Anthropogenic impacts continue to drive loss of function and degradation in marine ecosystems. Biodiversity within these systems has declined over time, and we are now facing the extinction of some of the world’s most important and charismatic species. Elasmobranchs (sharks, rays, skates and chimeras) are showing population declines globally, with more than 30% of species globally threatened with extinction (IUCN, 2022), and oceanic elasmobranch populations are thought to have decreased by at least two-thirds between 1971 and 2020 (Pacoureau et al., 2021). Elasmobranchs overall have a higher extinction risk than most other vertebrates, in part owing to their low fecundity, late maturation and long generation times, with large shallow water-dwelling species most at risk (Dulvy et al., 2014).

Although these declines are due to a cumulative range of anthropogenic impacts – habitat degradation, pollution and climate change, for example – overfishing, resulting in both targeted and incidental catch, is considered the most important driver (Dulvy et al., 2014; Dulvy et al., 2021). Across the world, elasmobranchs are fished by artisanal, recreational and commercial fisheries for their fins, livers and meat (Oliver et al., 2015; Tiralongo et al., 2018a) and it has been estimated from reported landings, unreported landings and discards that the global catch of sharks in 2010 was 1.41 million tonnes (Worm et al., 2013). As well as targeted fishing, a major problem facing elasmobranch populations is bycatch, the incidental capture of non-target species (Oliver et al., 2015). It has been estimated that bycatch makes up 40% of annual global marine catch (Davies et al., 2009). Bycaught elasmobranchs are increasingly being retained by fishers, due to their market value and decreases in target species catch, but a lack of management and legislation persists (Oliver et al., 2015; Pacoureau et al., 2021). Even if species are released alive, there can be sub-lethal impacts on individuals that could, in turn, lead to population impacts, as well as post-release mortalities (Wilson et al., 2014; Pacoureau et al., 2021).

Within the Mediterranean Sea, 88 species of elasmobranchs have been recorded (Serena et al., 2020), and many are facing declines in population size and range (Dulvy et al., 2016). Here, due to increased relative fishing pressure, 50% of skates and rays (hereafter referred to as batoids) and 56% of sharks, have regional International Union for Conservation of Nature (IUCN) Red List conservation status assessments that are elevated compared to their global populations (Dulvy et al., 2016). Historically, elasmobranch species diversity and richness were considered greatest in the western Mediterranean Sea, but in recent years there have been a greater number of local extinctions in the west and an increase in threats throughout the Mediterranean (Dulvy et al., 2016; Serena et al., 2020). There have been fewer studies of elasmobranch diversity in the eastern Mediterranean, although Türkiye alone has over three quarters of all Mediterranean elasmobranch species (Bengil and Basusta, 2018), with known nursery areas for Carcharhinus plumbeus (Nardo 1827, sandbar sharks), Rhinobatos rhinobatos (Linnaeus 1758, common guitarfish) and Glaucostegus cemiculus (Geoffroy St. Hilaire 1817, blackchin guitarfish) in the coastal waters of Türkiye (Bengil et al., 2020; Basusta et al., 2021).

Elasmobranchs make up just 1% of the annual fisheries catch in the Mediterranean (Bradai et al., 2018), however, fishing mortality due to bycatch is a major threat for this group (Bengil and Bengil, 2018). In most Mediterranean countries, elasmobranch bycatch is not reported by species, leading to underestimations of catches and poorly informed conservation efforts (Cashion et al., 2019; Giovos et al., 2021a). Many elasmobranch studies use data from large industrial fishing vessels such as trawlers and longliners (Molina and Cooke, 2012). Given that at least eighty percent of Mediterranean fishing vessels are small-scale vessels (<12m total length; FAO, 2020), and small-scale fisheries (SSF) have recently been identified as the second greatest contributor to bycatch of threatened elasmobranchs in the Mediterranean (after longline vessels; Tiralongo et al., 2018a; Carpentieri et al., 2021), understanding bycatch within the Mediterranean SSF fleet is urgently needed.

