First records of pregnant female sharks (Carcharhiniformes; Lamniformes) in the Ilha Grande–Sepetiba estuarine complex (Rio de Janeiro, Brazil): evidence of a reproductive hotspot within a marine protected area

Primeiros registros de fêmeas grávidas de tubarões (Carcharhiniformes; Lamniformes) no complexo estuarino Ilha Grande–Sepetiba (Rio de Janeiro, Brasil): evidências de um hotspot reprodutivo em uma área marinha protegida

Abstract

This study provides the first records of six pregnant sharks (spinner shark - Carcharhinus brevipinna (n = 2), blacktip shark - Carcharhinus limbatus (n = 2), and sand tiger shark - Carcharias taurus (n = 2)) in the Ilha Grande–Sepetiba estuarine complex, southeastern Brazil, highlighting the ecological importance of this region as a potential reproductive and nursery area. Reproductive data for carcharhinoid and lamnoid sharks had not previously been documented in this region. The pregnant females were examined allowing the assessment of embryo number, yolk content, and developmental stages (early and late). Variation among females provides new insights into reproductive dynamics and habitat use within this estuarine–coastal region. These findings strengthen the scientific basis for conservation and management strategies aimed at protecting shark populations in the Southwestern Atlantic and reinforce the regional importance of the Ilha Grande–Sepetiba estuarine complex for key elasmobranch life-history processes.

Keywords:
reproduction; shark ecology; nursery area; marine monitoring; conservation

Resumo

Este estudo apresenta o primeiro registro de seis tubarões grávidas (tubarão-rotador - Carcharhinus brevipinna (n = 2), tubarão galha-preta - Carcharhinus limbatus (n = 2) e tubarão-mangona - Carcharias taurus (n = 2)) no complexo estuarino Ilha Grande–Sepetiba, no sudeste do Brasil, evidenciando a importância ecológica da região como potencial área reprodutiva e de berçário. Até o momento, não havia registros de informações reprodutivas para tubarões carcharhiniformes e lamniformes nesta região. As fêmeas grávidas foram examinadas por meio de necropsias in situ e em laboratório, permitindo a avaliação do número de embriões, conteúdo vitelino e estágios de desenvolvimento (inicial e avançado). As variações observadas entre as fêmeas fornecem novos insights sobre a dinâmica reprodutiva e o uso de habitat nesse complexo estuarino-costeiro. Esses achados fortalecem a base científica para estratégias de conservação e manejo, reforçando a relevância regional do complexo Ilha Grande–Sepetiba para processos-chave da história de vida de elasmobrânquios no Atlântico Sudoeste.

Palavras-chave:
reprodução; ecologia de tubarões; área de berçário; monitoramento marinho; conservação

1. Introduction

Sharks are highly migratory animals that use coastal bays and estuarine environments during seasonal movements for feeding and reproduction (Reyier et al., 2014; Lana, 2012, 2016; Ebert et al., 2021; Pessoa et al., 2025; Lana et al., 2025). Along the Brazilian, coast 99 species of sharks have been registered, including the spinner shark Carcharhinus brevipinna (Valenciennes, 1839), the blacktip shark Carcharhinus limbatus (Valenciennes, 1839) and the sand tiger shark Carcharias taurus (Rafinesque, 1810) (Gomes et al., 2019; Ebert et al., 2021; Kotas et al., 2023). These species are coastal-pelagic and occur in warm temperate, tropical and subtropical oceans, from shallow inshore zones to the outer continental shelves down to 200 m depth (Otway and Ellis, 2011).

Carcharhinus brevipinna is characterized by the presence of black spots on the distal tips of paired and unpaired fins (pectoral, dorsal and lower lobe of caudal fin) and is often misidentified with C. limbatus. Both species are distinguished by the presence of black spots on the distal tip of the second dorsal and anal fins (vs. spot absent in C. limbatus), absence of a black spot on the pelvic fin (vs. present in C. limbatus), first gill slit more than four times greater than the eye diameter (vs. 2.5 times) and first dorsal fin origin posterior to the pectoral insertion (vs. opposite or anterior to the pectoral insertion) (Gomes et al., 2019; Ebert et al., 2021). Both Carcharhinus species possess a nictitating membrane below the eye, which is absent in the lamniform C. taurus. The sand tiger Carcharias taurus is characterized by brown spots on the upper body, a flattened-conical snout, small eyes, protruding spike-like teeth and small, equal-sized dorsal and anal fins.

