Books by Manuel Ballesteros
Papers by Manuel Ballesteros

Mediterranean Marine Science, 2016
In this Collective Article on “New Mediterranean Biodiversity Records”, we present additional rec... more In this Collective Article on “New Mediterranean Biodiversity Records”, we present additional records of species found in the Mediterranean Sea. These records refer to eight different countries throughout the northern part of the basin, and include 28 species, belonging to five phyla. The findings per country include the following species: Spain: Callinectes sapidus and Chelidonura fulvipunctata; Monaco: Aplysia dactylomela; Italy: Charybdis (Charybdis) feriata, Carcharodon carcharias, Seriola fasciata, and Siganus rivulatus; Malta: Pomacanthus asfur; Croatia: Lagocephalus sceleratus and Pomadasys incisus; Montenegro: Lagocephalus sceleratus; Greece: Amathia (Zoobotryon) verticillata, Atys cf. macandrewii, Cerithium scabridum, Chama pacifica, Dendostrea cf. folium, Ergalatax junionae, Septifer cumingii, Syphonota geographica, Syrnola fasciata, Oxyurichthys petersi, Scarus ghobban, Scorpaena maderensis, Solea aegyptiaca and Upeneus pori; Turkey: Lobotes surinamensis, Ruvettus pretios...

First assessment of the rocky intertidal communities of Fildes Bay, King George Island (South Shetland Islands, Antarctica)
Polar Biology, 2015
Studies on Antarctic intertidal fauna are comparatively scarce compared to those from the sublitt... more Studies on Antarctic intertidal fauna are comparatively scarce compared to those from the sublittoral and the deep sea. In order to contribute to filling this gap in knowledge, during the 2006 BENTART Spanish Antarctic Expedition, we conducted a quantitative and qualitative study on the fauna inhabiting the intertidal rocky platform of Fildes (=Maxwell) Bay, King George Island (South Shetlands Archipelago). This platform is dominated in its middle and lower area by the brown algae Adenocystis utricularis, which covers ca. 100 % of the rocky surface. Three quantitative samples of 20 × 20 cm were taken in the community of A. utricularis, and one additional sample of 20 × 20 cm in the tide pools community dominated by the gigartinacean rhodophyceae Iridaea cordata. In addition, qualitative samples of invertebrates that live under the boulders were also taken. Quantitative samples yielded a total of 9950 individuals belonging to 41 species. Mollusks were the most abundant group with a total of 9522 specimens belonging to 11 species, being the gastropod Eatoniella kerguelenensis regularis the most abundant with 6763 specimens. Crustaceans were represented by 295 specimens and 7 species and annelid polychaetes with 133 specimens and 20 taxa. The Shannon diversity indexes calculated for the quantitative samples of A. utricularis ranged between 0.803 and 1.030 with values of 1.577 for the I. cordata community. Our study confirms that the intertidal area of Fildes Bay hosts a surprisingly rich community in terms of abundance and biodiversity.

Journal of the Marine Biological Association of the United Kingdom, 2006
Studies on the opisthobranch molluscs inhabiting the continental platform in areas not accessible... more Studies on the opisthobranch molluscs inhabiting the continental platform in areas not accessible by SCUBA diving are rare in the subtidal bottoms of the Iberian Peninsula. Data obtained from trawls are not adequate for quantitative analysis, due to the indirect method used and the high number of variables which affect the capture of the samples. However, with the aim of obtaining qualitative information on the opisthobranch molluscs inhabiting the subtidal trawling grounds off Blanes, we took more than 500 samples over a two-year period, on the non-commercial fraction of the trawling fisheries, finding opisthobranchs in 306 samples. We analysed samples from 16 trawling sites, at depths between 50 and 300 m, by using ten fishing boats of the ‘Confraria de Pescadors del Port de Blanes’. We obtained 4.295 specimens of opisthobranchs belonging to 29 species. From these, 4 are Cephalaspidea s.s., 1 Anaspidea, 1 Umbraculacea, 3 Pleurobranchacea and 20 Nudibranchia (3 Arminina, 3 Dendrono...
Spatial patterns and diversity of bryozoan communities from the Southern Ocean: South Shetland Islands, Bouvet Island and Eastern Weddell Sea
Systematics and Biodiversity, 2012
In this study, we report new data on the biodiversity and the geographic and bathymetric distribu... more In this study, we report new data on the biodiversity and the geographic and bathymetric distribution of bryozoans collected during the ANT XXI/2 cruise (November 2003 to January 2004) in the Eastern Weddell Sea and Bouvet Island, and during the Spanish Antarctic expedition ECOQUIM (January 2006) in the South Shetland Islands. Our data on distribution were analysed together with previous
Journal of Molluscan Studies, 2002

