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Aka microterebrans sp. nov. (A) Ectosomal skeleton; (B) endosomal skeleton A. ruetzleri sp. nov.; (C) dermal membrane made of a layer of irregularly arranged oxeas; (D) irregular reticulum with stouter roundish meshes below the dermal membrane (arrow); (E) very thick spicule tracts showing a parallel course; skeletal organization of Aka maldiviensis; (F) ectosomal skeleton of the fistule; (G) peripheral part of the inner skeleton of the fistule; (H) supporting skeleton of the fistule. Scale bars: D, 200 μm; others, 1 mm.

Aka microterebrans sp. nov. (A) Ectosomal skeleton; (B) endosomal skeleton A. ruetzleri sp. nov.; (C) dermal membrane made of a layer of irregularly arranged oxeas; (D) irregular reticulum with stouter roundish meshes below the dermal membrane (arrow); (E) very thick spicule tracts showing a parallel course; skeletal organization of Aka maldiviensis; (F) ectosomal skeleton of the fistule; (G) peripheral part of the inner skeleton of the fistule; (H) supporting skeleton of the fistule. Scale bars: D, 200 μm; others, 1 mm.

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Three new species of the boring genus Aka, A. coralliirubri, A. microterebrans and A. ruetzleri are described respectively from the Mediterranean Sea, Celebes Sea and Caribbean Sea. The Indo-Pacific A. maldiviensis, known only for the structure of its epilithic fistules, is here described in more detail. The new species confirm that the sponges com...

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Context 1
... oxeas tangentially arranged, without forming regular network ( Figure 4A). Endosomal skeleton consisting of interconnected fibres (40-175 μm wide) of numerous, densely packed oxeas, forming elongated meshes (150-800 μm in diameter) ( Figure 4B) filled by numerous scattered oxeas. ...
Context 2
... oxeas tangentially arranged, without forming regular network ( Figure 4A). Endosomal skeleton consisting of interconnected fibres (40-175 μm wide) of numerous, densely packed oxeas, forming elongated meshes (150-800 μm in diameter) ( Figure 4B) filled by numerous scattered oxeas. ...
Context 3
... skeleton of fistules made of layer of loose, irregularly arranged oxeas ( Figure 4C). Under this dermal membrane the oxeas are arranged in paucispicular (3-5) tracts of fibres (50-150 μm wide) forming an irregular reticulum with roundish meshes (100-670 μm) ( Figure 4D). ...
Context 4
... skeleton of fistules made of layer of loose, irregularly arranged oxeas ( Figure 4C). Under this dermal membrane the oxeas are arranged in paucispicular (3-5) tracts of fibres (50-150 μm wide) forming an irregular reticulum with roundish meshes (100-670 μm) ( Figure 4D). This network increasingly stouter towards fistular lumen. ...
Context 5
... network increasingly stouter towards fistular lumen. In this zone tracts of 100-650 μm thickness and meshes 400- 1000 μm wide ( Figure 4E). In the basal part of fistules, inside the lower part of coral, very thick spicule tracts showing parallel course were observed ( Figure 4E). ...
Context 6
... this zone tracts of 100-650 μm thickness and meshes 400- 1000 μm wide ( Figure 4E). In the basal part of fistules, inside the lower part of coral, very thick spicule tracts showing parallel course were observed ( Figure 4E). Endosomal spicules irregularly arranged. ...
Context 7
... walls with a thin ectosomal skeleton formed by tangential, almost regular, reticulate meshes incorporating coral sand grains ( Figure 4F). Ectosome supported by fan- shaped, paucispicular (3-4) tracts of oxeas corresponding to peripheral part of inner skeleton ( Figure 4G). ...
Context 8
... walls with a thin ectosomal skeleton formed by tangential, almost regular, reticulate meshes incorporating coral sand grains ( Figure 4F). Ectosome supported by fan- shaped, paucispicular (3-4) tracts of oxeas corresponding to peripheral part of inner skeleton ( Figure 4G). Supporting skeleton of fistules consisting of long, stout, multispicular tracts of spicules (75-370 μm thick) forming irregular reticulum with circular or elongated meshes (113-603 μm wide) ( Figure 4H). ...
Context 9
... supported by fan- shaped, paucispicular (3-4) tracts of oxeas corresponding to peripheral part of inner skeleton ( Figure 4G). Supporting skeleton of fistules consisting of long, stout, multispicular tracts of spicules (75-370 μm thick) forming irregular reticulum with circular or elongated meshes (113-603 μm wide) ( Figure 4H). The skeleton inside the boring chambers is formed by disorderly and densely arranged spicules. ...