Carcharhinus limbatus and C. brevipinna are viviparous and placental, developing internally and being nourished by a placenta before birth. Gestation lasts 10 to 12 months, occurring in alternate years, with each female carrying 1-10 embryos in C. limbatus and 3-15 embryos in C. brevipinna (Gomes et al., 2019; Ebert et al., 2021). Females of C. brevipinna mature at about 8–10 years, whereas those of C. limbatus reach maturity around 4- 7 years. C taurus displays aplacental viviparity, with embryos developing precocious dentition to consume their uterine siblings (adelphophagy) and unfertilized oocytes supplied by the mother (oophagy) (Compagno, 1984; Branstetter and Musick, 1994). Male sand tigers reach maturity between 190-195 cm total length (TL) at the age of 6–7 years and females between 220-230 cm TL at 9–10 years (Lucifora et al. 2002; Goldman et al. 2006). The size at birth is between 95-105 cm TL (Compagno, 1984; Branstetter and Musick, 1994).

Previous studies have documented nursery areas, parturition grounds and reproductive patterns for sharks in different regions of the Southwestern Atlantic, including northeastern and southern Brazil (Yokota and Lessa, 2006; Bornatowski, 2008; Santander-Neto et al., 2020; Lucifora et al., 2002). More recently, fishers’ ecological knowledge has highlighted the long-term decline and overexploitation of shark populations in southeastern Brazil (Fogliarini et al., 2024). However, reproductive records for sharks within the Ilha Grande–Sepetiba estuarine complex remain undocumented. Therefore, this study provides the first documented records of pregnant female sharks within this protected coastal region, reinforcing its ecological importance as a potential reproductive hotspot for threatened shark species in southeastern Brazil.

2. Material and Methods

2.1. Study area

Sepetiba-Ilha Grande bays, an estuarine complex located along the southern coast of Rio de Janeiro State, southeastern Brazil (22º55’S-23º25’S; 44º42’W–43º34’W), approximately 60 km in length (Figure 1). This tropical region, with a mean annual rainfall of 1,400 mm, is recognized as a marine biodiversity hotspot due to the presence of important remnants of the insular Atlantic Forest and a high diversity of coastal ecosystems, including rocky shores, islands, sandy beaches, and mangroves (Belo, 2002; Cutrim et al., 2021a; Silva et al., 2022; Lana et al., 2025). In addition to its ecological relevance, the region also has major economic and social importance, supporting traditional communities, artisanal and commercial fisheries, and other productive activities. However, these ecosystems have been increasingly affected by intense urbanization and the presence of nuclear power plants, oil terminals, ports, and industrial facilities (Barrera-Alba et al., 2019; Mordente, 2020; Miranda et al., 2024; Pessoa et al., 2025). Together, the two bays cover approximately 3,100 km², with a perimeter of about 220 km and bathymetric variation ranging from 1 to 30 m (UNACOOP, 2011; Mordente, 2020). The region also includes important marine protected areas (MPAs), such as the Tamoios Ecological Station (ESEC Tamoios), established in 1990 between the municipalities of Angra dos Reis and Paraty, which protects 29 emergent areas and their surrounding marine waters within a 1-km radius, covering 5.69% of Ilha Grande Bay, while the entire bay serves as its buffer zone. In Sepetiba Bay, the Boto-Cinza Marine Protected Area, established in 2015 in the municipality of Mangaratiba, protects 240 km² of the bay’s 536 km² water surface.

Figure 1
Map of the study area with delimitations of marine protected areas and capture location of Sand Tiger sharks (red dot), Blacktip sharks (yellow dot) and Spinner sharks (blue dot).

Despite their close proximity, Sepetiba and Ilha Grande bays differ markedly in their hydrodynamic characteristics (Belo, 2002; Mordente, 2020). Both systems receive freshwater input from rivers and channels; however, Ilha Grande Bay is directly connected to the open ocean and is strongly influenced by tidal circulation, which introduces colder and more saline waters that mix with the warmer and less saline waters of Sepetiba Bay. In contrast, the sandy barrier of Restinga da Marambaia restricts the oceanic connection of Sepetiba Bay, making it more strongly influenced by continental freshwater inflows and only weakly affected by tidal forcing (Belo, 2002; Mordente, 2020).