Journal of Molluscan Studies, 1996
Six valid species of the nudibranch genus Dendrodons Ehrenberg, 1831 inhabit the Atlantic Ocean, ... more Six valid species of the nudibranch genus Dendrodons Ehrenberg, 1831 inhabit the Atlantic Ocean, including the Mediterranean and Caribbean Seas. Dendrodons Umbata (Cuvier, 1804), Dendrodons grandiflora (Rapp, 1827), Dendrodons nigra (Stimpson, 1855) (immigrant from the Red Sea), Dendrodons krebsu (M6rch, 1863), Dendrodons senegalensis Bouchet, 1975 and Dendrodons warta Marcus & Gallagher, 1976. Additional data about the biology and geographical distribution of these species are presented^ New evidence suggests that other species assigned to the genus Dendrodons, Dendrodons racemosa Pruvot-Fol, 1951 and Dendrodons minima Pruvot-Fol, 1951, must be included in the genus Doriopsilla Bergh, 1880. Three new species of Dendrodons are described from the Northeastern Atlantic and West Africa; Dendrodons angolensis, Dendrodons guineana and Dendrodons herytra. The variable external morphology makes species recognition difficult Instead, the diagnostic characters utilised to separate species are the shape of the male cirrus hooks, the structure of the reproductive system and features of the egg-mass.

Acta Zoologica, 2012
Figuerola, B., Ballesteros, M. and Avila, C. 2013. Description of a new species of Reteporella (B... more Figuerola, B., Ballesteros, M. and Avila, C. 2013. Description of a new species of Reteporella (Bryozoa: Phidoloporidae) from the Weddell Sea (Antarctica) and the possible functional morphology of avicularia.-Acta Zoologica (Stockholm) 94: 66-73. A new species of cheilostome bryozoan, Reteporella rosjoarum sp. n., belonging to the family of Phidoloporidae is described from the Weddell Sea (Antarctica) and illustrated with binocular microscope and SEM micrographs. SEM has been used to observe the essential characters to describe the new species and to compare it to similar species. Three samples from three different stations were collected at 332-597 m depth during the ANT XXI ⁄ 2 cruise of the R ⁄ V Polarstern (AWI, Bremerhaven, Germany) using a bottom trawl. The new species is characterized by the presence of giant vicarious spherical avicularia. The distinctive morphology of these avicularia discriminates this species from Antarctica congeners. The variability in avicularian morphometrics demonstrates that intraspecific variation between localities (Austasen and Drescher Inlet) may exist. It is also interesting to relate the morphology of the avicularia with the possible functions in Cheilostomata.

The Mediterranean Sea is a hotspot for biodiversity and endemic species (between 4 % and 18 % of ... more The Mediterranean Sea is a hotspot for biodiversity and endemic species (between 4 % and 18 % of known species are endemic, depending on the group), but this environment, favourable for native species, also favours exotic species. To ascertain how biodiversity has been impacted by intense human activity, we surveyed the marine life in the Barcelona Forum bathing area, an artificial beach receiving large amounts of waste from neighbouring areas and effluent from the nearby sewage treatment plant. Despite such eutrophic influences and the replacement of natural substrates with artificial ones, a surprisingly rich marine biota of 514 species was found. The groups with most species identified were molluscs (176), fish (88), crustaceans (65) and algae (50). These results include 15 exotic species that have settled in this ecosystem, such as the sea hare Bursatella leachii and the polychaete Branchiomma luctuosum, and some iconic natives such as the Fan mussel (Pinna nobilis) and the Dusk...

