 5 0

Fork Length (cm)

Figure 3. Size structure for oceanic whitetip and silky sharks seized in vessel Fu Yuan Yu Leng 999 (based on 188 and 257 individuals, respectively). Size to maturity for oceanic whitetip was based on Joung et al.39, by transforming Total Length to Fork Length49. Median Fork Length of captures with purse seines and 90% quantiles were approximated from Clarke et al.27 Figure S3. we created this fgure using Microsof Excel Spreadsheet (https://www. micro sof. com/ en- ww/ micro sof- 365/ excel ) and Adobe Creative Suite (https://www. adobe.com ).

fsheries in ABNJs and their compliance with international treaties, will be paramount to preserve biological and economical marine resources of developing countries in the Eastern Tropical Pacifc and tropical seas in general. Methods Sampling, morphological identifcation, and size of sexual maturity. Sampling was conducted onboard vessel Fu Yuan Yu Leng 999 in September and October 2017, in a collaborative efort between the National Army of Ecuador, the GNP Directorate, and the Galapagos Science Center (GSC) of Universidad San Francisco de Quito and the University of North Carolina at Chapel Hill (research permits PC-38-16 and PC-24- 17). Te procedure was performed under the time and logistic constraints imposed by the seizure’s legal process, the low temperatures of the vessel holds, and the identifying and measuring of frozen samples—with limited personnel and the vessel in motion—while destruction/disposal of the cargo was taking place. We tried to take a representative sample of all species present in the vessel, choosing a random sample of individuals within each species. Morphological identifcation was based on expert’s knowledge and specialized literature18, 28. Tissue samples and pre-caudal body length were taken for as many individuals as possible. Te total vessel cargo was weighed to the nearest 0.1 kg before disposal. Te size of sexual maturity was inferred from the literature to estimate the ratio of immature-to-mature individuals in our sample. For pelagic thresher (Alopias pelagicus) and silky shark (Carcharhinus falciformis), size of sexual maturity was inferred from pre- caudal length. For other species, we used conversion factors to calculate total length from pre-caudal length, as

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Figure 4. Trajectory of vessel Fu Yuan Yu Leng 999, including AIS–detected anomaly. Te vessel departed from Fuzhou on the Chinese coast 50 and was detained at Punta Pitt, San Cristobal Island, within the Galápagos Marine Reserve. Figure from Cutlip 17, its use authorized by Global Fishing Watch.

follows: for shortfn mako (Isurus oxyrinchus), blue shark (Prionace glauca), and smooth hammerhead (Sphyrna zygaena) we followed Mas et al.36; for oceanic whitetip (Carcharhinus longimanus) D´Alberto et al.37; for bigeye thresher (Alopias superciliosus), grey reef shark (Carcharhinus amblyrhynchos), tiger shark (Galeocerdo cuvier), and smooth-hound (Mustelus mustelus), FishBase (https://www. fshb ase. de ). For scalloped hammerhead (Sphy- rna lewini), we used the conversion factor for smooth hammerhead36. Since we lacked sex ID for all individuals, we defned minimum size at maturity using estimates for males only (which tend to be smaller than females). When a range was provided, we used the minimum size. To compare the size structure of silky and oceanic whitetip sharks found in the vessel with that of sharks fshed with long-lines and purse-seines27, we converted pre-caudal length to fork length using Oshitani et al.38 and Joung et al.39 conversion factors, respectively.

Molecular identifcation. We sequenced the mitochondrial gene Nicotinamide Adenine Dehydroge- nase subunit 2 (ND2), a barcode gene for sharks 40 (permits MAE-DNB-CM-2016-0041 and MAE-DNB-2018- 0759-O). We conducted genomic DNA extractions on samples of subcutaneous muscle tissue (2 × 2 mm). Extrac- tion followed a guanidine thiocyanate protein precipitation plus isopropanol DNA precipitation protocol 41. Quality and concentration of genomic DNA were measured with a NanoDrop ND-1000 spectrophotometer v3.0.1 (NanoDrop, TermoFisher Scientifc) or an Epoch Microplate Spectrophotometer (BioTek). Te ND2 gen was amplifed by PCR in 25 μl reactions comprised of 0.25 μl Paltinum Taq DNA Polymerase, Master Mix (Invitrogen), 2.50 μl 10X PCR Bufer, 1.5 mM of MgCl2, 0.50 μl of 10 μM of each dNTP, 0.5 μl of each primer (10 μM), and diferent amounts of ultrapure water with 1 μl of template DNA (adjusted to 100 ng/mL of DNA). Termocycling conditions included an initial denaturation at 94 °C/3 min; 35 cycles of denaturation at 93 °C/ 30 s, annealing at 58 °C/1 min, and extension at 72 °C/1 min; and a fnal extension at 72 °C/10 min. Te resulting amplicons were visualized via electrophoresis in a 2% agarose gel (UltraPure Agarose). Excess nucleo- tides and dNTPs were removed from PCR products using ExoSAP-IT PCR Product Cleanup Reagent (Applied Biosystems). Purifed amplicons were sequenced with big-dye chemistry and PCR primers, using capillary electrophoresis in an ABI3730xl sequencer. Inspection of chromatograms, contig assembly of forward and reverse sequences, and fnal sequence checking and edition were performed in Geneious 11.1.5 (Biomatters Ltd.42).