2.2. Sharks database

Data were obtained opportunistically from pregnant females of C. taurus, C. brevipinna, and C. limbatus recorded in the Ilha Grande–Sepetiba estuarine complex, southeastern Brazil. All sampling procedures and biological analyses were conducted under permits issued by the Brazilian Biodiversity Information and Authorization System (SISBIO), under license numbers 86236-2 and 82226-1, the latter currently valid and officially granted to Instituto PROSHARK.

Specimens were documented through two complementary sources: (i) monitoring artisanal fishing landings and incidental captures, and (ii) scientific monitoring activities conducted in the study area. Sampling events occurred in both Ilha Grande Bay and Sepetiba Bay, including areas within or adjacent to marine protected areas (MPAs), such as the buffer zone of the Tamoios Ecological Station (ESEC Tamoios) and the Boto-Cinza Marine Protected Area.

Pregnant females captured by artisanal fisheries were obtained from local fishers operating mainly with gillnets, including bottom-set gillnets in Ilha Grande Bay and gillnets in Sepetiba Bay. In these cases, biological data were collected directly at landing sites or at the capture location whenever possible. For specimens associated with scientific monitoring, individuals were recorded either after stranding events or during directed research activities conducted with the support of local fishers. In the latter case, one C. limbatus female was capture using drumlines equipped with straight hooks as part of a scientific monitoring and satellite tagging project, tagged with a MiniPAT satellite tracking device, and later recovered for biological analysis.

All available females were measured for total length (TL, m), and reproductive condition was assessed through necropsy. When immediate transport was not feasible, necropsies were conducted in situ; otherwise, specimens were transported to the laboratory for examination. During necropsy, the reproductive tract was inspected to confirm pregnancy and determine the number and developmental stage of embryos. Embryos were removed, counted, and measured for total length (cm), and qualitative observations were made regarding developmental condition, including external morphology and yolk sac condition when present. Specimens were examined and processed at the Chondrichthyes Laboratory (LABCHON), Federal University of Rio de Janeiro (UFRJ), where the biological material was officially cataloged under DOC No. 001, dated June 2024. As records were obtained opportunistically from fishery landings, strandings, and scientific monitoring, sampling was not standardized in time or effort.

The collected biological and reproductive data were used to characterize embryonic development, reproductive condition, and the ecological relevance of the Ilha Grande–Sepetiba estuarine complex for shark reproductive processes in southeastern Brazil.

3. Results

Pregnant females of three shark species (C. taurus, C. brevipinna, and C. limbatus) were recorded in the Ilha Grande–Sepetiba estuarine complex between April 25, 2006, and June 6, 2024, including areas within or adjacent to local marine protected areas.

Two pregnant females of C. taurus were recorded in the buffer zone of ESEC Tamoios, in Ilha Grande Bay, on April 25, 2006. Necropsy of the first female revealed two embryos measuring 76.9 and 77.2 cm TL, whereas the second female contained four embryos ranging from 18.8 to 20.5 cm TL (Figure 2D). Two additional pregnant females of C. brevipinna were recorded within the limits of the Boto-Cinza Marine Protected Area, in Sepetiba Bay, on August 30 and 31, 2023. Both females contained six to seven embryos in an advanced stage of development, all measuring more than 40 cm TL.

Figure 2
Pregnant females and embryos of spinner sharks (C. brevipinna - A), blacktip sharks (C. limbatus - B, C) and sand tiger sharks (C. taurus - D) recorded in Ilha Grande Bay during this study.

For C. limbatus, two pregnant females were documented in Ilha Grande Bay. The first, measuring 2.14 m TL, was found dead and stranded in the Piraquara region, within the limits of ESEC Tamoios, on June 11, 2020. Necropsy revealed an advanced stage of pregnancy, with five fully developed embryos measuring between 20 and 27 cm TL. The cause of death could not be determined (Figure 2B). The second female, measuring 2.30 m TL, was recorded on June 6, 2024, within the limits of ESEC Tamoios, after being previously tagged with a MiniPAT satellite transmitter. The individual was recovered and transported to the Chondrichthyes Laboratory at UFRJ for necropsy. Examination revealed four embryos ranging from 7.1 to 9.4 cm TL, all in an early stage of development, as indicated by the substantial yolk reserves present in the yolk sacs and the apparent absence of placental development (Figure 2C). Although necropsy findings did not allow determination of the cause of death, the observed conditions suggest a possible comorbidity; however, no conclusive cause could be established.