A checklist of 60 chondrichthyans was recently reported by a study which reviewed available data, studies and citizen science reports for the island of Cyprus (Giovos et al., 2021b). A published record of Hexanchus nakamurai (Teng 1962, bigeyed sixgill shark, now H. vitulus, Springer and Waller 1969, Atlantic sixgill shark) has since been retracted (Bengil et al., 2021), while a Cetorhinus maximus (Gunnerus 1785, basking shark) previously reported for Northern Cyprus (Kabasakal, 2013), and for the whole island (Giovos et al., 2021b) was based on a newspaper report which was later found to have been inaccurate, instead being Alopias superciliosus (Lowe 1841, bigeye thresher shark, Hakan Kabasakal pers. comm.). This brings the total species count for the entire island to 58 (30 sharks and 28 batoids). Most of the contributing records, however, are from studies focused in the area of the island under the effective control of the Republic of Cyprus (RoC), and until our current study, just fourteen species had been documented in Northern Cyprus (Turkish Republic of Northern Cyprus (TRNC); Oray and Karakulak, 2005; Çoker and Akyol, 2014; Akbora et al., 2019) a self-declared state recognised only by Türkiye and considered by the international community to be part of the Republic of Cyprus (hereafter referred to as Northern Cyprus).

The commercial fishing fleet of Northern Cyprus includes 300-400 SSF vessels [Northern Cyprus Department of Animal Husbandry (DAH)], predominantly fishing with gill nets, trammel nets and longlines. Interactions with marine vertebrate species, including elasmobranchs, marine turtles, Mediterranean monk seals (Monachus monachus, Hermann 1779) and dolphins are common, with an estimated 1,000 marine turtles caught in this fishery each year (Snape et al., 2013; Snape et al., 2018b; Beton et al., 2021). Trawl fishing is not permitted and there are no industrialised longline vessels. Although, marine protected areas (MPAs) have been established to protect sea turtles, monk seals, seabirds and some key habitats across more than a quarter of the coast of Northern Cyprus, (Snape et al., 2018a), few fisheries restrictions are implemented within or outside of these MPAs. Until May 2023, there were no formal protection measures for any elasmobranch species from commercial trade, other than for C. plumbeus and C. maximus of which catching or landing were prohibited (Fisheries Law 27-2000, 2022). This was partly as a result of the lack of data available for policy makers, however, as a result of data from this current study, in May 2023 the fisheries regulations were updated to prohibit trade of all species listed on Annex I of the Barcelona Convention. The authorities have also maintained a blanket ban on industrialised fishing, with no trawling permitted since the 1990s (Ulman et al., 2015), thus, habitats may be in favourable condition compared with other parts of the Mediterranean, where bottom trawling persists.

This study aimed to describe the diversity, distribution and life stages of elasmobranch species caught in the fisheries of Northern Cyprus, through deploying onboard observers and engaging small-scale fishers in data collection, to inform fisheries management and conservation action in line with global, regional and national biodiversity conservation strategies.

A total of 1,702 sets were recorded, 730 by onboard observers and 972 by self-reporting fishers. These included 1,367 demersal net sets, 282 demersal longline sets and 2 pelagic (surface/epipelagic) longline sets, with average set depths of 34.2 m (SD: 23.7 m, range: 1.0-250.0 m), 53.5 m (SD: 39.7 m, range: 6.0-296.0 m) and 162.2 m (SD: 92.4 m, range: 96.8-227.5 m), respectively. The number of observed sets by year and month together with the proportion of these that had elasmobranch bycatch is shown in Supplementary Figure 1 .

Using novel methods in an under-studied, yet regionally important fleet, this study helps further our understanding of elasmobranch diversity and distribution in Cyprus, and the contribution of this coast to Mediterranean biodiversity. It also provides detailed insight into the interaction of elasmobranchs with small-scale fisheries and their wider ecology in the study area. We recorded 36 elasmobranch species in the coastal waters of Northern Cyprus, 61% of which are considered globally threatened and 50% regionally threatened in the Mediterranean (IUCN, 2022). Of these species, two of the three Mediterranean endemic species (R. polystigma and R. radula) were recorded and 24 species (65%) were previously unpublished records for Northern Cyprus, bringing the total number of species recorded from 14 (Çoker and Akyol, 2014; Akbora et al., 2019) to 39. Nearly half (47%) of the 36 species we report are priority species under the GFCM or listed on Appendix II of CITES ( Figure 2 ), meaning that international trade should be controlled.

