Raja undulata 

Scope: Global

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Taxonomy [top]

Kingdom Phylum Class Order Family
Animalia Chordata Chondrichthyes Rajiformes Rajidae

Scientific Name: Raja undulata
Species Authority: Lacepède, 1802
Regional Assessments:
Common Name(s):
English Undulate Skate, Undulate Ray
French Raie Brunette
Spanish Raya Mosaica
Taxonomic Source(s): Eschmeyer, W.N. (ed.). 2015. Catalog of Fishes. Updated 6 April 2015. Available at: http://researcharchive.calacademy.org/research/ichthyology/catalog/fishcatmain.asp. (Accessed: 6 April 2015).

Assessment Information [top]

Red List Category & Criteria: Endangered A2bd+3d+4bd ver 3.1
Year Published: 2009
Date Assessed: 2003-09-03
Needs updating
Assessor(s): Coelho, R., Bertozzi, M.,Ungaro. N. & Ellis, J.
Reviewer(s): Kulka, D.W., Valenti, S.V. & Fowler, S.L. (Shark Red List Authority)
The Undulate Ray (Raja undulata) is a medium-sized, inshore skate that has a patchy distribution in the northeast and eastern central Atlantic, with discrete areas where it may be locally common (southwestern Ireland, eastern English Channel, southern Portugal). It also occurs in the Mediterranean Sea, where it appears to be uncommon. The Undulate Ray occurs in shelf waters down to about 200 m depth, although it is more common in shallow waters. This species is taken as utilised bycatch by trawl, trammel net and other demersal fisheries. Its patchy distribution means that populations are widely separated, possibly with little exchange. In the areas where it is known to be locally common, available data suggest declines have occurred. Time series catch data from Tralee Bay, southwestern Ireland (where this species forms a discreet subpopulation) show that catches of this species have declined by 60-80% since 1981, although they fluctuate each year. The species has also traditionally been observed in English beam trawl surveys in the eastern English Channel, but has been absent in recent years. French landings of this species from the Celtic Seas have declined steeply during the past decade. Another major area of occurrence is southern Portugal, where it is the most common skate captured by the mixed species trammel net fisheries. Species-specific landings data are not collected, but overall landings of Raja species have decreased by 29% between 1988 and 2004 in this area. This medium-bodied skate has limiting life-history characteristics that make it more vulnerable to exploitation than smaller skate species (three year generation period of ~45 years). Given the Undulate Ray's life-history characteristics, documented species-specific declines in some areas and the declines in aggregate catch data, significant declines are also inferred to have occurred off southern Portugal. Although no specific data are available from western Africa, intensive artisanal and demersal trawl fisheries operate throughout its inshore range there. Due to the patchy distribution of this inshore species, and that it is thought to have declined in major areas of its range, it is assessed as Endangered.

Geographic Range [top]

Range Description:This species has a patchy distribution in the eastern Atlantic, including the Mediterranean Sea, with discrete areas where it may be locally common (southwestern Ireland, eastern English Channel, southern Portugal).

Northeast and eastern central Atlantic: occurs from southern Ireland and southwestern England to the Gulf of Guinea, including the Canary Islands (Coelho and Erzini 2006).

Mediterranean Sea: western Mediterranean, a few records along the coasts of Israel and Turkey (Serena 2005). It has been recorded in the Ria Formosa coastal lagoon on the south coast of Portugal (Coelho et al. 2002), an important nursery ground for many bony fishes (Erzini et al. 2002). Smaller specimens of R. undulata have been reported in this coastal lagoon which may suggest that this species also uses these sheltered areas as nursery grounds (Coelho et al. 2002).
Countries occurrence:
Algeria; Benin; Cameroon; Côte d'Ivoire; Egypt (Egypt (African part)); Equatorial Guinea (Equatorial Guinea (mainland)); France (Corsica, France (mainland)); Gambia; Ghana; Greece (Greece (mainland), Kriti); Guinea; Guinea-Bissau; Israel; Italy (Italy (mainland), Sardegna, Sicilia); Lebanon; Liberia; Libya; Mauritania; Morocco; Portugal (Portugal (mainland)); Senegal; Sierra Leone; Spain (Baleares, Canary Is., Spain (mainland)); Syrian Arab Republic; Togo; Tunisia; Turkey (Turkey-in-Asia); United Kingdom (Great Britain); Western Sahara
FAO Marine Fishing Areas:
Atlantic – northeast; Atlantic – eastern central; Mediterranean and Black Sea
Additional data:
Lower depth limit (metres):200
Upper depth limit (metres):1
Range Map:Click here to open the map viewer and explore range.