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... Porifera research has been carried out in the Indonesia region since the late 19 th century (Breitfuss 1898;Hardwicke 1822;Herklots & Marshall 1868;Kieschnick 1896;Marshall 1875;Poléjaeff 1883;Quoy & Gaimard 1833;Ridley 1884bRidley , 1885Schulze 1887;Sollas 1888;Thiele 1899;Topsent 1897) and early to mid-20 th century (Annandale 1924;Brøndsted 1934;Burton 1930b;Dendy 1913;Hentschel 1912;Ijima 1927;Lévi 1964;Reid 1968;Thiele 1900Thiele , 1903Vosmaer 1911) including surrounding regions (Lévi 1961;Wilson 1925). In addition, several modern ecological studies have been conducted on Indonesian sponge communities (Amir 1992;Bell & Smith 2004;Rovellini et al. 2019;van Soest 1989van Soest , 1990de Voogd et al. 1999de Voogd et al. , 2004, including taxonomic studies with novel species descriptions (Alvarez et al. 2016;Azzini et al. 2007;Becking 2013;Calcinai et al. 2006Calcinai et al. , 2007Calcinai et al. , 2013Calcinai et al. , 2017aLim & Setiawan 2021;Marlow et al. 2021;Muricy 2011;van Soest et al. 2021;van Soest & de Voogd 2015;de Voogd 2003de Voogd , 2004de Voogd & van Soest 2002, 2007de Weerdt & van Soest 2001). These studies give significant information on sponge diversity and distribution throughout the Indonesian archipelago. ...
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... According to the findings of Calcinai et al. (2003Calcinai et al. ( , 2004, it is possible to attribute the boring agent of the Sciacca red coral to a species of the genus Siphonodictyon based on the micro-layered concentric pattern reaching the centre of the pits. In the Mediterranean Sea, two species of Siphonodictyon, Siphonodictyon insidiosa (Johnson, 1899) and Siphonodictyon coralliirubri Calcinai, Cerrano and Bavestrello, 2007, are recognized (Calcinai et al. 2007). The average size of the pits suggests that, very likely, S. coralliirubri is responsible for the bioerosion recorded in the Sciacca red coral. ...
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... Unfortunately, it was not possible to remove fragments of etched material for an accurate microscopical analysis. Nevertheless, the large excavations extending to the entire scleraxis leaving free only a thin sheet of carbonate are very similar to those recorded in dead red coral from the Alboran Sea and attributed to Delectona ciconiae Bavestrello, Calcinai and Sarà, 1996 [47], while the spongiosus micro-excavation are very similar to those of Siphonodictyon corallirubri (Calcinai, Cerrano and Bavestrello, 2007) [48]. ...
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... The resulting substrate chip is mechanically dislodged and discarded via cell transport and water currents (Warburton 1958;Rützler and Rieger 1973). Sponge chip diameters commonly range from 15 to 100 μm, but were most frequently assessed as being between 30 and 60 μm in diameter (Rützler and Rieger 1973;Fütterer 1974;Rützler 1975;Schönberg 2000;Calcinai et al. 2002Calcinai et al. , 2007. They are therefore predominantly part of the silt fraction of the sediment and are most abundant in the coarse silt fraction (Table 1; e.g. ...
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Bioeroding sponges are important macroborers that chemically cut out substrate particles (chips) and mechanically remove them, thereby contributing to reef-associated sediment. These chemical and mechanical proportions vary with elevated levels of partial pressure of carbon dioxide (pCO2). To assess related impacts, the morphometric parameters “chip diameter” and “etching fissure width” were analyzed for Cliona orientalis Thiele, 1900, hypothesizing that their dimensions would differ with different pCO2 exposures (72 h at ca. 400, 750 and 1700 μatm). Under ambient conditions, we obtained a mean chip diameter of 21.6 ± 0.7 μm and a mean fissure width of 0.29 ± 0.01 μm. Chips were evenly distributed across the medium and coarse silt fractions regardless of treatment. We could not find a reliable pCO2 treatment effect for chip diameter and fissure width, but we observed strong data variability not related to our key questions. A hierarchical data design further reduced the test power. Fissure width was the more sensitive, but also more variable parameter. Sample size analyses nevertheless indicated that we had processed enough data. Thus, we reject our scenario of an increase in fissure width and consequent reduction in chip size to explain why chemical sponge bioerosion increases more strongly than the mechanical counterpart. Instead, we propose that a lowered ambient pH may favor respiratory acid build-up in the sponge tissue, possibly leading to a less localized bioerosion, causing bias towards more chemical bioerosion. Overall, this does not seem to affect the morphometry of sponge chips and the quality of sponge-generated sediment.
