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Equinodermos (Asteroidea; Echinoidea) de la demarcación marina noratlántica española

  • Autores: Laura María García Guillén
  • Directores de la Tesis: M.E. Manjón-Cabeza (dir. tes.), María del Pilar Ríos López (codir. tes.)
  • Lectura: En la Universidad de Málaga ( España ) en 2023
  • Idioma: español
  • Tribunal Calificador de la Tesis: Gonzalo Giribet (presid.), Francisca Moya Ruiz (secret.), Luis José Míguez Rodríguez (voc.)
  • Programa de doctorado: Programa de Doctorado en Diversidad Biológica y Medio Ambiente por la Universidad de Málaga
  • Materias:
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  • Resumen
    • Echinoderms (Asteroidea; Echinoidea) North Atlantic Spanish Marine Sub-Division Spain is divided into five marine subdivisions: North Atlantic Spanish Marine Sub-Division (NAMD), South Atlantic Spanish Marine Sub-Division (SASD), Strait of Gibaltar and Alborán Sea Spanish Marine Sub-Division (GSMD), Levantine Balearic Spanish Marine Sub-Division (LBMD) and Canary Islands Spanish Marine Sub-Division (CMD). This study is framed within NAMD, which includes the Bay of Biscay and the Iberian waters of the Atlantic Ocean. NAMD is comprised between the coordinates 48° N and 36° N and the western limit is 11° W. NAMD includes the Avilés Canyon System (ACS), the Galician Bank (GB) and Cachucho (CA) (which are part of this thesis).

      These areas were studied by the projects LIFE + INDEMARES Avilés Canyon System, LIFE + INDEMARES Galician Bank, ECOMARG and SponGES, which were carried out all around the area. These are multidisciplinary projects where geomorphology, hydrology and biodiversity were studied. The importance of these areas resides in their high productivity and the existence of vulnerable habitats. In addition, due to these places being traditional fishing areas with overfishing, regularization is required. ECOMARG began in 2000, and their surveys were carried out throughout the entire demarcation, intensively on Le Danois Bank and adjacent areas in the CA, with exploratory sampling also carried out at stations in the ACS and BG. Its objective was the study of demersal and benthic communities, which led to the identification of vulnerable, threatened, or declining habitats through the identification of deep-sea sponge aggregates, cold water coral reefs, seamount communities, pennatulacean communities and burrowing megafauna (OSPAR, 2003). All these results contributed to the attribution of the status of marine protected area (Ministry of the Environment, Rural and Marine, 2011). In 2021, the second management plan was approved, requesting an extension of the protected area (Ministry of the Presidency, Relations with Parliament and Democratic Memory, 2021).

      As a result of the obligation to define measures for the conservation and protection of Spanish waters, the LIFE+ INDEMARES project was created in 2009 to carry out the inventory and designation of the Natura 2000 Network in Spanish marine areas. This project is carried out in 10 marine areas within 3 Spanish biogeographical regions: Atlantic Region, where the ACS and BG project are framed; Mediterranean Region and Macaronesian Region (Ministry of Agriculture, Food and Environment, 2009). The results of this project have led to the protection of 10 Site of Community Importance (SCI). ACS and GB are Site of Community Importance (SCI- ESZZ12003 and SCI-ESZZ1200 respectively), while CA is a Marine Protected Area (MPA) within the Natura 2000 Network and recognized as a Vulnerable Marine Ecosystem (VME), where Echinoderms play an important role in these communities and habitats.

      The actions that began with the LIFE+ INDEMARES project are still active today, as in 2017 the LIFE IP INTEMARES project was awarded and will continue until 2024. Its objective is to complete and improve knowledge for effective marine management of the Natura 2000 Network as well as to ensure the maintenance of protected areas, to know their status, and to monitor the application of current legislation.