Ecología química en el bentos antártico
ABSTRACT El estudio de las interacciones entre los organismos mediadas por sustancias químicas, l... more ABSTRACT El estudio de las interacciones entre los organismos mediadas por sustancias químicas, lo que se denomina ecología química, nos da información sobre la biología de las especies implicadas, el funcionamiento de la comunidad en que viven y nos puede proporcionar además el conocimiento de nuevas sustancias que pueden ser útiles para el ser humano. En el marco de nuestros estudios sobre la ecología química de los invertebrados antárticos, en el proyecto ECOQUIM (2003-2006) se recolectaron numerosas muestras biológicas a partir de las cuales se obtuvieron extractos y sustancias naturales de invertebrados antárticos. En el proyecto ACTIQUIM (2007-actualidad), se ha evaluado el papel ecológico de estos extractos y sustancias naturales mediante experimentos de ecología química in situ (repelencia, defensa ante distintos tipos de macro y micro-depredadores, toxicidad, citotoxicidad y actividad antifouling) y también se ha analizado la actividad antitumoral de algunos compuestos obtenidos recientemente. Nuestros resultados demuestran, entre otras cuestiones, que las defensas químicas en los organismos antárticos son elevadas en las esponjas, los cnidarios, los tunicados, los briozoos, los equinodermos y los moluscos y que por lo tanto, estos grupos pueden ser una fuente rica en metabolitos secundarios con bioactividad, tanto a nivel ecológico como farmacológico. The study of the relationships between organisms mediated by chemicals (chemical ecology) can provide valuable information about the biology of the species, their role within the community and our knowledge of new molecules potentially useful to human beings. Within our ECOQUIM Project (2003-2006), we collected many biological samples that allowed us to obtain extracts and bioactive natural products from Antarctic benthic invertebrates. During the ACTIQUIM Project (2007 and on) the ecological activities of the marine natural products obtained have been determined by in situ chemical ecology experiments (repellence, defence, toxicity, cytotoxicity and antifouling). Also, the pharmacological potential has been analyzed by testing the antitumoral activity of selected compounds recently described. Our results show that chemical defences are abundant in Antarctic sponges, cnidarians, echinoderms, bryozoans, tunicates and mollusks, and that these zoological groups can be considered as a rich source of bioactive secondary metabolites of importance both in ecology and in pharmacology.
Comunidades bentónicas de sustrato duro de la Isla de Formentera (Baleares)

Antarctic marine ecosystems are immersed in an isolated, relatively constant environment where th... more Antarctic marine ecosystems are immersed in an isolated, relatively constant environment where the organisms inhabiting their benthos are mainly sessile suspension feeders. For these reasons, physical and chemical biotic interactions play an essential role in structuring these marine benthic communities (Dayton et al., 1974; Orejas et al., 2000). These interactions may include diverse strategies to avoid predation (e.g. Iken et al., 2002), competition for space or food (e.g. Bowden et al., 2006) and avoiding fouling (e.g. Rittschof, 2001; Peters et al., 2010). For instance, in the marine benthos, one of the most extended effective strategies among sessile soft-bodied organisms is chemical defense, mediated by several bioactive natural products mostly considered secondary metabolites (e.g. Paul et al., 2011). The study of the “chemical network” (chemical ecology interactions) structuring the communities provides information about the ecology and biology of the involved species, the f...

