Molecular phylogenies confirm the presence of two cryptic Hemimycale species in the Mediterranean and reveal the polyphyly of the genera Crella and Hemimycale (Demospongiae: Poecilosclerida)

Este artículo contiene 24 páginas, 4 figuras, 3 tablas.Background: Sponges are particularly prone to hiding cryptic species as their paradigmatic plasticity often favors species phenotypic convergence as a result of adaptation to similar habitat conditions. Hemimycale is a sponge genus (Family Hymedesmiidae, Order Poecilosclerida) with four formally described species, from which only Hemimycale columella has been recorded in the Atlanto-Mediterranean basin, on shallow to 80 m deep bottoms. Contrasting biological features between shallow and deep individuals of Hemimycale columella suggested larger genetic differences than those expected between sponge populations. To assess whether shallow and deep populations indeed belong to different species, we performed a phylogenetic study of Hemimycale columella across the Mediterranean. We also included other Hemimycale and Crella species from the Red Sea, with the additional aim of clarifying the relationships of the genus Hemimycale. Methods: Hemimycale columella was sampled across the Mediterranean, and Adriatic Seas. Hemimycale arabica and Crella cyathophora were collected from the Red Sea and Pacific. From two to three specimens per species and locality were extracted, amplified for Cytochrome C Oxidase I (COI) (M1–M6 partition), 18S rRNA, and 28S (D3–D5 partition) and sequenced. Sequences were aligned using Clustal W v.1.81. Phylogenetic trees were constructed under neighbor joining (NJ), Bayesian inference (BI), and maximum likelihood (ML) criteria as implemented in Geneious software 9.01. Moreover, spicules of the target species were observed through a Scanning Electron microscope. Results: The several phylogenetic reconstructions retrieved both Crella and Hemimycale polyphyletic. Strong differences in COI sequences indicated that C. cyathophora from the Red Sea might belong in a different genus, closer to Hemimycale arabica than to the Atlanto-Mediterranean Crella spp. Molecular and external morphological differences between Hemimycale arabica and the Atlanto- Mediterranean Hemimycale also suggest that Hemimycale arabica fit in a separate genus. On the other hand, the Atlanto-Mediterranean Crellidae appeared in 18S and 28S phylogenies as a sister group of the Atlanto-Mediterranean Hemimycale. Moreover, what was known up to now as Hemimycale columella, is formed by two cryptic species with contrasting bathymetric distributions. Some small but consistent morphological differences allow species distinction. Conclusions: A new family (Hemimycalidae) including the genus Hemimycale and the two purported new genera receiving C. cyathophora and Hemimycale arabica might be proposed according to our phylogenetic results. However, the inclusion of additional Operational Taxonomic Unit (OTUs) appears convenient before taking definite taxonomical decisions. A new cryptic species (Hemimycale mediterranea sp. nov.) is described. Morphologically undifferentiated species with contrasting biological traits, as those here reported, confirm that unidentified cryptic species may confound ecological studies.The research has been funded by MARSYMBIOMICS project (Spanish MINECO, CTM2013-43287-P), BluePharmTrain (FP7 People-INT, Ref. 2013-667786), and Grup Consolidat SGR-120, to Maria J. Uriz. Leire Garate benefited from a fellowship within the Benthomics project (Spanish MICINN, CTM-2010-22218-C02-01).Peer reviewe

Les esponges litorals

S'han identificat un total de 98 espècies d'esponges (11 Calcarea i 87 Demospongiae) en la zona litoral (O-55 m) de l'Arxipèlag de Cabrera, la qual cosa representa un nombre relativament alt d'espècies si es compara amb els catàlegs coneguts d'altres zones litorals considerades com a ben estudiades (p. e. litoral català Nord). Les anàlisis de classificació i d'ordenació permeten de diferenciar un grup d'espècies esciòfiles d'un altre grup format per dos subgrups: un d'espècies fotòfiles i un d'àmplia distribució batimètrica. Dins del grup esciòfil se separen les espècies de coves, les del coral· ligen i les comunes a coves, coral· ligen i comunitats d'algues esciòfiles. No apareixen grups que corresponguin a les esponges de les diferents comunitats fotòfiles i hemiesciòfiles. La irradiància que arriba al fons està correlacionada negativament amb el nombre d'espècies d'esponges. La biomassa d'esponges, pel contrari, es més alta en certes comunitats fotòfiles (p. e. la d'algues fotòfiles soprepasturades) i a les coves semifosques, que en comunitats amb irradiàncies intermèdies (p.e. comunitat d'algues esciòfiles)A total of 98 sponge species (11 Calcarea and 87 Demospongiae) were identified in the littoral zone around the Cabrera Archipelago. That represents a relatively high number of species compared with other well-known littoral areas of the Iberian Peninsula. Cluster and correspondence analysis differentiated a group of sciaphilous species from another group made up of two subgroups (photophilic species and those with a broad bathymetric distribution). Within the sciaphilous group three subgroups were distinguishable: species from caves, species exclusively dwelling in the coralligenous, and species common to caves, coralligenous, and sciaphilic algal communities. No groups corresponding to the different photophilic and hemi-sciaphilous communities were detected. Irradiance at the bottom level was negatively correlated with the number of sponge species. In contrast, the sponge coverage was higher in some photophilic (p.e. that of overgrazed photophilic algae) and semi-dark caves communities than in communities with mean levels of irradiance (e.g. sciaphilic algae biocoenosis

