Deep-Sea Research Part I 131 (2018) 75–86

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Deep-Sea Research Part I

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Sponge assemblages on the deep Mediterranean continental shelf and slope T

(Menorca Channel, Western Mediterranean Sea)
⁎
Andreu Santína, , Jordi Grinyóa, Stefano Ambrosoa, Maria J. Urizb, Andrea Goria,
Carlos Dominguez-Carrióa, Josep-Maria Gilia
a
Institut de Ciències del Mar (ICM-CSIC), Barcelona, Spain
b
Centre d’Estudis Avançats de Blanes (CEAB-CSIC), Blanes, Girona, Spain

A B S T R A C T

Sponge assemblages on continental shelves and slopes around the world have been known about for centuries.
However, due to limitations of the traditional sampling systems, data about individual sponge species rather
than assemblages have been reported. This study characterizes sponge assemblages over a wide bathymetric
range (~50–350 m depth) and covering the entire continental shelf and the upper slope of the Menorca Channel,
an area soon to be declared a Marine Protected Area (MPA) as part of the Natura 2000 Network. Quantitative
analysis of 85 video-transects (a total linear distance of 75 km), together with representative collections to
confirm species identifications, allowed us to discriminate six major assemblages. Differences in the assemblages
mainly corresponded to differences in substrate type and depth. On the inner continental shelf, a semi-scia-
philous Axinellid assemblage dominated the rocky outcrops. Maërl beds on the inner continental shelf were
dominated by Haliclona (Reniera) mediterranea, whereas the horny sponge Aplysina cavernicola and several other
haliclonids mostly dominated maërl beds and rocky substrates of the outer shelf. Soft sediments on the shelf
break hosted a monospecific Thenea muricata assemblage, whereas rocky substrates of the shelf break were
characterized by a mixture of encrusting, columnar and fan-shaped sponges. Finally, the upper slope was
dominated by Hamacantha (Vomerula) falcula and the hexactinellid Tretodictyum reiswigi. Overall, sponge di-
versity showed its highest values above the shelf break, plummeting severely on the upper slope. Despite this
diversity decrease, we found very high densities (> 70 ind./m2) of sponges over vast areas of both the shelf
break and the upper slope.

1. Introduction In general terms, sponge assemblages show a high variability in

Sponges play a significant role in the functioning of marine benthic
ecosystems, (Gili and Coma, 1998; Van Soest et al., 2012), where some
of them are key bioengineering species (Rützler, 1975; Wulff and Buss,
1979; Buhl-Mortensen et al., 2010) thanks to their three-dimensional
structures and long lifespans (McMurray et al., 2010). Sponges can
modify environmental properties and provide suitable habitats for a
wide variety of other marine species (Van Soest, 2007; Bell, 2008), from
bacteria to commercial fishes (Buhl-Mortensen et al., 2010; Hogg et al.,
2010; Van Soest et al., 2012). Sponge assemblages dominate several,
distinctive, hard-bottom environments (Lissner et al., 1986), being one
of the most important structural organisms at several locations around
the world (e.g. Indo-Pacific, Bell and Smith, 2004; Atlantic, Hogg et al.,
2010; Antarctica, Göcke and Janussen, 2013; Mediterranean Sea,
Bertolino et al., 2015; Caribbean Sea, Maldonado et al., 2016).
their species composition across different environments, with diversity
values that tend to increase with depth in littoral habitats (Roberts and
Davis, 1996; Bell and Smith, 2004). Nonetheless, this pattern has barely
been studied in areas beyond SCUBA depth range (> 60 m) (Vacelet,
1969; Bourny-Esnault, 1971; Vacelet et al., 1994), and sponge assem-
blages of the continental shelf and slope have been explored with less
intensity. Although the Mediterranean sponge fauna is among the
world's most studied (Van Soest et al., 2012), information is minimal for
sponges from Mediterranean deep-sea areas compared to shallow wa-
ters (Bo et al., 2012), with most of the data deriving from indirect
sampling methods (Vacelet, 1969; Voultsiadou, 2005) or from by-catch
specimens collected by bottom trawlers.
Recent technological developments and increased availability of
towed video equipment, remotely operated vehicles (ROVs), manned
submersibles and autonomous underwater vehicles (AUVs) have

⁎
Corresponding author.
E-mail addresses: santin@icm.csic.es, andreusantin@gmail.com (A. Santín).