Update of the catalog of opisthobranch mollusks (Gastropoda: Heterobranchia) from the Catalan Coa... more Update of the catalog of opisthobranch mollusks (Gastropoda: Heterobranchia) from the Catalan Coasts.—An extension of the checklist of opisthobranch species (Gastropoda: Heterobranchia) known from the Catalan Coasts is presented, based on numerous unpublished findings by the authors and reports confirmed with pictures posted in several Internet platforms. A total of 53 species are added to the previous catalog: 4 Cephalaspidea, 9 Runcinacea, 3 Anaspidea, 8 Sacoglossa and 29 Nudibranchia (11 Doridacea, 10 Aeo lidacea, 2 Dendronotacea and 6 Cladobranchia incertae sedis). Data for the different reports of each of these species are provided, along with some biological, distribution or taxonomical remarks of interest. Finally, an updated and taxonomically sorted list of all opisthobranch species known for Catalonia is provided, including a total of 257 species, of which 9 are basal Heterobranchia, 35 Cephalaspidea, 13 Runcinacea, 10 Anaspidea, 22 Sacoglossa, 8 Pleurobranchomorpha, 2 Umb...

Sea slugs in the broad sense (until recently called «opisthobranchs» and now within the Heterobra... more Sea slugs in the broad sense (until recently called «opisthobranchs» and now within the Heterobranchia subclass of the Gastropoda), have aroused the interest of naturalists since they began to be studied by the end of the XVIII century. Their colorful and varied shapes make them very attractive to be studied and photographed. Numerous are the published works written to spread the knowledge of the biodiversity of these molluscs in all marine regions, however very few studies have been carried out to know these species that inhabit highly anthropized areas, such as the interior of commercial ports, marinas, artificial substrates or the coastal areas of large cities. Here are the results of a biodiversity study of marine heterobranchs conducted along the coast of the city of Barcelona (Spain). Numerous specimens of 73 species of sea slugs have been found, belonging to 5 orders: 4 Aplysiida, 5 Cephalaspidea, 3 Runcinida, 47 Nudibranchia and 4 Pleurobranchida, and 1 superorder: 10 Sacogl...

First record of the gastropod Philinopsis wildpretii (Ortea, Bacallado and Moro, 2003) (Cephalaspidea, Aglajidae) in the Iberian Peninsula
Marine Biodiversity, 2016
The cephalaspidean gastropod Philinopsis wildpretii (Ortea, Bacallado and Moro, 2003) was first d... more The cephalaspidean gastropod Philinopsis wildpretii (Ortea, Bacallado and Moro, 2003) was first described for the Canary island of Gran Canaria (Ortea et al. 2003). Initially included in the genus Melanochlamys, it was later changed by the same authors to the genus Spinoaglaja (Ortea et al. 2013) based on external and shell morphology. Ornelas-Gatdula and Valdés (2012), using molecular and morphological data, consider Spinoaglaja to be a synonym of Philinopsis, as is also suggested by Camacho-García et al. (2014) in their phylogeny of the Aglajidae. WoRMS (accessed April 25, 2016) accepts this species under the name Melanochlamys wildpretii, and in the CLEMAMdatabase (accessed April 25, 2016) it is considered synonymous with Spinoaglaja aeci. According to the latest published molecular analysis, we consider this species as belonging to the genus Philinopsis until the status of Spinoaglaja is completely resolved in a future study. Since its description, only a few specimens have been observed, all in the Mediterranean Sea: only shells in the Greek islands (Manousis et al. 2012) and in the island of Malta (Romani and Pagli 2015), five living specimens in Sardinia (Trainito and Doneddu 2014), one in the French Mediterranean (Horst and Juan 2014) and one in Porto Cesareo (Lecce, Italy) (12/11/2015, F. Vitale, SalentoSommerso). In this paper we present the first record of its presence in the Iberian Peninsula waters, along the Catalan coast (NE Spain), one animal being observed in each of the following locations (Fig. 1): Tossa deMar (M. Ballesteros, 12/ 13/2011), Sa Tuna (Begur) (X. Salvador, 12/05/2013), Morro del Vedell (Palamós) (E. Madrenas, 10/03/2014) and Cap de Creus (E. Madrenas, 10/31/2015). These observations suggest that the species has a wider distribution than formerly known, and given that all records since its description have been in the Mediterranean Sea, it should be considered as a kind of Atlanto-Mediterranean species or, as some authors have found (Ornelas-Gatdula and Valdés 2012), the Canarian and Mediterranean specimens could correspond to two different sympatric cryptic species with minute external differences. According to the original description, the animals from the Canary Islands have digging habits and live in mud or sand bottoms with half-buried stones covered with turf of algae, between 4 and 15 m deep; they move swiftly on the sediment and are able to bury themselves within seconds. Mediterranean specimens were found in shallow sandy bottoms near rocks where the algae Acetabularia acetabulum and Liagora viscida were growing (Horst et al. 2014), and also on leaves and rhizomes of Posidonia oceanica and Cymodocea nodosa meadows with Caulerpa prolifera and C. cylindracea (Trainito et al. 2014). The specimens observed in the Catalan coast of the Iberian Peninsula have been found among rhizomes of Posidonia oceanica and masses of photophilic algae.. Communicated by V. Urgorri