Grazing on fleshy seaweeds by sea urchins facilitates sponge Cliona viridis growth

7 páginas, 4 figurasWe studied possible interactions among invertebrate species and algal assemblages in a shallow sublittoral community of the Mediterranean Sea using a 2-step approach. First, we analysed the general pattern by correlation analysis. Thereafter, we experimentally assessed cause–effect relationships between the species or assemblages that were clearly related in the correlation study. The abundance of Cliona viridis (Schmidt 1862) was positively correlated with that of sea urchins and negatively correlated with fleshy algal cover. These relationships were confirmed by field experiments: treatments without fleshy algae, owing to both natural and simulated sea urchin grazing, promoted C. viridis growth significantly more than controls. Our results support the hypothesis that sea urchin grazing on seaweeds increases light availability at the basal stratum, which favours the primary production of the symbiotic zooxanthellae and thus C. viridis growth through nutrient transfer. We found a network of interactions with several signs, directions and strengths: a strong positive indirect interaction (‘facilitation’) between sea urchins and C. viridis, a negative direct interaction between sea urchins and seaweeds (‘predation’), a negative direct interaction between seaweeds and C. viridis (‘shading-interference’) and a weaker but significant direct interaction between C. viridis and Pione vastifica (‘space competition’). The multiple interactions observed suggest a cascade that involves 4 trophic levels: primary producers, herbivorous, carnivorous, and filterfeeders. This cascading process can have negative cryptic implications on the environment, as the excavating sponge C. viridis is a bio-eroder that may strongly impact the shallow sublittoral landscape by producing substrate weakening and instability. The knowledge acquired on the sign and strength of these multi-species interactions is useful to model and predict the responses of shallow benthic communities to anthropogenic disturbances (i.e. overfishing and eutrophication).This research was partially funded by grants from the European Community (SPONGES project COOP-CT-205-017800) and the CICYT (Spain) (INTERGEN, CTM2004-05265-CO2/MAR) to M.J.U.Peer reviewe

High genetic diversity, phenotypic plasticity, and invasive potential of a recently introduced calcareous sponge, fast spreading across the Atlanto‑Mediterranean basin

16 páginas, 6 tablas, 8 figuras.Sponges are considered poor invaders, and no genetic studies on introduced sponges have been performed up to now. Paraleucilla magna is the first calcareous sponge introduced to the Mediterranean and Northeastern Atlantic. The study aimed at investigating the genetic makeup and connectivity of the introduced populations of P. magna and at exploring signs of local phenotypic adaptation, to gain insight on the species invasive potential. Ten populations along the species introduction range (Brazil, Açores, Madeira, and continental Europe) were genetically characterized by using nine microsatellite markers. Most populations were genetically structured as suggested by significant Dst and Fst values, significant differences among populations (AMOVA) and the presence of private alleles. The analyzed populations belonged to three genetically homogeneous groups (K) according to the Bayesian algorithm (structure software) and the UPGMA dendrogram. Genetic diversity within populations was higher than expected. Recurrent introductions of non-randomly selected individuals from the native sources may have contributed to the heterozygote deficit found in all populations by forming pedigree structures with mating among relatives. Moreover, the species biological cycle was monitored in a population established on native Mediterranean assemblages (41°40′27″N, 2°47′25″E) and compared with the species cycle in other habitats. Contrasting life spans, growth habits, and reproduction cycles, depending on the habitat conditions, were recorded. To summarize, high genetic diversity, phenotypic local adaptation, and high reproduction rates altogether allow predicting the fast proliferation of P. magna in newly colonized regions and point to its strong invasive potential.This study has been partially funded by MARSYMBIOMICS project ref. CTM2013-43287-P (Spanish Ministry of Economy and Competitiveness (MINECO)—FECYT agency, and Consolidate Group Award 2009-SGR655 (Generalitat of Catalonian) to MJU. MG has benefited from a FPU fellowship from the Spanish MINECO and JF from a Marie Curie (EU) fellowship, associated, respectively, with the BENTHOMICS and BIOCAPITAL projects to MJU.Peer reviewe