https://doi.org/10.1016/j.dsr.2017.11.003
Received 8 April 2017; Received in revised form 9 November 2017; Accepted 16 November 2017
Available online 06 December 2017
0967-0637/ © 2017 Elsevier Ltd. All rights reserved.
A. Santín et al.
significantly increased our access to deep environments (Longo et al.,
2005; Bo et al., 2015; Maldonado et al., 2016; Pierdomenico, 2016;
Pineda et al., 2016). Most studies based on these technologies, however,
have been conducted below 150 m depth, leaving intermediate depths
relatively unexplored. Moreover, most studies have focused on an-
thozoan species (Longo et al., 2005; Danovaro et al., 2010; Buhl-
Mortensen and Buhl-Mortensen, 2014), and only a few have focused on
sponge assemblages (Bo et al., 2012; Bertolino et al., 2013, 2015).
Novel work has revealed the presence of highly diverse sponge as-
semblages on the continental shelf of the Mediterranean Sea (Pedel and
Fabri, 2011; Bertolino et al., 2015). Nevertheless, these areas have been
intensely impacted by fishing practices, especially bottom trawling and
long line fishing, which have decimated their sponge assemblages
(Pansini and Musso, 1991; Pronzato, 2003). Consequently, relatively
well preserved benthic communities are mostly confined to locations
not frequented by bottom trawlers (Bo et al., 2015), generally thanks to
outcropping rocks that would damage trawl nets (Díaz et al., 2015). As
for the continental shelf and slope of the Menorca Channel, trawling
pressure during the past decades has mainly been restricted to a few
locations, leaving large areas relatively unimpacted (Grinyó, 2016).
This area will soon be declared a Marine Protected Area under the
Natura 2000 network; thus, a detailed characterization of the species
and assemblages currently present on its bottoms is required to provide
tools for management and as baselines for monitoring changes in its
diversity in coming years.
The current study aims to: (i) determine the best size of sampling
unit to study sponge assemblages on the Balearic continental shelf and
upper slope, (ii) characterize the sponge assemblages in deep habitats of
the continental shelf and upper slope in the Menorca Channel and (iii)
evaluate the variation of sponge diversity with depth.
Fig. 1. Location of the video transects in the study area. (a) Enlargement showing video tracks 1–10 in Cap Formentor. (b) Enlargement showing video tracks 75–82 in the Menorca
Canyon's head. The location of the survey area in the Mediterranean Sea is shown in the upper right corner. Projected view (UTM Zone 31N (WGS84) with geographic (WGS84)
coordinates indicated for reference.
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Deep-Sea Research Part I 131 (2018) 75–86
2. Material and methods
2.1. Study area
The Menorca Channel is located between Mallorca and Menorca
Islands (39°53'0.73"N, 3°29'51.16"E) (Fig. 1) in the Balearic Archipelago
(western Mediterranean Sea), and is part of the Balearic Promontory
(Acosta et al., 2002). The Channel's continental shelf extends between
40 and ~110 m depth, and mainly consists of maërl substrates of
varying densities, alternating with patches of coastal detrital sediments
and coralligenous outcrops (Barberá et al., 2012). Deeper, distinctively
smooth relief and large extents of detrital sediments and fine sand with
a few isolated patches of outcropping rock characterize most of the
continental shelf break and continental slope at 150–350 m depth
(Grinyó, 2016). Vertical walls and rock outcrops are the dominant
substrate near Cap Formentor (Fig. 1a) and in the Menorca Canyon's
head (Fig. 1b). Hydrologically, the Menorca Channel is located in a
border zone between the Balearic and the Algerian sub-basins. The
northern part of the study area is mainly influenced by a branch of the
Liguro-Provençal-Catalan current (also termed the Northern Current),
known as the Balearic Current (Balbín et al., 2012) and its associated
front (Ruiz et al., 2009). The Balearic Current flows over the northern
slope of the Balearic archipelago at ~200 m depth (López García et al.,
1994; Ruiz et al., 2009). The southern slope is not influenced by a
steady current (Amores and Monserrat, 2014), but mesoscale gyres
arrive intermittently after detaching from the Algerian Current and the
Almería-Oran front (Millot, 1987).
A. Santín et al.
2.2. Video recording
Eighty-five video transects, covering a total distance of 75 km, were
recorded between 50 and 350 m depth during six different surveys in the
frame of the LIFE+ INDEMARES and ENPI-ECOSAFIMED projects on board
the R/V “Garcia del Cid” (September 2010, April 2011, October 2011, June
2012), the R/V “Miguel Oliver” (August 2011) and the R/V "SOCIB"
(August 2014) (Fig. 1; Supplementary material 1). The first 20 video
transects during the first two surveys were recorded with the manned
submersible JAGO (IFM-GEOMAR), whereas the remaining 65 video
transects were recorded during the last four surveys using the ROV Nemo,
operated by Gavin Newman. Both vehicles were equipped with depth
sensors, a grabber and an HD camera (1920×1080) that recorded images in
a digital format. Two parallel laser beams provided scales on the images,
which were required for the spatial-density analyses. Both JAGO and Nemo
moved at a constant speed of ~0.3 knots, with transect lengths ranging
between 80 and 2375 m. Transects were haphazardly set along the study
area in order to cover as much of its extension as possible, with areas
suspected to be dominated by rocky substrates explored more intensively
(Fig. 1; Supplementary material 1).
In order to confirm the taxonomic identifications of the species
observed in the video footage, individuals were collected along several
transects and preserved for morphologic analyses using 10% formalin
in seawater. Additionally, a small tissue sample from each individual
was preserved in absolute ethanol for genetic determination.
2.3. Video analysis
Quantitative video analysis followed the methodology described in Gori
et al. (2011), Ambroso at al. (2013) and Grinyó et al. (2016), using the
software Final Cut Pro 7 (Apple Inc.). Pauses and loops were removed from
the footage to avoid overestimation of transect length. Video sequences
recorded too far above the seafloor or with poor image quality were dis-
carded from the analysis. The remaining video footage was considered
suitable for analysis (93.4%, linear distance of 70 km). Every sponge ob-
served within the laser beams (marking a width of 0.3 m) as the vehicle
cruised over the seabed was annotated and assigned a time reference, this
being the time elapsed since the beginning of the video transect. A similar
procedure was used to characterize substrate type, depth and slope along
each transect. Seabed substrate types were classified in four categories
based on the Wentworth scale (Wentworth, 1922): sands, cobbles and
pebbles, maërl, and outcropping rock. Seabed slope was classified as hor-
izontal (0–30°), sloping (30–80°) or vertical (80–90°). Slopes were estimated
from the video and the depth sensor. Depth was recorded as the time re-
ference of any 0.1 m depth change. Time references were transformed into
distances (d) from the beginning of the video transect according to the
vehicles speed (d = t· v, where t is the time reference expressed in seconds,
and v is the velocity expressed in meters per second).
2.4. Species identification
Species identifications were carried out on collected individuals and
HD images from the video transects using previous taxonomic studies
on Atlantic-Mediterranean and Balearic sponge faunas (e.g., Uriz, 1986;
Bibiloni, 1990; Hooper and Van Soest, 2002). Whenever possible, close-
up images of relevant parts of the sponges (e.g. inhalant/exhalant
structures, surface patterns) were recorded to help in species identifi-
cations. Sponges were identified to the lowest possible taxonomic level.
2.5. Data analysis
2.5.1. Determination of sampling units
In order to determine the minimal areas necessary to adequately
characterize communities, we calculated a species-area curve as sug-
gested by Weinberg (1978). A species-area curve is drawn by assessing
the number of species present in a first quadrat of a given area, then
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Deep-Sea Research Part I 131 (2018) 75–86
doubling that surface and recounting the species present, and then
enlarging the sample successively. Initially, the species area-curves tend
to rise sharply until reaching a break point, after which they approach
to a stable asymptote. The area at that point is assumed to be the
minimal area characterizing the sampled community. In order to obtain
an objective estimate of the asymptotic value, several approaches have
been given (Cain and Castro, 1959; Weinberg, 1978). We selected
Weinberg's (1978) stabilization criterion, that is, the minimal commu-
nity-defining area is reached when doubling the sample area results in
an increase of less than 10% in the number of species counted. This
criterion was favored as it fits better than others for marine invertebrate
communities (Weinberg, 1978).
This species-area analysis is of key importance for the later statis-
tical analyses and hence is detailed here. Raw analyzed video data were
transformed into a matrix of 1 m2 samples in which all sponge in-
dividuals were considered. Species-area curves were generated with R
software (R Development Core Team, 2014) by means of the function
specaccum in the vegan package (Oksanen et al., 2016).
The minimal area for the study of sponge assemblages in the
Menorca Channel was selected as 20 m2 (Fig. 2).
2.5.2. Sponge occupancy and assemblage composition
For statistical analysis, transects were divided into strings of ad-
jacent 20 m2 sampling units (0.3 m width and 66.66 m long), adding to
a total of 926 sampling units. For all sampling units, the numbers of
individuals of each identified species were tabulated. Sponge species
were then evaluated according to occurrence (frequency of appearance
of each species in the set of sampling units), abundance (number of
individuals per sampling unit) and density (ind./m2).
To explore the possible assemblages present, the sponge species
abundance data were square-root transformed and distances between
pairs of samples were calculated using the Bray-Curtis dissimilarity
index. The dissimilarity matrix was then represented as a dendrogram
using the hclust function of the R vegan package, and groups were de-
termined using a 20% similarity threshold. To determine the statistical
significance of differences among groups, an ANOSIM test was run
using the anosim function of the vegan package. Additionally, a SIMPER
analysis (Similarity Percentage; Clarke and Warwick, 1994) was per-
formed to identify the species contributing most to the dissimilarities
Fig. 2. Species-area curve for sponge assemblages in the Menorca Channel. Black line
represents the mean species richness, whereas the grey area represents the 95% con-
fidence interval. Dot-dashed lines and the dot at their intersection mark the threshold
where the minimal assemblage defining area is reached.
A. Santín et al. Deep-Sea Research Part I 131 (2018) 75–86