      This thesis is based on the study of Echinoderm (Asteroidea; Echinoidea) in NAMD. For this purpose, it tries to inventory and review asteroid and echinoid fauna collected during the INDEMARES project in the ACS and compare the new findings with previous studies Official Spanish Checklist (IEEM: ¿Inventario Españnol de Especies Marinas¿, 2017, 2020) to update our knowledge on the diversity and distribution of the asteroids and echinoids species. During the surveys carried out within the project LIFE + INDEMARES-Avilés Canyon System (2010¿2012), a total of 445 specimens of asteroids, belonging to 25 species, were collected from 36 stations in a depth range between 266 and 1476 m. The most frequent species were Nymphaster arenatus (Perrier, 1881) (30.55%) and Henricia caudani (Koehler, 1895) (25%). After reviewing public datasets, two species should be considered as new records for Spanish waters: Radiaster tizardi (Sladen, 1882) and Henricia sexradiata (Perrier, 1881), and 4 species expand their bathymetric range: Novodinia pandina (Sladen, 1889), H. caudani, H. sexradiata (Perrier, 1881) and Myxaster perrieri Koehler, 1895. Regarding echinoids, a total of 287 specimens were sampled at 35 stations with a depth ranging between 510 and 1476 m. Twelve species of echinoids were identified, the most frequent being Araeosoma fenestratum (Thomson, 1872) (48.57%), Cidaris cidaris (Linnaeus, 1758) (42.85%) and Phormosoma placenta Thomson, 1872 (28.57%). One species should be considered as a new record in Spanish waters: Gracilechinus affinis (Mortensen, 1903); and the species Echinocardium flavescens expands its known bathymetric range (from 325 to 552 m).

      In addition, the present study tries to inventory and review asteroid and echinoid fauna collected during the INDEMARES project in the GB, which is a seamount located 180 km away from the Galician coast (Northwest Spain), in the Northeast Atlantic Ocean. The summit occurs at a depth between 650 and 1500 m, with the maximum depth reaching 4000 m (the abyssal bottom). The water masses, twists, eddies, and geomorphology favour the retention of nutrients and larvae, thus, being an area rich in nutrients. It is a hotspot of biodiversity and an important place for benthic communities. This study aims to inventory and review the asteroid and echinoid fauna collected during the LIFE+INDEMARES project in GB, compare the new findings with previous studies Official Spanish Checklist (IEEM: ¿Inventario Español de Especies Marinas¿, Manjón-Cabeza et al., 2017, 2020) and update our knowledge of the diversity and distribution of known species. In this study a total of 272 asteroid specimens belonging to 19 species were found at 45 stations in depths between 765¿1764 m, as part of the LIFE+INDEMARES-Galician Bank (2010¿2011) surveys. The most frequently encountered species were Plinthaster dentatus (Perrier, 1884), Peltaster placenta (Müller & Troschel, 1842) and Henricia caudani (Koehler, 1895). Circeaster americanus (A.H. Clark, 1916) and Hymenaster giboryi (Perrier, 1894) are new observations from this area. For several species, including Henricia caudani, Pedicellaster typicus M. Sars, 1861, Podosphaeraster thalassae Cherbonnier, 1970 and Hymenaster giboryi, known bathymetric range has been extended. In case of echinoids, a total of 139 echinoids belonging to 8 different species were sampled from 27 stations, between 744 and 1764 m depth. The most frequent species were Cidaris cidaris (Linnaeus, 1758) and Sperosoma grimaldii Koehler, 1897. After public datasets analysis, it turns out that one species should be considered as a new record in Spanish waters: Echinocyamus grandiporus Mortensen, 1907, and two for the North Atlantic Spanish Marine Sub-Division in IEEM: Holanthus expergitus (Lovén, 1871) and Plexechinus hirsutus Mortensen, 1905. Besides, the bathymetric range of Echinocyamus pusillus (O.F. Müller, 1776) was extended to 1313 m depth.