La auditoría ética en trabajo social: un instrumento para mejorar la calidad de las instituciones sociales

La auditoría ética en trabajo social: un instrumento para mejorar la calidad de las instituciones sociale

Faunal affinities of the Sponges (Porifera) of the Balearic Islands with those of other biogeographical areas

An evaluation is made of the affinities of the marine sponge fauna of the Balearic Islands (158 species; Bibiloni, 1990) with those of other biogeographical areas and regions of the world, as well as with those of other Mediterranean areas (on the basis of the literature and interpreting the synonymies), by the simple method of calculating the percentage of Balearic species cited for the other areas considered. With respect to other Mediterranean areas, the greatest affinity of the Balearic sponge fauna is with those in the Gulf of Lyons (79.3%) and the Tyrrhenian Sea (56.8%); with respect to other areas in the world, the greatest affinity is with those in the Mauritanian-Senegalian (46.2%) and boreal regions (45.6%). If the results are compared with those of Vacelet (1980) for the Demospongiae of the whole Mediterranean, the sponges of the Balearic Islands show marked subtropical characteristics.Afinidades faunísticas de las esponjas (Poríferos) de las islas Baleares con las de otras áreas biogeográficas. Se estiman las afinidades de la fauna de esponjas marinas de las islas Baleares (158 especies; Bibiloni, 1990) con las de otras regiones y provincias biogeográficas mundiales, así como con las de otras regiones mediterráneas (a partir de la bibliografía e interpretando las sinonimias), mediante el simple procedimiento de calcular el porcentaje de especies de Baleares citadas para las demás áreas consideradas. La mayor afinidad de la fauna de esponjas de Baleares con las de otras áreas del Mediterráneo es con el golfo de León (79,3%) y el mar Tirreno (56,8%); cuando se consideran otras áreas mundiales, la mayor afinidad es con las regiones mauritano-senegaliense (46,2%) y boreal (45,6%). Si se comparan los resultados con los de Vacelet (1980) para las demosponjas de todo el Mediterráneo, las esponjas de las Baleares manifiestan un marcado carácter subtropical

Benthic assemblages in two Mediterranean caves: species diversity and coverage as a function of abiotic parameters and geographic distance

16 páginas, 5 figuras, 4 tablasBenthic assemblages of two Mediterranean submarine caves were compared. Species coverage and number of species were lower in internal (dark) communities than external. This feature was specially marked in the less illuminated cave. Ordination analyses performed on species coverage per community for each cave separately, distinguished several benthic communities from the outermost to the innermost zone of each cave. Cluster analyses on species coverage, taking into account all communities in both the caves, established similarities among communities: algal-dominated communities clustered according to the level of light received independently of the cave they inhabited, while animal-dominated communities were more similar within each cave than between the caves. Moreover, among the abiotic parameters measured irradiance was the only factor that clearly diminished from the entrance to the innermost zone in both the caves. In contrast, water movement and particulate organic matter varied di¡erently in each cave. Results indicate that the di¡erent topography, depth and geographic location of the two caves determine water movement, light penetration and nutrient availability along the caves. These factors are responsible for determining species abundance and diversity, as well as species growth habit in each community.This research was partially supported by INTERREG-IIIA K115C n 0123 (EU) and CICYT (Comision Interdepartmental de Ciencia yTechnología) REN2001-2312-CO3/MAR grants. It also benefited from Governments of Catalonia and Spain fellowships to R. MartíPeer reviewe