among assemblages. That was done using PRIMER v6 software (Clarke 3. Results
and Warwick, 2001).
3.1. Sponge diversity and assemblage composition

2.5.3. Relationships with environmental parameters
The environmental factors determining sponge assemblage compo-
sition were explored by canonical correspondence analysis (CCA), with
species abundances square-root transformed. CCA is a constrained
multivariate ordination technique for identifying possible relationships
between species composition (response variables) and their habitat
(explanatory variables) (Ter Braak, 1994; Greenacre, 2013). The en-
vironmental variables used in our CCA where depth, slope and substrate
type. Depth was coded into five fuzzy categories that reduce the in-
formation contained onto a categorical scale (Greenacre, 2013). The
fuzzy-coding approach allows for a more accurate depiction of non-
linear relationships among environmental data and biological gra-
dients, as well as improving the explained variance (Greenacre, 2013).
Since the inclusion of rare or sporadic species in multivariate analyses
might constrain or cause distortions in the final results (Marchant,
2002; Poos and Jackson, 2012), sponge species observed less than 5
times were excluded from the CCA analysis, as were sampling units
without sponges. As a result, the CCA was conducted on a reduced
dataset of 380 sampling units which contained 87 sponge taxa.
2.5.4. Variation of sponge diversity with depth
Sponge species richness or beta-diversity was compared among the
continental shelf (50–110 m depth), shelf break (110–180 m depth) and
the upper slope (180–350 m depth) by evaluating the slopes of rando-
mized species accumulation curves (Ugland et al., 2003) with the
speaccum function of the vegan package. Mean species accumulation
curves and their standard deviation from random permutations of data
(n = 100) give an estimate of the rate at which new species are en-
countered as sampling effort increases (Vetter et al., 2010; De Leo et al.,
2014), avoiding the distortions due to variations in species abundance
or sampling effort (Gotelli and Colwell, 2001).
Sponge species richness and diversity (quantified by means of the
exponent of Shannon's diversity index) were also computed at a finer
scale of 20 m depth intervals over the studied depth gradient
(50–350 m depth). The exponent of Shannon's information entropy was
preferred over other diversity indices (e.g. Shannon's entropy or the
Simpson index) because it weights all species by their frequency
without favouring either rarest or most common species (Jost, 2006).
Sampling effort was also evaluated based on the number of sampling
units in each 20 m depth interval.
Sponges occurred in 76% of the 926 sampling units. A total of
40,523 sponge individuals were recorded along all transects, from
which 32,234 individuals could be identified at least to genus level (109
sponge taxa; species list given in Supplementary material 2). Sponges
belonging to the classes Calcarea, Demospongiae, Homoscleromorpha
and Hexactinellida were identified in the video footage. Demospongiae
represented more than 98.8% of all sponge individuals and was dis-
tributed along the whole bathymetric range. Calcarea and Homo-
scleromorpha were only present on the continental shelf, whereas
Hexactinellida were limited to the upper slope.
Haliclona cf. elegans and Rhabderemia sp. were the most abundant
and frequent species, each accounting for 20% of the observed sponge
individuals, and they were present in 19% and 17.7% of the sampling
units, respectively (Table 1). Beside these two species, the most fre-
quent sponges were Axinella damicornis, Dictyonella sp. 1, Haliclona
mediterranea, Aplysina cavernicola, Poecillastra compressa, Axinella ver-
rucosa, Hexadella cf. dedritifera and Hamacantha falcula. The most
abundant sponges, beside H. cf. elegans and Rhabderemia sp., were A.
cavernicola, H. mediterranea, H. falcula, A. damicornis, Haliclona implexa,
Hamacantha sp., Haliclona sp. 1, and Dictyonella sp. 1 (Table 1). The
remaining species accounted individually for less than 2% of the overall
observed sponge abundances (Supplementary material 2).
Cluster analysis separated six assemblages at a 20% similarity
threshold, being significantly different from each other (ANOSIM,
p < 0.001). Average similarity in species composition within assem-
blages ranged between 31% and 62% (Fig. 3; Table 2).
The Thenea muricata assemblage was monospecific, with only T. muricata
individuals. The Axinellid assemblage was characterized by A. damicornis and
A. verrucosa, contributing 85.5% and 9.6% of the total individuals present,
respectively. The Haliclona mediterranea assemblage was characterized by H.
mediterranea (82.7%), Siphonochalina sp. 2 (10.1%) and Tethya aurantium
(3.3%). The “Haliclonid-Aplysina” assemblage was the most diverse, with the
greatest number of species required to reach a 95% cumulative contribution:
H. cf. elegans (52.5%), H. mediterranea (10.8%), A. cavernicola (5.8%), A.
damicornis (4.7%), Axinella cf. pseudominuta (4.2%), H. implexa (4.1%),
Haliclona sp. 1 (3.4%), Haliclona sp. 10 (2.8%), Rhabderemia sp. (1.9%),
Haliclona poecillastroides (1.8%), Haliclona fulva (1.7%) and Hamacantha sp.
(1.7%). The “Rhabderemia-Dictyonella” assemblage was characterized by
Rhabderemia sp. (51.3%), Dictyonella sp. 1 (28.0%), H. falcula (5.6%), P.
compressa (5.5%), H. cf. dedritifera (5.3%). Finally, the “Hamacantha-
Tretodictyum” assemblage consisted of H. falcula (76.6%), Tretodictyum reiswigi
(10.3%), Haliclona mucosa (6.6%) and H. cf. dedritifera (2.8%).