      So far, it has been possible to make a study on macro and megafauna in the Avilés Canyon System, during several oceanographic cruises carried out from 2009 to 2017. The biodiversity of ACS is summarized and its description is herein updated after sampling surveys (ECOMARG, INDEMARES, SponGES, INTEMARES) conducted by the Spanish Institute of Oceanography (IEO). This study has updated previous knowledge in the canyon area from past national and international projects, their reports and publications, as well as data collected in the context of regional projects designed to gain new insight into the diversity of marine invertebrates and fishes from the ACS. Samples were taken using a range of sampling gears (Rock dredge, Beam trawl, Trawl gear GOC-73, Suprabenthic sledge, Box corer and Remoted operated vehicle), from 55 to 2291 m in depth. A total of 1015 species were identified at the ACS, and among them 74 correspond to Filum Echinodermata. As a result of the research projects carried out in the area in the last fifteen years, important information is now available, which suggests that the ACS houses a large number of species with a high ecological value, that it represents a biodiversity hotspot in terms of the presence of sponge aggregations and coral reefs in certain regions, and that it sustains important fisheries due to the abundance of commercial species. Given the relevance of the species and habitats occurring in the ACS, there is a need to implement a conservation and management plan of the area in order to maintain habitats in good state of preservation.

      A first ecological approach is included in this work, which analyzes the fauna composition of the community of brittle and sea stars associated with sponge aggregations located in Avilés Canyons System and El Cachucho, Marine Protected Area (MPA). Diverse sampling methods were used depending on bottom morphology, such as rock dredges and specific samplers for sedimentary bottoms, mainly beam trawl models. These fields are made up of sponge and coral species that build a very appropriate substrate for the proliferation of benthic species, which together create VME that are highly relevant for management and conservation. Among these benthic species, echinoderms are of great interest due to their value as indicators of good habitat. In total, 1261 specimens were collected (934 brittle stars and 327 starfishes), belonging to 42 species (28 ophiuroids and 14 asteroids) from INDEMARES AVILÉS, ECOMARG, and SponGES projects surveys. Specimens were distributed among four sponge aggregations (F: fields) that were considered according to the sponge records obtained in the same stations 36). These fields were defined and named based on the five most common sponge species: Aphrocallistes beatrix Gray, 1858 and Regadrella phoenix Schmidt, 1880 (F1: Avilés Canyon); Pheronema carpenteri (Thomson, 1869) (F2: Intraslope basin of Le Danois Bank); Asconema setubalense Kent, 1870 (F3: Le Danois Bank); and Neoschrammeniella aff. bowerbankii (Johnson, 1863) (F4: Corbiro Canyon). Faunistic results show that Ophiactis abyssicola (M. Sars, 1861) (55.55% occurrence), Brisinga endecacnemos Asbjørnsen, 1856, Ophiolycus purpureus (Düben & Koren, 1846), and Peltaster placenta (Müller & Troschel, 1842) (33.33%) were the most frequently found species in F1; Psilaster andromeda (Müller & Troschel, 1842) (80%), Pseudarchaster parelii (Düben & Koren, 1846) (60%), and Nymphaster arenatus (Perrier, 1881) (46.67%) in F2; Ophiura carnea Lütken, 1858 (71.43%) and Ophiacantha smitti Ljungman, 1872 (42.86%) in F3; and Ophiacantha densa Farran, 1913, and Henricia caudani (Koehler, 1895) (100%) in F4. The asteroid and ophiuroid species collected seem to be composed of four different communities that fit to areas with morphological and biological features, related to the presence of species specialized in the use of the resources they can find there. In general, the abiotic factor controlling this structure is depth. This assemblage structure, which favors the dissimilarity between the canyons and the bank, is not so clear, since the deepest stations are located on the intra-slope basin of ¿El Cachucho¿, Marine Protected Area (MPA), therefore, using it a priori could lead to misunderstandings. Once the structure of the echinoderm community was known, we compared the expected and obtained results to analyze evidence which should prove the existence of any association between echinoderms and sponges, which enabled us to refute the hypothesis.