Endosymbiotic calcifying bacteria across sponge species and oceans

Este artículo contiene 14 páginas, 7 figuras, 1 tabla.From an evolutionary point of view, sponges are ideal targets to study marine symbioses as they are the most ancient living metazoans and harbour highly diverse microbial communities. A recently discovered association between the sponge Hemimycale columella and an intracellular bacterium that generates large amounts of calcite spherules has prompted speculation on the possible role of intracellular bacteria in the evolution of the skeleton in early animals. To gain insight into this purportedly ancestral symbiosis, we investigated the presence of symbiotic bacteria in Mediterranean and Caribbean sponges. We found four new calcibacteria OTUs belonging to the SAR116 in two orders (Poecilosclerida and Clionaida) and three families of Demospongiae, two additional OTUs in cnidarians and one more in seawater (at 98.5% similarity). Using a calcibacteria targeted probe and CARD-FISH, we also found calcibacteria in Spirophorida and Suberitida and proved that the calcifying bacteria accumulated at the sponge periphery, forming a skeletal cortex, analogous to that of siliceous microscleres in other demosponges. Bacteria-mediated skeletonization is spread in a range of phylogenetically distant species and thus the purported implication of bacteria in skeleton formation and evolution of early animals gains relevance.The research has been funded by MARSYMBIOMICS project (Spanish MINECO, CTM2013-43287-P) BluePharmTrain (FP7 People-INT, Ref. 2013- 667786), and Grup Consolidat SGR-120, to MJU. LG benefited from a fellowship within the Benthomics project (Spanish MICINN, CTM-2010-22218-C02-01).Peer reviewe

Snapshot of a Bacterial Microbiome Shift during the Early Symptoms of a Massive Sponge Die-Off in the Western Mediterranean

30 páginas, 7 figuras, 1 tabla.Ocean warming is affecting marine benthic ecosystems through mass mortality events that involve marine invertebrates, in particular bivalves, corals, and sponges. Among these events, extensive die-offs of Ircinia fasciculata sponges have been recurrently reported in western Mediterranean. The goal of our study was to test whether the temperature-related mass sponge die-offs were associated with or preceded by an early unbalanced bacterial microbiome in the sponge tissues. We took advantage of the early detection of disease and compared the microbiomes of healthy vs. early diseased I. fasciculata tissues. Our results showed a microbiome shift in early diseased tissues. The abundance of Gammaproteobacteria and Acidobacteria increased and that of Deltaproteobacteria decreased in diseased vs. healthy tissues. The change in community composition was also noticeable at the operational taxonomic unit (OTU) level. Diseased tissues contained more bacterial sequences previously identified in injured or stressed sponges and corals than healthy tissues. Bacterial diversity increased significantly in diseased tissues, which contained a higher number of low abundance OTUs. Our results do not support the hypothesis of one particular pathogen, whether a Vibrio or any other bacteria, triggering the Northwestern Mediterranean mass mortalities of I. fasciculata. Our data rather suggest an early disruption of the bacterial microbiome balance in healthy sponges through a shift in OTU abundances, and the purported consequent decline of the sponge fitness and resistance to infections. Opportunistic bacteria could colonize the sponge tissues, taking benefit of the sponge weakness, before one or more virulent pathogens might proliferate ending in the mass sponge die-off.AcknowledgmentsThis study has partially been funded by projects MarSymbiOmics (MINECO, I+D+I ofExcellence, CTM2013-43287-P), the Benthic Ecology Consolidate Team 2014-SGR-120(Generalitat de Catalunya), and BluePharm Train FP7 People-INT, to MU The work of PGwas supported by the Agence Nationale de la Recherche (ANR) through the projectsEUREKA (ANR-14-CE02-0004-01).Peer reviewe

Vertical transmission and successive location of symbiotic bacteria during embryo development and larva formation in Corticium candelabrum (Porifera: Demospongiae)

7 páginas, 8 figuras.This study reports on the transfer of heterotrophic bacteria from parental tissue to oocytes in the Mediterranean bacteriosponge Corticium candelabrum (Homosclerophorida) and the description of the successive locations of the microsymbionts during embryo development through transmission and scanning electron microscopy. Eight different types of symbiotic bacteria are described morphologically. These eight bacteria morphotypes are found in both adult individuals and larvae. Symbiotic bacteria are transferred to oocytes, but not to spermatocytes. Bacteria are first located at the oocyte periphery below a thick collagen layer and then they migrate to the oocyte cytoplasm, forming spherical clusters. After cleavage, the bacteria can be found in the free space between blastomeres but mainly accumulate at the embryo periphery below the follicular cells that surround the embryo. Once the blastocoel is formed, the symbiotic bacteria move to this central cavity where they actively divide by bipartition, increasing their number considerably. Many examples of phagocytosed bacteria in the proximal zone of the larval cells are observed at this stage. Consequently, bacteria may represent a complementary source of energy for free larvae and settlers before they are able to capture food from the surrounding water.This study was partially funded by the project INTERGEN CTM2004-05265-C02-02/MAR from the CICYT (Spain)Peer reviewe