PLOS ONE, 2016
Although several studies are devoted to determining the diversity of Antarctic heterobranch sea s... more Although several studies are devoted to determining the diversity of Antarctic heterobranch sea slugs, new species are still being discovered. Among nudibranchs, Doto antarctica Eliot, 1907 is the single species of this genus described from Antarctica hitherto, the type locality being the Ross Sea. Doto antarctica was described mainly using external features. During our Antarctic research on marine benthic invertebrates, we found D. antarctica in the Weddell Sea and Bouvet Island, suggesting a circumpolar distribution. Species affiliation is herein supported by molecular analyses using cytochrome c oxidase subunit I, 16S rRNA, and histone H3 markers. We redescribe D. antarctica using histology, micro-computed tomography (micro-CT), and 3D-reconstruction of the internal organs. Moreover, we describe a new, sympatric species, namely D. carinova Moles, Avila & Wägele n. sp., and provide an anatomical comparison between the two Antarctic Doto species. Egg masses in both species are also described here for the first time. We demonstrate that micro-CT is a useful tool for non-destructive anatomical description of valuable specimens. Furthermore, our high resolution micro-CT data reveal that the central nervous system of both Doto species possesses numerous accessory giant cells, suggested to be neurons herein. In addition, the phylogenetic tree of all Doto species sequenced to date suggests a scenario for the evolution of the reproductive system in this genus: bursa copulatrix seems to have been reduced and the acquisition of a distal connection of the oviduct to the nidamental glands is a synapomorphy of the Antarctic Doto species. Overall, the combination of thorough morphological and anatomical description and molecular analyses provides a comprehensive means to characterize and delineate species, thus suggesting evolutionary scenarios.
[Redescription of Baptodoris cinnabarina Bergh, 1884 (Opisthobranchia, Doridina, Platydorididae) and taxonomic discussion of other species of the genus Baptodoris Bergh, 1884]
Iberus, 1999
Triterpene glycosides from Antarctic sea cucumbers III. Structures of liouvillosides A4 and A5, two minor disulphated tetraosides containing 3-O-methylquinovose as terminal monosacharide units from the sea cucumber Staurocucumis liouvillei (Vaney)
Nat Prod Res, 2011
Two new minor triterpene glycosides, liouvillosides A(4) (1) and A(5) (2) have been isolated from... more Two new minor triterpene glycosides, liouvillosides A(4) (1) and A(5) (2) have been isolated from the sea cucumber Staurocucumis liouvillei living in Antarctic and Sub-Antarctic waters. The structures of the new glycosides were elucidated using extensive nuclear magnetic resonance (NMR) spectroscopy ((1)H- and (13)C-NMR spectroscopy, (1)H-(1)H COSY, 1D-TOCSY, HMBC, HSQC and NOESY), ESI-MS and MALDI TOF MS. The glycosides 1 and 2 are disulphated tetraosides having very rare 3-O-methylquinovose as terminal monosaccharide.
Second addition to the malacological fauna of the littoral of Garraf (NE of the Iberian Peninsula)
Iberus, 2008
Una nueva especie de Doriopsilla Bergh, 1880 (Mollusca: Nudibranchia) del Archipiélago de Cabo Verde
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Books by Manuel Ballesteros
Papers by Manuel Ballesteros