      Other relevant objectives have been proposed in order to make a new reliable catalog for NAMD (1) to analyze the open access asteroids and echinoids data base reliability, (2) to compare these results with the IEEM including the new SELUMA own dataset and their georeferenced.

      This reliability analysis was made based on following criteria: 1) records published in scientific journals; (2) records from prestigious international institutions; (3) "Human observations" records have only been taken into account when they come from data reviewed by an internationally renowned Echinoderm specialist.

      The databases analysed for this purpose were open-access databases: Global Biodiversity Information Facility, GBIF.org. (2022), Ocean Biodiversity Information System, OBIS (2022), the Official Spanish Checklist (IEEM: ¿Inventario Español de Especies Marinas¿. Asteroidea: Manjón-Cabeza et al. 2017, 2020) as well as it was reviewed all the bibliography which compound the main surveys in the area since 19th: Lightning and Porcupine (Wyville-Thomson, 1873); Challenger (Aggasiz, 1881; Sladen, 1889), Travailleur and Talisman (Perrier 1885, 1894; Bernard, 1895; Mortensen, 1927). Hirondelle (Perrier, 1891; Koehler, 1921); Caudan (Koehler, 1895); Danish Ingolf-expedition (Liberkind 1935, Mortensen 1903, 1907); Princesse Alice (Koehler, 1909, 1921); Huxley (Morgan, 1913); Michael Sars (Grieg, 1932); Thalassa (Cherbonier, 1969, 1969-1970); Noratlante (Cherbonier y Sibuet, 1972); Hespérides 76 (Monteiro-marques, 1980); BIOGAS (Laubier y Monniot 1985; Sibuet, 1977); Cantábrico 83 (López-Ibor, 1987); COCACE (Louzao et al., 2010); EUSKALBENTOS I, II,III (Martínez et al., 2005, 2006, 2007); ITSASTEKA (Garmendia et al., 2019); BURATO 4240, GRAN BURATO y GALINCLIMARCH (Salgado-Peniza, 2016). In addition, other works as Inventario dos Equinodermos de Galicia (Echinodermata) (Besteiro and Urgorri, 1988); (included in Besterio and Urgorri, 1988: Acuña et al., 1984; Fischer-piette and Seoane, 1962, 1963; Gili et al., 1979; Guerra et al., 1986; Míguez, 1980, 1984; Mora, 1980; Olaso, 1979, 1980, 1983; Olaso and Pereda, 1982; Polo et al., 1979; Román and Pérez, 1982; Susaeta, 1913). Cristobo et al., 1988; Míguez, 1997.

      In general terms, this study was motivated by the need to have a frame of reference with which to compare or simply consult the results obtained in the different projects. The percentage of reliability in the study of the public databases has turned out to be more reliable than initially expected, 94.66% for asteroids and 97.05% for echinoids. However, these values are lower in IEEM, even though a revision of the one published in 2017 and 2020 (uncorrected synonymies) has been performed. These values were 89.77% for asteroids and slightly better for echinoids (92.30%).

      In the case of asteroids, unreliable records from databases accounted for 41.19% of the total records. These contained 19 species, of which 10 were not included in the reliable records (Anseropoda placenta (Pennant, 1777); Asterias amurensis Lutken, 1871; Asterias forbesi (Desor, 1848); Asterias rubens Linnaeus, 1758; Asterina gibbosa (Pennant, 1777); Ceramaster vincenti (Perrier, 1894); Echinaster (Echinaster) sepositus (Retzius, 1783); Henricia sanguinolenta (O.F. Müller, 1776); Marthasterias glacialis (Linnaeus, 1758) and Neomorphaster margaritaceus (Perrier in Milne-Edwards, 1882). In the case of Ceramaster vicenti, the search for this species in the various publications and WoRMS (2022) has been unsuccessful, although it was recorded by Cherbonier (1969) in his work on the samples collected in the Thalassa campaign. In fact, it may be an error or a synonymy (although not referred to) of the former Archaster echinulatus (Perrier, 1875), now Cheiraster (Barbadosaster) echinulatus (Perrier, 1875), but it is a species with an exclusively Caribbean distribution (GBIF, 2022). It is most likely that the listing given by Cherbonier (1969) was an error in the specific name, as it could refer to Ceramaster grenadensis, or to Peltaster placenta, which at the time was identified as Ceramaster placenta (Perrier, 1885), or Pentagonaster vicenti (Müller and Troschel, 1842), both synonymies of its currently accepted name. In view of this dilemma, this species will not be considered reliable and is excluded from the list, therefore we will focus on the remaining 9 species.