4. Discussion

This study provides the first documented records of pregnant spinner sharks (Carcharhinus brevipinna), blacktip sharks (Carcharhinus limbatus), and sand tiger sharks (Carcharias taurus) within the Ilha Grande-Sepetiba estuarine complex southeastern Brazil. Until now, no reproductive records of pregnant females had been documented for this region, representing an important contribution to the understanding of elasmobranch reproductive ecology and population dynamics along the southeastern Brazilian coast. Although these species exhibit migratory behavior, the occurrence of pregnant females at different embryonic developmental stages suggests that the Ilha Grande–Sepetiba complex may function as a potential reproductive, refuge, and nursery-related habitat for large coastal sharks.

Previous studies have documented reproductive and nursery areas for sharks in different regions of the Southwestern Atlantic, including northeastern and southern Brazil (Yokota and Lessa, 2006; Bornatowski, 2008; Santander-Neto et al., 2020; Lucifora et al., 2002). In this context, our findings reinforce the regional ecological importance of this protected coastal system for critical elasmobranch life-history stages. These records are further supported by ongoing monitoring efforts coordinated by Instituto PROSHARK, which have documented seasonal aggregations of blacktip sharks (C. limbatus) and tiger sharks (Galeocerdo cuvier) in Ilha Grande Bay associated with reproductive behavior, habitat use, and satellite-tracked movements (Lana et al., 2025, 2026), as well as reproductive behavior of mobulid rays within the same region, including Mobula cf. tarapacana, Mobula cf. birostris and Mobula mobular (Pessoa and Lana, 2025; Pessoa et al., 2025). Similarly, other marine megafauna species, such as sea turtles, rays, and seabirds, also use this region during critical life-history stages (Cutrim et al., 2021a, b). Together, these findings reinforce the ecological importance of the Ilha Grande–Sepetiba estuarine complex as a potential reproductive and aggregation hotspot for large elasmobranchs in southeastern Brazil.

This record provides valuable insights into the biology of coastal pelagic sharks, suggesting the existence of a potential primary nursery area. This hypothesis is supported by the repeated occurrence of pregnant C. limbatus females during similar periods across different years. Additionally, satellite and acoustic tracking data generated through the ongoing monitoring program coordinated by Instituto PROSHARK have indicated extensive use of Ilha Grande Bay by multiple shark species over time, including C. limbatus and Galeocerdo cuvier (Lana et al., 2025, 2026), demonstrating strong site fidelity and reinforcing the ecological importance of this region for large coastal sharks. This fidelity may be particularly significant, as the seasonal movements of these sharks are closely linked to water temperature, which directly influences their movement patterns and habitat selection (Lana et al., 2025). Currently, ongoing research initiatives in the region and across Rio de Janeiro State are expanding this evidence base through the continued deployment of satellite and acoustic telemetry, generating new data on the spatial behavior, habitat use, and movement patterns of blacktip sharks and other shark species occurring in Ilha Grande Bay and along the coast of Rio de Janeiro. By integrating technological innovation with long-term ecological monitoring, these initiatives reinforce the scientific basis for regional management and conservation planning in the southwestern Atlantic. Collectively, they are increasing temporal resolution and taxonomic coverage, thereby advancing a more robust understanding of ecological dynamics within this estuarine–coastal system.

Nursery area selection by Carcharhiniform sharks typically occurs in shallow, protected coastal zones characterized by warmer temperatures, high prey availability, and reduced predation risk, which enhances neonate survival (Lana et al., 2025). The seasonal aggregation of blacktip sharks (C. limbatus) and spinner sharks (C. brevipinna), previously reported in regional monitoring studies (Rolim et al., 2025; Lana et al., 2025), together with recent records of tiger sharks (Galeocerdo cuvier) aggregations and satellite-tracked movements obtained through Instituto PROSHARK monitoring efforts (Lana et al., 2026), further support this hypothesis, as such aggregation patterns are frequently associated with reproductive behavior and habitat use by large coastal sharks.

The environmental conditions within this marine protected area (MPA) also favor its role as a nursery, as estuarine systems provide refuge for pregnant females, while the occurrence of pregnant females through capture suggests site fidelity during gestation, a common trait in sharks that use nursery areas. Protecting these habitats is crucial for conservation, as nursery areas are vulnerable to habitat degradation and overfishing. Future research should focus on confirming the presence of neonates, seasonal movement patterns, and small-scale habitat use to strengthen conservation strategies. These priorities are already being addressed through ongoing multidisciplinary research efforts conducted under the Integrated Shark and Ray Monitoring Program of Rio de Janeiro State, coordinated by Instituto PROSHARK, particularly within the Ilha Grande–Sepetiba estuarine complex. These efforts include telemetry, drone-based surveys, and field-based monitoring, generating integrated data on movement patterns, habitat use, and population structure within this estuarine–coastal system.