Population [top]

Population:There appears to be a discrete population of Raja undulata in Tralee Bay (ICES 2007). Angling records from Tralee Bay (Ireland) indicate a peak in records of undulate ray in 1981-1982, with lower (but stable) catches since then (ICES 2007). Undulate ray have traditionally been observed in English beam trawl surveys in the eastern English Channel, but have been absent for the most recent two years (ICES 2008). It is uncommon in the Bay of Biscay (ICES 2007). Determining accurate stock trends in surveys is problematic due to the patchy distribution of the species.

The species is also common off southern Portugal (Coelho and Erzini 2006), where it forms a separate population to that on the Portuguese west coast (Moura et al. 2007). Raja undulata appears to be uncommon in the Mediterranean Sea, with only occasional records (Bertozzi et al. 2003, Baino et al. 2001).
Current Population Trend:Decreasing
Additional data:
Population severely fragmented:No

Habitat and Ecology [top]

Habitat and Ecology:This skate occurs in shelf waters to about 200 m depth, on sandy and muddy substrates and appears to be more common in shallow waters (Stehmann and Bürkel 1984, Coelho and Erzini 2006). Smaller specimens have been reported in coastal lagoons (specifically in the Ria Formosa coastal lagoon on the South coast of Portugal), which suggests that this species may use these sheltered habitats as nursery areas (Coelho et al. 2002). Like other skates, reproduction is oviparous. Estimated asymptotic maximum size is110 cm total length (TL) (Coelho and Erzini 2002). Significant regional differences have been observed in size at first maturity of this species: In Portugal, females mature (50% maturity) at 76.2 cm TL in the Algarve (southern) region and at 83.8 cm TL in the Peniche (western) region, while males mature (50% maturity) at 73.6 cm TL in the Algarve and at 78.1 cm TL in Peniche. Age at first maturity for the Algarve population has been estimated to occur at 8.98 years in females and 7.66 years in males (Coelho and Erzini 2006). An annual reproductive cycle has been observed, but differences occur between the different populations: In southern Portugal this species breeds mainly during the winter (Coelho and Erzini 2006), while in the Portuguese west coast it breeds during the winter and the spring (Moura et al. 2007). These differences seem to be related to water temperature, with reproduction restricted to periods of colder water. The maximum observed age for this species is 13 years old (Coelho and Erzini 2002) but longevity has been estimated at 21-23 years (Coelho et al. 2002). The population on the South Portuguese coast has a very slow growth rate (k=0.11) (Coelho and Erzini 2002). Natural mortality for this population has been estimated to be 0.20-0.219 year-1 (Coelho et al. 2002).

Generation period for this species varies from 14.9-15.9 years in females and from 14.3-15.3 in males. Generation period was estimated using the following formula: average reproductive age = age at maturity + 0.5* (longevity - age at maturity).

R. undulata shows a dietary ontogenetic shift within three major length groups: 20-55 cm TL; 55-75 cm TL and 75-100 cm TL. This shift goes from small semi-pelagic to larger and benthic prey and from a generalised to a specialised diet. Shrimps and mysids are especially important in the diet of smaller specimens, while in larger specimens brachyuran crabs have an increased importance (Moura et al. 2008).

Use and Trade [top]

Use and Trade: Flesh marketed for human consumption. In Portugal, larger specimens can reach relatively high market prices (R. Coelho pers. obs.).

Threats [top]

Major Threat(s): This species is a bycatch of trawl, trammel net and other demersal fisheries operating within its range. Raja undulata has a patchy distribution and declines have been documented in areas where is it was formerly considered locally abundant. Catch records are available from two charter-angling vessels in Tralee Bay (southwestern Island), where this species forms a discreet population. These data show that catches of this species declined from a high of 80-100 fish per year in 1981 to 20-30 fish per year in the mid 1990s (a decline of 60-70%), before increasing to 40-60 per year in 2001-2003. Catches now appear to be declining again, with less than 20 fish recorded in 2005, although they fluctuate each year (ICES 2007). R. undulata has traditionally been observed in English beam trawl surveys in the eastern English Channel, but has been absent for the most recent two years (ICES 2008). ICES advice (2008) is now for no target fishing in the North Sea, English Channel and Celtic Seas (ICES 2008).