Table 1
Most abundant and frequent sponge species in the study area. Occupancy (frequency of occurrence in the set of sampling units), abundance (number of individuals), mean and maximum
density and bathymetric range are given for each species.

Species Sampling units N° individuals Mean density ± SD Max. density Depth-range

2 2
Number (%) Number (%) Individuals/m Individuals/m m

A. cavernicola 105 (11.33) 3882 (12) 5 ± 4.9 53 72–179

A. damicornis 131 (14.14) 1555 (4.8) 2.4 ± 2.5 20 52–100
A. verrucosa 85 (9.2) 583 (1.8) 2.2 ± 2.2 18 52–100
Dictyonella sp. 1 128 (13.82) 1075 (3.3) 2 ± 1.4 8 92–246
Haliclona sp. 1 66 (7.1) 1149 (3.7) 2.8 ± 2.9 26 75–113
H. cf. elegans 176 (19) 6481 (20.1) 5.3 ± 6.8 71 61–116
H. implexa 66 (7.1) 1414 (4.4) 5 ± 6.4 45 73–110
H. mediterranea 119 (12.85) 1799 (5.6) 2.6 ± 2.8 25 60–110
Hamacantha sp. 51 (5.5) 1204 (3.6) 2.8 ± 2.3 14 95–174
H. falcula 83 (9) 1557 (4.8) 3.4 ± 3 17 102–346
H. cf. dedritifera 85 (9.2) 447 (1.8) 2 ± 2.3 18 86–321
P. compressa 88 (9.5) 282 (0.9) 1.3 ± 0.8 10 95–246
Rhabderemia sp. 164 (17.7) 6509 (20.2) 6.2 ± 6.5 43 72–228

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A. Santín et al. Deep-Sea Research Part I 131 (2018) 75–86

Fig. 3. Dendrogram representing the sample clustering based

on a Bray-Curtis dissimilarity matrix. Red-dashed line re-
presents a 20% similarity threshold. Groups are differently
shaded on a grey-scale to ease visual group identification.
Assemblages are labeled as follows: Axinellids (Ax); Haliclona
mediterranea (H-m); “Haliclonid-Aplysina” (H-A); Thenea
muricata (T-m); “Rhabderemia-Dictyonella” (R-D);
“Hamacantha-Tretodictyum” (H-T).

Table 2 characterized by A. damicornis and A. verrucosa (the Axinellid assemblage;