The most abundant threatened elasmobranch species ( Figure 2 ) caught in our study were D. pastinaca (VU), Mustelus mustelus (Linnaeus 1758, common smoothhound EN) and T. marmorata (VU), differing from the top three previously reported by Carpentieri et al. (2021) for small-scale fisheries in the eastern Mediterranean (C. plumbeus, M. mustelus and M. mobular). They also differ from the top three most abundant threatened species in both the western Mediterranean (Alopias spp., P. glauca and I. oxyrinchus) and central Mediterranean (C. plumbeus, M. mustelus and G. cemiculus). This regional variation in elasmobranch species composition may be driven by habitat or prey availability, or intensity of threats such as coastal development or fisheries activity, especially the lack of trawling activity in Northern Cyprus.

Three species that had previously been reported in Northern Cyprus, but were not found in this study were: Etmopterus spinax (Linnaeus 1758, velvet belly lanternshark), Galeus melastomus (Rafinesque 1810, blackmouth catshark) and Mustelus punctulatus (Risso 1827, blackspotted smoothhound; Oray and Karakulak, 2005; Çoker and Akyol, 2014). E. spinax and G. melastomus have typically been recorded in deep water trawls (Bengil and Basusta, 2018), and thus may be unlikely to have been caught in this SSF fleet where trawling is prohibited. M. punctulatus has apparently been shifting periodically in the Mediterranean since the 1920s (Colloca et al., 2017), and although it had previously been reported in the area of Cyprus under the effective control of the RoC Authorities (Hadjichrisophorou, 2006) there have been no reports since 1984. It is not surprising that our study did not find all 58 species considered to be present in waters of the whole island, because, compared to Northern Cyprus, the fisheries in the RoC controlled area are more diverse (with active trawl and pelagic longline fleets), because data availability for the latter region is far greater (partly due to the isolation of Northern Cyprus from many data collection frameworks), and because our study was restricted to include systematic methods and opportunistic observations during only recent years. In addition, it is worthy of note that fishermen from the Karpaz (Zone 3) region talk about the rare presence and capture of Sphyrna spp. (hammerhead sharks) but there have been no recent records in the area.

Although there were few species for which we had multiple measurements, clearly both adult and juvenile life stages of many species are present in coastal waters of Northern Cyprus. Many coastal elasmobranch species are known to use inshore nursery grounds with high productivity that offer protection from predators (Heupel and Simpfendorfer, 2002; Tiralongo et al., 2018b), and the large bycatch event of S. blainville of both juvenile and adult pregnant females, suggests there is a nursery ground for this species at least in the Karpaz region, with previous records supporting this (Bengil, 2022). From the Squatina size classes and birthing events reported, and the presence of new-born I. oxyrinchus it appears that Northern Cyprus may also be an important breeding and nursery area for these Critically Endangered and Endangered species.

A large proportion of the species found in Northern Cyprus are coastal – A. superciliosus, Bathytoshia lata (Garman 1880, brown stingray), Dasyatis marmorata (Steindachner 1892, marbled stingray), Taeniurops grabatus (Geoffroy St. Hilaire 1817, round stingray), D. pastinaca, A. bovinus and Raja asterias (Delaroche 1809, starry skate) – or found on the continental shelf and upper slopes, making them vulnerable to interactions with small-scale fisheries that are concentrated along the coast (Carpentieri et al., 2021). The majority of elasmobranch species caught by small-scale fisheries in the Mediterranean are demersal species (Carpentieri et al., 2021) caught in trammel nets or trawls, however, in Northern Cyprus trawling is not permitted, and the majority of fishers use trammel nets. Gear modifications or bycatch reduction technology could be considered to mitigate the impact of this fishery, however, a more detailed analysis of the drivers of bycatch is required to allow for greater planning and prioritisation of mitigation measures to more discrete métiers. Greater temporal analysis of catches could also help focus conservation efforts to seasonal métiers. With only Türkiye recognizing Northern Cyprus, it often falls outside international conventions, research and conservation networks which creates a challenging environment for funding the monitoring and management of the fishery.