Raja undulata is captured in large quantities as bycatch in the mixed species trammel net fishery that operates off the southern coast of Portugal (Coelho et al. 2002). The species is retained and marketed for human consumption. It is mainly captured in shallow waters, with CPUE values in this fishery decreasing from 1.91 specimens/1000m net at 10-30m depth to 0.03 specimens /1,000 m net at more than 90 m depth (Coelho et al. 2005). Portuguese official fisheries statistics for landings of Raja spp. in the Algarve have decreased 29.1% between 1988 and 2004 (DGPA 1988-2004). Raja undulata is the most common skate species in this area and its size makes it more vulnerable to depletion than smaller skate species, and these declines may under-reflect changes in the population of this species (Erzini et al. 2001, Coelho et al. 2005).

It is also a known bycatch of the Spanish demersal trawl fleet operating in the Cantabrian Sea, southern Bay of Biscay, which targets a mixture of gadoids and flatfish at depths of 100-300 m over the continental shelf (ICES 2007). Species-specific French landings data for the Celtic Seas report 12 t of R. undulata in 1995, 6 in 1996, 10 in 1997, after which landings fell to 2 t in 1998, 1 t in 1999, to 0 t in 2000-2001 (ICES 2007).

This species' preference for shallow waters places it within the range of intensive artisanal coastal fisheries operating along the western coast of Africa (Walker et al. 2005). Although no specific details are currently available on catches, this species is presumably a utilised bycatch of these, and demersal trawl fisheries operating in this area. Exploitation of the continental shelf is also high in the Mediterranean Sea (Massuti and Moranta 2003).

Like many other larger skates, this species possesses life history characteristics that may increase vulnerability to exploitation, reduce rate of recovery and increase the risk of extinction: including delayed age at maturity, long generation time (14-15 years), low fecundity, and consequently slow population growth (Dulvy et al. 2000).

Conservation Actions [top]

Conservation Actions: There is no species-specific management in place for Raja undulata. Tralee Bay is voluntarily closed to commercial fishing to protect regionally important elasmobranchs such as R. undulata and angel shark, which are only found in localised populations on the Irish West coast (ICES 2007). Recreational fishing continues in this area.

The most recent ICES advice (2008) is for no target fishing in the North Sea, English Channel and Celtic Seas. Species-specific monitoring of catches and landings is required to determine the proportion of Raja undulata taken.

Classifications [top]

9. Marine Neritic -> 9.1. Marine Neritic - Pelagic
9. Marine Neritic -> 9.4. Marine Neritic - Subtidal Sandy
9. Marine Neritic -> 9.5. Marine Neritic - Subtidal Sandy-Mud
9. Marine Neritic -> 9.6. Marine Neritic - Subtidal Muddy
9. Marine Neritic -> 9.10. Marine Neritic - Estuaries
10. Marine Oceanic -> 10.1. Marine Oceanic - Epipelagic (0-200m)
13. Marine Coastal/Supratidal -> 13.4. Marine Coastal/Supratidal - Coastal Brackish/Saline Lagoons/Marine Lakes
3. Species management -> 3.1. Species management -> 3.1.1. Harvest management
5. Law & policy -> 5.1. Legislation -> 5.1.1. International level
5. Law & policy -> 5.4. Compliance and enforcement -> 5.4.1. International level

In-Place Research, Monitoring and Planning
In-Place Land/Water Protection and Management
  Conservation sites identified:Yes, over entire range
In-Place Species Management
In-Place Education
5. Biological resource use -> 5.4. Fishing & harvesting aquatic resources -> 5.4.3. Unintentional effects: (subsistence/small scale)
♦ timing:Ongoing    
→ Stresses
  • 2. Species Stresses -> 2.1. Species mortality

5. Biological resource use -> 5.4. Fishing & harvesting aquatic resources -> 5.4.4. Unintentional effects: (large scale)
♦ timing:Ongoing    
→ Stresses
  • 2. Species Stresses -> 2.1. Species mortality

1. Research -> 1.2. Population size, distribution & trends
1. Research -> 1.3. Life history & ecology
1. Research -> 1.6. Actions
3. Monitoring -> 3.1. Population trends

♦  Food - human
 Local : ✓   National : ✓  International : ✓ 

Bibliography [top]

Bauchot, M.L. 1987. Raies at autres batoidés. In: M. Fisher, M. Schneider and M.-L. Bauchot (eds), Fiches FAO d'Identification des Espècs pour les Besoins de la Peche. Méditerranée et Mer Noire. Zone de Peche 37. Revision 1. II, pp. 847-885. FAO, Rome.

Bertozzi, M., Pasolini P., Hemida F., Capapé C. and Tinti, F. 2003. Species diversity in Algerian skates. Proceedings of the 7th European Elasmobranch Association Meeting: 38. San Marino.