SIMPER analysis. Species that constitute up to 95% of cumulative contribution of each Fig. 5; Supplementary material 3). Horizontal maërl beds of the inner
similarity group are present. The average similarity values were as follows: Thenea continental shelf were mainly characterized by the Haliclona mediterranea
muricata = 45.63%; Axinellids = 61.69%, Haliclona mediterranea = 43.41%, “Haliclonid-
Aplysina” = 51.75%; “Rhabderemia-Dictyonella” = 31.21%; “Hamacantha-Tretodictyum”
assemblage (Fig. 6; Supplementary material 4). On horizontal maërl beds,
= 51.25%. cobble grounds and rocky substrates of the outer continental shelf the
sponge fauna was characterized by Haliclona species (mainly H. cf. elegans,
Assemblage Species Contribution % Cumulative H. implexa, Haliclona sp. 1), A. cavernicola and Hamacantha sp. (the “Hali-
contribution %
clonid-Aplysina” assemblage; Fig. 7; Supplementary material 5). On sandy
Thenea muricata T. muricata 100 100 grounds of the shelf break T. muricata was the only sponge species (the
Axinellids A. damicornis 85.55 85.55 Thenea muricata assemblage; Fig. 8a). Rhabderemia sp. and Dictyonella sp. 1
A. verrucosa 9.62 95.18 dominated sloping rocky grounds of the shelf break, along with the fan-
Haliclona H. mediterranea 82.74 82.74
shaped sponge Poecillastra compressa (the “Rhabderemia-Dictyonella” assem-
mediterranea
Siphonochalina sp. 2 10.08 92.83 blage; Fig. 8b-g; Supplementary material 6). Finally, sloping and vertical
T. aurantium 3.28 96.1 rocky substrates of the upper slope were dominated by H. falcula and T.
“Haliclonid- H. cf. elegans 52.53 52.53 reiswigi (the “Hamacantha-Tretodictyum” assemblage; Fig. 8h-i; Supplemen-
Aplysina” tary material 7).
H. mediterranea 10.84 63.37
A. cavernícola 5.72 69.1
Supplementary material related to this article can be found online at
A. damicornis 4.66 73.76 http://dx.doi.org/10.1016/j.dsr.2017.11.003.
A. cf. pseudominuta 4.2 77.95
H. implexa 4.06 82.02
Haliclona sp. 1 3.41 85.43 3.3. Variation of sponge diversity with depth
Haliclona sp. 10 2.76 88.19
Rhabderemia sp. 1.88 90.07 Beta-diversity of sponges decreased with depth, being considerably
H. poecillastroides 1.76 91.83
higher on the continental shelf compared to the shelf break and slope
H. fulva 1.72 93.55
Hamacantha sp. 1.71 96.25 (Fig. 9). Both species richness and the exponent of Shannon's Index (Fig. 10)
“Rhabderemia- Rhabderemia sp. 51.26 51.26 presented peaks at 70–90 m depth (Fig. 10). We observed an acute plum-
Dictyonella” meting of species numbers and the exponent of Shannon's index at the
Dictyonella sp. 1 28.02 79.28 beginning shelf break (~110–130 m depth). Below that, a small increase in
H. falcula 5.58 84.86
P. compressa 5.53 90.39
species richness, and an evenness-diversity peak occurred at the transition
H. cf. dedritifera 5.27 95.65 between the shelf break and the upper slope (170–190 m depth). Finally,
“Hamacantha- H. falcula 76.61 76.61 that was followed by gradual decreases in these measures down to
Tretodictyum” 250–270 m depth, where both species richness and diversity were lowest.
T. reiswigi 10.30 86.91
Diversity values remained more or less stable on the remaining upper slope.
H. mucosa 6.64 93.84
H. cf. dedritifera 2.84 96.68
4. Discussion

3.2. Relationship with environmental parameters

4.1. Representative size of the sampling unit

The environmental variables used in our analysis explained ~34.4% of

The area selected to determine sponge assemblages in our study (20 m2)
the total inertia in the CCA, with the first two axes accumulating 15.86% of
considerably exceeds that of other work performed in neritic environments
the variance. Named based on their dominant contributing species (Fig. 4b),
(Weinberg, 1978; Coppari et al., 2016). In coastal areas, environmental fea-
six sponge assemblages (identical to those identified by the cluster analysis)
tures change dramatically at very small scales (Garrabou et al., 2002), so that
were differentiated in the CCA (Fig. 4a), which shows the relations of their
small sample areas can include large parts of the regional faunal diversity.
presence to the examined environmental factors (Fig. 4c). Depth strongly
Conversely, environmental conditions on continental shelves and slopes
segregates the Menorca Channel sponge assemblages into those occurring
worldwide generally remain constant over vast areas (Snelgrove and Smith,
on the continental shelf, the shelf break and the upper slope. On horizontal
2002). In these deeper environments, small-scale (e.g. random disturbances;
rocky bottoms of the inner continental shelf the sponge fauna was
Rex, 1981) and regional-scale processes (e.g. barriers and re-colonization;

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A. Santín et al. Deep-Sea Research Part I 131 (2018) 75–86

Fig. 4. Canonical correspondence analysis (CCA) ordination biplots. (a) The sampling
unit distribution (n = 380) are color-coded based on their main substrate type as follows:
blue: maërl, yellow: cobbles and pebbles; red: rocky substrates; green: medium
sands. Assemblages are represented with black ovals as follows: Axinellids (Ax);
Haliclona mediterranea (H-m); “Haliclonid-Aplysina” (H-A); Thenea muricata (T-m);
“Rhabderemia-Dictyonella” (R-D); “Hamacantha-Tretodictyum” (H-T). (b) Biplot re-
presenting most contributing species. Length of the lines shows the contribution of each
species to the CCA. The scale of greys indicates the degree of correlation; lines in black
indicate the highest correlations. (c) The biplot shows the different environmental vari-
ables. Depth is set as a continuous variable with 5 subdivisions, represented with a light
blue line and dep1, dep2 … for the subdivisions. Substrate and sloping are presented with
the following acronyms: M: maërl; R: rock; CP: cobbles and pebbles; MS: medium sands;
Hor: horizontal; Slo: slope; Ver: vertical. Acronyms used for all sponge species can be
found at the Supplementary Material 2.

Gage, 2004) that alter biological diversity are more widely spaced. Thus, in
order to obtain a representative area that encompasses both the small and
regional scale processes affecting species diversity in these deep environ-
ments, larger sampling units are required. That was confirmed by our results.