      Of these 9 species, 6 are recorded in the literature (Anseropoda placenta, Asterias rubens, Asterina gibbosa, Echinaster (Echinaster) sepositus, Marthasterias glacialis and Neomorphaster margaritaceus), which confirms their reliability, and 3 others are definitely unreliable in the D-NOR (Asterias amurensis, Asterias forbesi and Henricia sanguinolenta).

      On the other hand, the rest of the results obtained in the bibliographic searches (600 records) provide 9 more reliable species (Astropecten aranciacus (Linnaeus, 1758); Astropecten platyacanthus (Philippi, 1837); Chaetaster longipes (Bruzelius, 1805); Freyastera sexradiata (Perrier, 1885); Henricia caudani (Koehler, 1895); Henricia oculata (Pennant, 1777); Pedicellaster typicus M. Sars, 1861; Poraniomorpha (Poraniomorpha) hispida (M. Sars, 1872) and Tethyaster subinermis (Philippi, 1837)).

      As for the IEEM we started from 62 of which 20.96% of the species were not recorded in public databases or in the literature. However, the incorporation of our data validates 4 of them (Hacelia attenuata Gray, 1840; Novodinia pandina (Sladen, 1889); Odontaster mediterraneus (von Marenzeller, 1893); Persephonaster patagiatus (Sladen, 1889)) therefore the degree of reliability of the remaining 9 species included in the IEEM should be considered.

      In the case of echinoids, unreliable records from databases constituted 77.82% of the total records. These contained 9 species, 7 of which were not included in the reliable records (Echinocardium cordatum (Pennant, 1777); Echinocardium mortenseni Thiéry, 1909; Echinus esculentus Linnaeus, 1758; Paracentrotus lividus (Lamarck, 1816); Psammechinus miliaris (P.L.S. Müller, 1771); Spatangus purpureus O.F. Müller, 1776; Sphaerechinus granularis (Lamarck, 1816)). In the case of Echinocardium mortenseni, it appears in MNHN as of doubtful identification with the code MNHN-IE-2002-1171, obtained in the Thalassa expedition (1968), west of Galicia. Therefore, it should be considered as unreliable.

      Of these 7 species, 6 are validated by finding their records in the literature review (Echinocardium cordatum, Echinus esculentus, Paracentrotus lividus, Psammechinus miliaris, Spatangus purpureus and Sphaerechinus granularis). The remaining 11 species were only found as D-NOR records in the literature (495 in total): Echinocardium flavescens (O.F. Müller, 1776); Echinocardium pennatifidum Norman, 1868; Genocidaris maculata A. Agassiz, 1869; Gracilechinus elegans (Düben & Koren, 1844); Histocidaris elegans (A. Agassiz, 1879); Hygrosoma luculentum (A. Agassiz, 1879); Peripatagus cinctus Koehler, 1895; Plexechinus hirsutus Mortensen, 1905; Pourtalesia miranda A. Agassiz, 1869; Spatangus raschi Lovén, 1869 and Stereocidaris ingolfiana Mortensen, 1903.

      In the list of echinoids from IEEM (Manjón-Cabeza et al., 2017, 2020), 27 species were catalogued, of which only 4 were not recorded in public databases or in the literature (33 species). However, the incorporation of our data validates 1 of them (Histocidaris purpurata (Thomson, 1872)), so that only 3 IEEM species should be considered as unreliable.


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