Breder, C.M. and Rosen, D.E. 1966. Modes of reproduction in fishes. T.F.H. Publications, Neptune City, New Jersey, USA.

Coelho, R. and Erzini, K. 2002. Age and growth of the undulate ray, Raja undulata, in the Algarve (southern Portugal). Journal of the Marine Biological Association of the United Kingdom 82(6): 987-990.

Coelho, R. and Erzini, K. 2006. Reproductive aspects of the undulate ray, Raja undulata, from the south coast of Portugal. Fisheries Research 81: 80-85.

Coelho, R., Bentes, L., Correia, C., Gonçalves, J.M.S., Lino, P.G., Monteiro, P., Ribeiro, J. and Erzini, K. 2002. Fisheries biology of the undulate ray, Raja undulata, in the Algarve (southern Portugal). NAFO Scientific Council Research Document.

Coelho, R., Erzini, K., Bentes, L., Correia, C., Lino, P.G., Monteiro, P., Ribeiro, J. and Gonálves, J.M.S. 2005. Semi pelagic longline and trammel net elasmobranch catches in the Algarve (southern Portugal): catch composition, catch rates and discards. Journal of Northwest Atlantic Fisheries Science 35: 531–537.

DGPA. 2001. Recursos da pesca. Série estatística. Direcção-Geral das Pescas e Aquicultura. 1988-2001. Available at: http://www.dg-pescas.pt/.

Dulvy, N.K., Metcalfe, J.D., Glanville, J., Pawson, M.G. and Reynolds, J.D. 2000. Fishery stability, local extinctions and shifts in community structure in skates. Conservation Biology 14: 283-293.

Erzini, K., Bentes, L., Coelho, R., Correia, C., Lino, P., Monteiro, P., Ribeiro, J. and Gonçalves, J. 2002. Recruitment of sea breams (Sparidae) and other commercially important species in the Algarve (Southern Portugal). Final Report. Commission of the European Communities, Project DG XIV Ref. 99/061. University of the Algarve, Faro.

Erzini, K., Puente, E., Stergiou, K., Hernando, J.A. (Coordinators). 2001. Trammel net selectivity studies in the Algarve (Southern Portugal), gulf of Cadiz (Spain), Basque country (Spain) and Cyclades islands (Greece). Final Report. Commission of the European Communities, Project DG XIV Ref. 98/014.

IUCN. 2009. IUCN Red List of Threatened Species (ver. 2009.2). Available at: www.iucnredlist.org. (Accessed: 3 November 2009).

Massuti, E. and Moranta, J. 2003. Demersal assemblages and depth distribution of elasmobranchs from the continental shelf and slope off the Balearic Islands (western Mediterranean). ICES Journal of Marine Science 60: 753-766.

Moura, T., Figueiredo, I., Farias, I., Serra-Pereira, B., Coelho, R., Erzini, K., Neves, A. and Gordo, L.S. 2007. The use of caudal thorns for ageing Raja undulata from the Portuguese continental shelf, with comments on its reproductive cycle. Marine and Freshwater Research 58: 983-992.

Moura, T., Figueiredo, I., Farias, I., Serra-Pereira, B., Neves, A., Borges, M.F. and Gordo, L.S. 2008. Ontogenetic dietary shift and feeding strategy of Raja undulata Lacepède, 1802 (Chondrichthyes: Rajidae) on the Portuguese continental shelf. Scientia Marina 72: 311-318.

Stehmann, M. and Bürkel, D.L. 1984. Rajidae. In: P.J.P. Whitehead, M.-L. Bauchot, J.-C. Hureau, J. Nielsen and E. Tortonese (eds), Fishes of the North-eastern Atlantic and Mediterranean, pp. 163-196. UNESCO, Paris.

Walker, P., Cavanagh, R.D., Ducrocq, M., and Fowler, S.L. 2005. Northeast Atlantic (Including Mediterranean and Black Sea). In: Fowler, S.L., Cavanagh, R.D., Camhi, M., Burgess, G.H., Cailliet, G.M., Fordham, S.V., Simpfendorfer, C.A. and Musick, J.A. (eds) (eds), Sharks, Rays and Chimaeras: The Status of the Chondrichthyan Fishes, pp. 71-94. IUCN/ SSC Shark Specialist Group, Gland, Switzerland and Cambridge, UK.

Citation: Coelho, R., Bertozzi, M.,Ungaro. N. & Ellis, J. 2009. Raja undulata. In: The IUCN Red List of Threatened Species 2009: e.T161425A5420694. . Downloaded on 30 July 2016.
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