The Piraquara de Fora region, in Ilha Grande Bay, has been formally recognized as an Important Shark and Ray Area (ISRA) under the IUCN Important Shark and Ray Areas initiative (IUCN SSC Shark Specialist Group, 2025), highlighting the aggregation of Carcharhinus limbatus and the ecological relevance of the site. The proposal was developed with Instituto PROSHARK in collaboration with other researchers working in the region. This designation reinforces the scientific importance of ongoing shark research in the area and elevates Ilha Grande Bay as a site of global conservation significance. The recognition represents a major milestone for the State of Rio de Janeiro and for Brazil, strengthening the country’s contribution to international marine biodiversity targets and advancing global efforts to safeguard critical habitats for sharks and rays in the Southwestern Atlantic (IUCN SSC Shark Specialist Group, 2025; Lana et al., 2025).

Our findings provide robust scientific evidence supporting the conservation and management of C. limbatus, C. brevipinna, C. taurus, and other shark species inhabiting the Ilha Grande–Sepetiba estuarine complex. By documenting patterns of occurrence and ecological relevance within this system, this study advances the understanding of critical habitats in the Southwestern Atlantic and strengthens the regional knowledge base necessary for informed decision-making. In the context of increasing anthropogenic pressures and fisheries interactions, generating reliable biological and spatial data is essential to guide adaptive management, marine spatial planning, and public policy development. The results presented here contribute directly to ongoing conservation initiatives and offer a solid foundation for integrated coastal–marine governance strategies aimed at ensuring the long-term persistence of shark populations in Brazil and their role within global marine biodiversity conservation frameworks.

Acknowledgements

Authors wish to thank the teammates from the Rio de Janeiro Marine Aquarium (AquaRio) that helped during the laboratory work. Angra dos Reis City Hall and Angra dos Reis Municipal Environmental Institute (IMAAR) for logistical support at sea during the research. Thanks to Eletronuclear S.A. team (LMA - Environmental Laboratory) that provided all the support, as well as credits to veterinarian Renato Velloso da Silveira and the “Tartaruga Viva Program” (“Tartaruga Viva Program” is a requirement of federal environmental licensing, conducted by Ibama - Operating License No. 1217/2014 – 1st Renewal) for carrying out the anatomopathological examination. We also thank Leonardo Neves (UFRRJ) for his contribution of data on sand tiger sharks (C. taurus) and participation in the initial drafting of the article. We would like to offer a special thanks to local fishermen from APMIM at Marambaia Island and Tamoios Ecological Station (ESEC Tamoios) in Ilha Grande-Sepetiba estuarine complex for their essential support during the field work and collaboration with research. This study was initially financed through the project “Marine and Fisheries Research” is an offset measure established under a consent decree agreed between the company PRIO and the Federal Public Prosecutors’ Office in Rio de Janeiro. Was implemented by FUNBIO over the years 2021-2023 (grant n. 02/2021), support received through the Fundação Educacional Ciência e Desenvolvimento (FECD). Karla D.A. Soares was financially supported by Fundação Carlos Chagas Filho de Amparo à Pesquisa do Estado do Rio de Janeiro (FAPERJ - APQ1/2023). Fernanda O. Lana was financially supported by the FECD (2022-2023) and Rio de Janeiro Marine Aquarium (AquaRio) throughout the months of June, July and August of 2024 to carry out field expeditions.

Data Availability Statement

The datasets generated during and/or analyzed during the current study are available from the corresponding author on reasonable request.

• Ethics and authorization statement
  This research was conducted under valid ethical approval granted by the relevant Animal Use Ethics Committee (CEUA), approval number 017-24. All procedures involving animals complied with applicable national regulations and institutional guidelines for the ethical treatment and handling of wildlife.

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Edited by

• Editor:
  Takako Matsumura Tundisi

Publication Dates

• Publication in this collection
  14 Aug 2026
• Date of issue
  2026

History

• Received
  13 Feb 2026
• Accepted
  20 May 2026