Since the decision of fishers to retain (for trade or bait) or release was influenced by the presence of onboard observers and interaction with the authors, we have not presented data on survivorship and release rates after hauling. However, there is currently little incentive to release specimens. The relatively low vessel mortality rates (% dead on hauling) for many species illustrate the opportunity to drastically reduce bycatch mortality rates through a successful release programme in this fishery. While momentum is being gained from an ongoing education and awareness raising programme in collaboration with local authorities, the current legal trade in endangered elasmobranch species must be tackled. Recent legislative changes are welcomed but enforcement remains a challenge. Meanwhile, it is commendable and highly positive for marine conservation, that the authorities have been able to eliminate bottom trawling. However, small-scale fishers in Northern Cyprus are currently permitted to use drift nets which are banned across the rest of the Mediterranean because of their high impacts on large pelagic species including elasmobranchs. To our knowledge, drift nets are not in use and have never been noted on any vessels or in ports, yet, their use should be prohibited to avoid their possible future use.

A larger number of individuals were caught in coastal Zone 3, indicative of the higher number of active vessels and greatest observer coverage compared to other zones ( Figure 1 ). When looking at BPUE however, several hotspots around Mağusa (Famagusta Bay), west of Cape Koruçam and the eastern Karpaz region can be seen ( Figure 5 ). Existing MPAs ( Figure 1 ) appear to overlap with these areas of high elasmobranch BPUE and so may be well-placed for continued work to designate strictly protected areas within them. Indeed, two candidate Important Shark and Ray Areas have been identified in Northern Cyprus which also overlap with these hotspots and the Akdeniz MPA on the west coast, and the two Karpaz MPAs ( Figure 1 ; https://sharkrayareas.org/e-atlas/). In the management plans for these MPAs, that were primarily put in place to protect marine turtles, monk seals, seabirds and seagrass beds, fishing with set nets is prohibited to 30 m deep and to 1.5 km offshore, although this has never been enforced. Further analysis with larger observed effort will be useful in confirming such hotspots, refining MPA management plans, effectively integrating no-take zones to protect multiple taxa and developing new MPAs. Further studies should aim to increase onboard observation and fill the spatial gaps in our survey coverage, especially along the west and central north coasts.

The results of this study have been shared with the Northern Cyprus authorities and a proposal to expand the list of protected species and introduce the first no-take zones was accepted and policy came into effect in May 2023. Meanwhile, the strong reaction of some fishers participating in the project, sharing videos of themselves releasing threatened elasmobranch species, are an encouraging reaction to a programme of education delivered during this project. The recognition of the value and diversity of elasmobranchs among fishers is critical, so that retention moves from being the norm, to becoming a taboo, since enforcement of elasmobranch trade bans also has inherent challenges. Bycatch mitigation gear for dolphins (Snape et al., 2018b) and turtles has been trialled in this fishery, and could be expanded to elasmobranchs in certain métiers, or at least for fishers operating in MPAs.

This study has underlined the value of collaborating with SSFs to identify marine biodiversity. Results support the existence of important elasmobranch species which should be protected from expanding fishing pressure and trade, through the development of strictly protected areas/no-take zones, development of mitigation, education, and most importantly, implementing and policing the recent ban on elasmobranch trade and no-take zones. Although management of small-scale fisheries is extremely difficult, high community engagement, education and engaging fishers through SSF co-management, can effect change (Piovano et al., 2012). As data in this location continues to grow, species-specific analysis of critically endangered elasmobranchs must be undertaken to further inform conservation planning and management of fisheries. More fine-scale distribution data would also be of benefit, to improve the accuracy and identification of species richness hotspots, and reliability of management actions. With nearly half (44%) of Mediterranean elasmobranch diversity across less than 1% of its coastline, Northern Cyprus should be considered a regional conservation priority for this taxonomic group.

The original contributions presented in the study are included in the article/ Supplementary Material . Further inquiries can be directed to the corresponding authors.

The animal study was approved by University of Exeter Ethics Committee. The study was conducted in accordance with the local legislation and institutional requirements.

RTES, DB, BJG and ACB contributed to the conception and design of the study. MaO, EGTB, JLP, DB, CC, MeO, RTES, collected data. MaO, EGTB, JLP, RTES and ACB developed the first draft of the manuscript and all authors contributed to further drafts. All authors contributed to the article and approved the submitted version.