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To evaluate the similarity between sequences from the vessel’s samples and those on GenBank (https://www. ncbi.nlm.nih.gov/genbank/), we used BLAST (https://blast.ncbi.nlm.nih.gov/Blast.cgi). To place the sample sequences in a phylogenetic framework, we consulted the Chondrichthyes Tree of Life (https://sharkrays.org) to assemble a list including all shark species from the Pacifc Ocean, and downloaded the sequences for these species from Genbank. Locality information for sequences of the species present in the vessel was gathered, when possible, in an attempt to relate the seized samples to general geographic regions in the Pacifc Ocean. Additionally, we sequenced samples from nine individuals of the Galapagos shark, Carcharhinus galapagensis, obtained from the Galápagos Marine Reserve (GNPD permit 065-2013). Sequences of samples from vessel Fu Yuan Yu Leng 999, the Pacifc Ocean (gathered from GenBank), and Galapagos shark were aligned using MAFFT 7 43. All sequences were inspected, translated into amino acids (to look for potential stop-codons), and trimmed to the size of the ND2 mitochondrial gen using Mesquite 3.644. To reduce the size of our matrix and facilitate phylogenetic analyses, we eliminated duplicates (identical sequences) in all datasets separately, using sRNAtoolbox45. Ten, we compiled the unique sequences from each dataset and re-aligned them in MAFFT 7. We used IQ-TREE46, as implemented in the IQ-TREE web server47 (http://iqtree.cibiv.univie.ac.at/) to infer the best-ft model of nucleotide substitution, estimate the maximum likelihood tree, and assess nodal support. Te best-model ft was estimated using ModelFinder48 in “Auto” function and “FreeRate heterogeneity”; the best- ft model (GTR + F + R7) was chosen using the Bayesian information criterion. Nodal support was obtained using 1000 ultrafast bootstrap replicates, 1000 maximum iterations, and minimum correlation coefcient of 0.99. As outgroup, we used the rabbit fsh, Chimaera monstrosa (GenBank accession number JQ518716). Data availability All the DNA sequences generated during this study are available on GenBank (Supplementary Data S1). All morphological data used is available from the Supplementary Information Data S1.

Received: 19 March 2021; Accepted: 30 June 2021

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Acknowledgements We are grateful to the national army of Ecuador, Fuerzas Armadas Nacionales, and especially Comander Dario Ortega (Zona Naval de San Cristóbal, Galápagos), for their support during the vessel seizure. To the Galápagos National Park Directorate for its institutional support and to its staf—Galo Quezada, Daniel Lara-Solis, Jorge Carrión, Gabriel Vásquez, and Maryuri Yépez—Galápagos National Park rangers, and Ingrid Jaramillo for their assistance during sampling. Special thanks to our volunteers Diego Iglesias, Jacob Salinas, Jens Mayorga, Alice Skehel, Jaime Grijalva, and Jason Castañeda, who assisted during sample collection under such hard conditions. Gabriela Gavilanes and Nathalia Valencia assisted during sequencing procedures. Finally, we want to thank GSC staf—Carlos Mena, Stephen Walsh, Philip Page, Sofa Tacle, Sylvia Sotamba, Cristina Vintimilla, and Soledad Sarzosa—for their continuous support. Tis study was funded by Te National Geographic Society (Grant Award

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NGS-421C-18), Universidad San Francisco de Quito (HUBI 10140), and a generous donation from Michael Smith towards shark conservation in the Galápagos (USFQ, HUBI 12204). Author contributions E.B.: Study concept and design of molecular component; acquisition of funding; fnal data analyses; drafing of the manuscript. N.O.-G.: acquisition and preliminary analyses of data; partial drafing of the manuscript. D.A.P.: comparative data compilation; morphological data analysis; provision of Galápagos shark samples; critical revision of the manuscript. A.H.: comparative data compilation; partial drafing of the manuscript; critical revision of the manuscript. D.P-R.: acquisition of research permits; sampling logistics; critical revision of the manuscript. S.C.: morphological identifcation; photography; critical revision of the manuscript. J.P.M-P.: sample collection and sampling logistics; review of the manuscript. E.E.: sample collection and sampling logistics. J. S.: sample collection and taking of morphological measurements; L.D.M-R.: sample collection and taking of morphological measurements. A.V.: sample collection and taking of morphological measurements. J.A.C.: acquisition of funding; critical revision of the manuscript. M.L.T.: laboratory support; critical revision of the manuscript. W.B.: logistic and institutional support; critical revision of the manuscript. D. R.: logistic and institutional support. M.H.: morphological identifcation; partial drafing of the manuscript; critical revision of the manuscript. J.M.G.: Study concept and design of molecular component; sampling logistics; acquisition of funding; critical review of the manuscript.

Competing interests Sebastián Cruz (SC) declares competing interest given its involvement as subject expert hired during the legal inspection of vessel Fu Yuan Yu Leng 999 for the morphological identifcation of shark species. Additional information Supplementary Information Te online version contains supplementary material available at https://doi.org/ 10.1038/s41598-021-94126-3. Correspondence and requests for materials should be addressed to E.B. Reprints and permissions information is available at www.nature.com/reprints. Publisher’s note Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional afliations. Open Access Tis article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. Te images or other third party material in this article are included in the article’s Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by/4.0/.

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