... Alboran Sea, 100-200 m, from Corallium rubrum community, but maybe not in the coral), Maldonado(1992, as Cliona, Alboran Sea, 70-120 m, in Corallium rubrum),Cruz-Simó (2002, as Cliona, Canary Islands, depth not stated, in Dendrophyllia ramea),Hansson (1999, Scandinavia, depth and substrate not specified) Siphonodictyon corallirubri(Calcinai et al., 2007b) Tyrrhenian Sea, depth unknown, in Corallium rubrum AM:Calcinai et al. (2010, Mediterranean, depth not stated, in precious coral) Madeira, depth not stated, in bivalve shells AM:? Topsent (1904, as Cliona labyrinthica),? Alander (1942, as Aka labyrinthica, W Norway and Sweden, 85-300 m, in Lophelia),Melone (1965, as Cliona labyrinthica, 60-75 m, Strait of Bonifacio in Corallium rubrum), ? ...
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... n. are, on average, considerably larger than in E. cretacea could be explained by fusion of several smaller chambers. On the other hand, even though there are at least some cases where extant species of Siphonodictyon develop from one growth type into another (Calcinai et al., 2007), such fusion of chambers would be quite unusual for Siphonodictyon (C. H. L. Schönberg, personal communication, 2017). ...
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The dependence of skeletal alteration on time spent in the taphonomic active zone (TAZ) can generate a taphonomic clock, which can be used to quantify scales of time averaging and rates of skeletal production and recycling in the fossil record. However, the strength of the taphonomic clock is variable in present-day shallow marine environments and it is unclear how this strength varies with depth. Here, we assess the strength of the taphonomic clock in Nautilus macromphalus dead shells that were collected in cool-water, sediment-starved, epi- and mesobathyal environments off New Caledonia and range in postmortem age from few decades to several millennia. We find that, first, differences in the onset and extent of alteration states in the epibathyal zone (< 750 m) segregate well-preserved shells with red stripes (less than ∼ 200 years) from encrusted shells with faded colors and extensively bored by sponges (∼ 400 years), and from strongly fragmented, bored, and coated shell relicts (> 1,000 years). The onset of dissolution and formation of clay-goethitic coating occurs earlier (∼ 200 years) in the mesobathyal zone (> 750 m) than in the epibathyal zone. Clay-goethitic rims and boring infills show signs of microbial binding, pelletization, and coccolith dissolution and can represent nascent stages of glauconitization. Second, shells several centuries old show differences between right and left flanks (1) in the degree of encrustation and sponge bioerosion in the epibathyal zone and (2) in the extent of clay-goethitic coating in the mesobathyal zone. The upper flanks are ultimately removed in both depth zones, leaving relict half-shells that are millennia old. Third, the depth dependence of alteration can reflect a bathymetric decline in disintegration rates by heterotrophic borers due to a reduced productivity in the mesobathyal zone and an increase in dissolution rates due to the proximity of the aragonite compensation depth. The between-flank asymmetry in preservation implies (1) horizontal position of shells close to the sediment-water interface for several decades or centuries without being overturned or subjected to reworking and (2) shell exposure to two distinct taphonomic regimes separated by few centimeters, with the upper flank located in the TAZ and the lower flank facing a less aggressive sediment zone. The stable exposure of shells in the taphonomic active zone and slow alteration rates in bathyal environments allow discriminating between within-habitat time-averaged assemblages on one hand and environmentally or stratigraphically condensed assemblages on the other hand.
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... According to the latest available revision of coralligenous biodiversity (Ballesteros 2006), 142 species of sponges have been recorded associated with this community. Adding to this list the species recorded on the coralligenous of Apulia (Sarà 1968 Calcinai et al. 2007b) and the data of the present study, the total number of sponge species hitherto associated to the coralligenous community increases to 273 ( Table 2). ...
... As to the boring sponges, Cliona janitrix is indicated by Ballesteros (2006) and Calcinai et al. (2007b) as the key species in the bio-erosive processes involving Corallium rubrum, whereas Cliona viridis has the same role in the coralligenous matrix (Rosell et al. 1999). According to our data Cliona celata Grant, 1826, C. schmidtii (Ridley, 1881), Spiroxya corallophila (Calcinai, Cerrano & Bavestrello, 2002), S. heteroclita Topsent, 1896 and Siphonodictyon insidiosum (Johnson, 1899) may also be considered important in the bio erosive processes acting upon the coralligenous structure. ...
... 3 B)and Pachastrella monilifera Schmidt, 1868(Fig. 3 C)were generally recorded as massive but also described as insinuating by Pulitzer-Finali (1970, 1983)andCalcinai et al. (2007b). ...
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