4.2. Sponge assemblages in the Menorca Channel

The community analyses revealed six sponge assemblages residing on

the continental shelf and upper slope of the Menorca Channel. They were
segregated by depth, substrate type and, to a lesser extent, slope. Two of
them were exclusively present on the inner continental shelf, segregated
primarily by substrate composition. That is the case of the Axinellid as-
semblage, which mostly occurs atop horizontal, coralligenous, rocky out-
crops. Its characteristic species (Axinella damicornis and Axinella verrucosa)
are common sciaphilous inhabitants on rocky bottoms of the entire
Mediterranean continental shelf (Bourny-Esnault, 1971; Pansini and Musso,
1991; Kefalas et al., 2003; Bertolino et al., 2013). Although Axinella spp.
have been observed to develop assemblages down to 130 m depth in other
areas of the Mediterranean Sea (Bertolino et al., 2015), the Axinellid as-
semblage in the Menorca Channel was mostly restricted to the inner, more
illuminated, continental shelf. Benthic communities (and sponge popula-
tions in particular) and their dynamics are strongly influenced by irradiance
(Uriz et al., 1992; Canals and Ballesteros, 1997; Kefalas et al., 2003). In the
Balearic Islands, the absence of fluvial systems and the scarce terrestrial
inputs result in an exceptionally transparent water column (Canals and
Ballesteros, 1997), which translates into relatively strong irradiance: >
0.05% of surface irradiance down to ~90 m depth. As a result, algal beds
reach deeper and attain greater biomasses in this area compared to other
Mediterranean sites (Ballesteros, 2006). Algal shading has been suggested to
enable sciaphilic sponges to inhabit otherwise less-suitable environments
(Woodin and Jackson, 1979; Uriz et al., 1992). Indeed, a dense and diverse
algal canopy was observed together with the Axinellid assemblage, partially
covering the rocky surface and shading the substrate below. In contrast, in
areas where the canopy was sparser or absent, less-schiaphilic sponges, such
as Crambe crambe or Cliona viridis, were observed. In this regard, some
sponges in the Mediterranean Sea are known to display marked seasonality
(Laubenfels, 1952; Uriz et al., 2017), which possibly correlates with the
diminishing of the algal canopy in winter. In those conditions, less-schia-
philic sponges that were sparse or had shrunk during summer might colo-
nize the newly available, alga-free rocky substrate (Woodin and Jackson,
1979; Becerro et al., 1994). Thus, we infer that the observed Axinellid as-
semblage might be a more complex semi-sciophilous assemblage than that
we observed, likely influenced by the seasonal shading provided by the
algal canopy.
The Haliclona mediterranea assemblage showed an almost identical
depth distribution with the Axinellid one, although the Haliclona cluster

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A. Santín et al. Deep-Sea Research Part I 131 (2018) 75–86

Fig. 5. Axinellid assemblage. (a) Close up of Axinella damicornis in association with Parazoanthus axinellae, (b) close up of Axinella verrucosa, (c) A. damicornis and A. verrucosa facies atop
of a partially sand-covered rock, (d) A. damicornis individuals observed under a dense algal canopy, with Crambe crambe growing on the exposed areas of the rocky bottom. Scale bar: 1 cm
(a, b) and 10 cm (c, d).

only occurred on horizontal maërl beds. It was mainly dominated by
Haliclona (Reniera) mediterranea, a well-known inhabitant of
Mediterranean coralligenous communities (Griessinger, 1971;
Ballesteros, 2006; Bertolino et al., 2013). Species belonging to this as-
semblage have also been found in maërl beds, but rather sparsely, on
the Tunisian shelf, at depths of 50–60 m where trawling takes place (El
Lakhrach et al., 2012). This pattern contrasts with that observed in our
study area, where H. mediterranea occupied vast areas of the inner
continental shelf. No other records can be found in the literature that
restrict this assemblage to maërl beds, despite the prevalence and wide
distribution of its dominant species along the northwestern edges of the
Mediterranean basin (Griessinger, 1971).
Around 70 m depth, we observed that the Axinellid, Haliclona
mediterranea and “Haliclonid-Aplysina” assemblages occasionally over-
lapped. It is common for assemblages to change gradually from one to
another across a variable transition zone (ecotone), where the numbers
of species often increase due to assemblage overlap (Bellan-Santini,
1985). In this transition zone, H. cf. elegans and H. mediterranea were
the dominant species, co-occurring with A. damicornis and A. verrucosa,
where the highly sciaphilous A. cavernicola (Maldonado and Uriz, 1995)
was completely absent. So, this zone appears to be the transition depth
(70 m) between the inner (Axinellid and Haliclona mediterranea
assemblages) and outer (“Haliclonid-Aplysina”) shelf assemblages, the
shift most likely being triggered by changing light penetration
(Ballesteros, 2006).
From the outer continental shelf down to the upper limit of the shelf
break (~70–110 m depth) the “Haliclonid-Aplysina” assemblage occurs
on horizontal maërl beds as well as on vertical rocks. This agrees with
previous studies in the Mediterranean Sea, where Aplysina cavernicola
and Haliclona cf. elegans have been observed inhabiting both horizontal
maërl (Griessinger, 1971; Maldonado and Uriz, 1995) and vertical
coralligenous substrates (Uriz et al., 1992; Cocito et al., 2002). As de-
scribed by Vacelet (1969) for offshore rocky-bottom (roche du large)
communities, we observed two distinctive morphologies of Haliclona
poecillastroides: (1) laminar, in close contact with the substrate, mostly
present on sloping or vertical surfaces, and (2) fan-shaped, present al-
most exclusively on horizontal surfaces (Fig. 8c).
Ballesteros (1991) and Augier and Boudouresque (1975) defined
two well-differentiated algal communities, the Lythophyllo-Halimedetum
tunae and the Rodriguezelletum-strafforellii, occurring on maërl and
coralligenous substrates. We reported the co-occurrence of the Axinellid
and Haliclona mediterranea assemblages and the Lythophyllo-Halime-
detum tunae algal community and, to a lesser extent, co-occurrence of
the “Haliclonid-Aplysina” and Rodriguezelletum-strafforellii groupings.

Fig. 6. Haliclona mediterranea assemblage. (a) Close up of Haliclona (Reniera) mediterranea, (b) H. mediterranea and Siphonochalina sp. 2 growing on maërl beds within a facies of Flabellia
spp., (c) A close up of Phorbas tenacior individual growing next to H. mediterranea. Peyssonnelia spp. and Flabellia spp. algae can be seen. Scale bar: 10 cm.

81
A. Santín et al. Deep-Sea Research Part I 131 (2018) 75–86

Fig. 7. “Haliclonid-Aplysina” assemblage. (a) Close up of Aplysina cavernicola, (b) Close up of Haliclona cf. elegans growing among laminar red algae (“mixing area”), (c) Overview of
“Haliclonid-Aplysina” at outer areas of the continental shelf, with H. cf. elegans and the fan-shaped Haliclona poecillastroides, d, e) View of the transition zone (70 m depth; upper limit of
the “Haliclonid-Aplysina”), with H. cf. elegans, H. mediterranea, Axinella damicornis and the orange-encrusting Haliclona (Halichoclona) fulva. Red laminar algae and the green-encrusting
Palmophyllum crassum dominate the algal community f) view of the A. cavernicola (yellow) H. cf. elegans (pink) growing atop of maërl beds at the lower distribution of the assemblage.
Scale bars 4 cm (a, b) and 10 cm (c-f).

We thus propose that the sponge assemblages and algal communities
are associated in the area and are probably limited by the same driving
factor: the gradient in irradiance penetration (Uriz et al., 1992;
Ballesteros, 2006).
At the shelf break (~110–180 m depth) sponge assemblages were
clearly segregated by substrate type. Sandy-muddy bottoms were home
to the Thenea muricata assemblage, with only Thenea muricata in-
dividuals present. T. muricata is a fairly common species on similar
bottoms around the Mediterranean Sea (Pansini and Musso, 1991),
ranging from ~50 m depth to the abyssal plain (Uriz and Rossell,
1990). It is known to be susceptible to bottom trawling (Pansini and
Musso, 1991) and is a common species in the trawling by-catch in the
Menorca Channel (Massutí and Reñones, 2005). Thus, trawling, com-
bined with the limited availability of suitable soft bottoms (Grinyó,
2016) could explain its narrow distribution in our study area.
The “Rhabderemia-Dictyonella” assemblage mainly occurred on

Fig. 8. Thenea muricata assemblage. (a) Close up of Thenea muricata; “Rhabderemia-Dictyonella” assemblage (b) Close up of Rhabderemia sp., (c) Rhabderemia sp. facies on a vertical wall,
(d) Facies of Dictyonella sp. 1 on an horizontal rocky surface, (e) Facies of the fan-shaped sponge Phakellia robusta growing on a vertical wall, (f) Pachastrella monilifera (white) and
Poecillastra compressa next to a facies of the gorgonian Viminella flagellum, (g) Poecillastra compressa individuals; “Hamacantha-Tretodictyum” assemblage, (h) Hamacantha (Vomerula)
falcula (grey-blue) and several Tretodictyum reiswigi individuals growing on a vertical wall, (i) Close up of T. reiswigi individuals. Scale bar 1 cm (a, b, i) and 10 cm (c-h).

82
A. Santín et al.
rocky substrates at the shelf break (110–180 m depth) and on the upper
slope (180–350 m depth). Rhabderemia sp. and Dictyonella sp. 1 domi-
nated this assemblage, following the distribution pattern described by
Vacelet (1969). Vertical and sloping rocky bottoms in this zone were
inhabited primarily by encrusting sponges (Rhabdaremia sp.), whereas
columnar sponges (Dictyonella sp. 1) were more characteristic of hor-
izontal rocky bottoms. Fan-shaped sponges (> 20 cm width), such as
Poecillastra compressa, and, to a lesser extent, Phakellia robusta, Pa-
chastrella monilifera and Phakellia hirondellei were also present in this
assemblage.
Bo et al. (2012) have highlighted the paramount roles of large, fan-
shaped sponges in providing habitat, shelter and food to a wide range of
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Deep-Sea Research Part I 131 (2018) 75–86
Fig. 9. Randomized species accumulation curves calculated
for each depth range. (a) Continental shelf (50–110 m depth);
(b) shelf break (110–180 m depth); (c) upper slope
(180–350 m depth). Coloured areas along each line represent
a 95% confidence interval for each bathymetric range.
associated, vagile fauna. However, in our results, the “Rhabderemia-
Dictyonella” assemblage was dominated by encrusting sponges (~63%),
with fan-shaped (~6%) and erect species (~28%) being considerably
less abundant. This is in contrast with the results of Bo et al. (2012) who
reported fan-shaped species dominating over encrusting ones in the
Bari Canyon (Southern Adriatic Sea). Such differences could be ex-
plained by the much higher sedimentation rates reported in the Bari
Canyon (Turchetto et al., 2007) compared to the Balearic Archipelago
(Pasqual et al., 2015; Grinyó et al., 2017), as higher sediment accu-
mulation can limit the presence of encrusting sponges that can become
smothered.
Finally, the “Hamacantha-Tretodictyum” assemblage occurred on
Fig. 10. Distribution with depth of species
richness and the exponential of Shannon's
index. Depth is divided into 20 m intervals,
with the number of species and diversity
represented as black dots. Black-to-grey
scale histograms represent the total numbers
of sampling units for each substrate type and
degree of slope (see legend) over the studied
bathymetric range.
A. Santín et al.
rocky walls on the upper slope at ~230–350 m depth. It consisted
mostly of the encrusting sponge Hamacantha (Vomerula) falcula and the
hexactinellid Tretodictyum reiswigi, both known to be frequent on the
upper slope (below 400 m depth) along the French (Pedel and Fabri,
2011; Fourt et al., 2017) and Italian (Taviani et al., 2017) Mediterra-
nean coasts. Moreover, T. reiswigi has only been known previously in
the Balearic archipelago from the Emile Baudot seamount (Maldonado
et al., 2015). Its presence in the Channel is a new addition to the known
Menorca sponge fauna and supports the view that T. reiswigi is very
likely widespread around the Mediterranean basin (Bourny-Esnault
et al., 2017).
4.3. Depth-diversity distribution
Diversity of megabenthic sponge species in the Menorca Channel
was higher than that reported for other Mediterranean littoral and
neritic areas (Bibiloni, 1990; Maldonado and Uriz, 1995; Cocito et al.,
2002; El Lakhrach et al., 2012). More than 100 large sponge species
were observed between 50 and 350 m depth, which is a high diversity
in agreement with previous records from the Balearic Sea (Bibiloni,
1990). The sponge assemblages observed in the Menorca Channel fol-
lowed the bathymetric pattern commonly described for other areas:
Demospongiae contributed the dominant species over the entire range,
from 50 to 350 m depth, whereas calcareous and homoscleromorph
sponges were restricted to the inner continental shelf (Van Soest et al.,
2012) and hexactinellid sponges appeared at around 200 m depth
(Reid, 1968).
Highest diversity values were concentrated on the outer continental
shelf (70–110 m depth), followed by an abrupt decline that continued
progressively downward. This pattern may be a response to several,
interacting environmental factors: light intensity, substrate type, ha-
bitat heterogeneity, other competing organisms and hydrodynamic
patterns. Highly illuminated shallow environments tend to be domi-
nated by algae, which may exclude the presence of sessile organisms
with slower growth rates (Maldonado and Young, 1996; Witman and
Dayton, 2001). This phenomenon, termed “depth emergence”, is the
increase of macroinvertebrate abundance and diversity with depth,
coinciding with the disappearance of the algal turf (Witman and
Dayton, 2001). Thus, the initial increase we observed in species rich-
ness and diversity (from 50 to 90 m depth) could be driven by the de-
creased light intensity, favoring the presence of a more diverse in-
vertebrate fauna (Kefalas et al., 2003). The presence of well-developed
coralligenous outcrops and maërl beds, characterized by the more di-
verse “Haliclonid-Aplysina” assemblage (> 50 species), also enhances
sponge diversity close to the shelf break. In addition, cryptic species
could potentially increase the observed diversity values, if they could be
successfully determined and identified (Bertolino et al., 2013). Ocean
currents and tides have also been suggested to favor the development of
dense aggregations of benthic suspension feeders on the shelf break, as
the area provides an abundant and stable food supply to them (Thiem
et al., 2006). Finally, the merging of continental shelf and slope faunas
on the shelf break may play a role in establishing the high shelf-break
diversity, a sort of mid-domain effect (Colwell and Lees, 2000). How-
ever, the abrupt decline in sponge diversity occurring just below the
shelf break, which is also observed in other areas (e.g., the Bahamian
Slope, Maldonado and Young, 1996), could derive from the dis-
appearance of coralligenous outcrops and maërl beds, coupled with a
marked overall decrease in the heterogeneity of substrates. At the
Menorca Channel's shelf break and upper slope, soft sediments domi-
nated over hard substrates. Sponge diversity at the shelf break and
upper slope is strongly linked to the presence of hard and vertical
substrates, where encrusting sponges dominate the community
(Maldonado and Young, 1996). On the contrary, deep soft sediments
harbor very low sponge diversity around the Mediterranean Sea (Fourt
et al., 2017). This could explain the observed diversity pattern and also
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Deep-Sea Research Part I 131 (2018) 75–86
explain the diversity peak at 170–190 m that coincided with the pre-
sence of vertical walls.
5. Conclusions
More than 40,000 sponges individuals belonging to 109 taxa were
observed to inhabit the deep areas of the Menorca Channel area. This
high number of species reflects the interactions among multiple en-
vironmental factors, which creates a variety of niches and favors de-
velopment of multiple sponge assemblages. Overall, six different as-
semblages were identified inhabiting distinctive shelf and slope
environments, with depth and substrate type being the important fac-
tors establishing these species associations. The high densities of par-
ticular species in many locations indicate that a pool of millions of
individuals inhabits the whole study area. Thus, we believe that the
Menorca Channel harbors one of the best-conserved, best-developed
sponge assemblages recorded and remaining to date in the Western
Mediterranean. Due to the imminent establishment of this area as a
marine protected area (MPA), it is of paramount importance to increase
our knowledge about the distributions, population size structures and
vulnerabilities to anthropogenic stressors of its most representative
species. We need a quantified and objective faunal baseline to support
implementation of effective management and monitoring measures.
The present study is a contribution towards meeting that goal.
Acknowledgments
The authors thank the crews of the R/V García del Cid, Miguel
Oliver and SOCIB, the JAGO team (J. Schauer and K. Hissmann) from
IFM-GEOMAR (Kiel, Germany), the NEMO ROV pilot Gavin Newman,
and Charles Miller, Michael Greenacre, David Díaz, Creu Palacín and
Carlota Ruiz for their aid. This work was funded by the European
project LIFE+ INDEMARES ‘‘Inventario y designación de la red Natura
2000 en áreas marinas del estado español” (LIFE07/NAT/E/000732)
and the EU-ENPI CBC-Med project ECOSAFIMED ENPI-ECOSAFIMED -
II-B/2.1/1073 (“Towards Ecosystem Conservation and Sustainable
Artisanal Fisheries in the Mediterranean basin”) (2014-15).
Appendix A. Supplementary material
Supplementary data associated with this article can be found in the
online version at http://dx.doi.org/10.1016/j.dsr.2017.11.003.
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 Coordinates
Main Main Date
Transect Depth (m)
Substrate slope
Latitude Longitude
129–223 MS H June - July 2012
1 39,99426838 3,101521851
117–293 R S Mar. - Apr. 2011
2 39,999445 3,120466667
117–223 R H June - July 2012
3 39,99666186 3,1233274
115–143 R H June - July 2012
4 39,9974344 3,1254479
200–224 MS H June - July 2012
5 39,99320853 3,139124453
144–207 R S Mar. - Apr. 2011
6 40,01159 3,143505
135–190 R S Oct. - Nov. 2011
7 40,01098167 3,145908333
155–207 CBP S June - July 2012
8 40,01082904 3,147817893
120–248 R S June - July 2012
9 40,016884 3,15115
103–229 MDS H Mar. - Apr. 2011
10 40,01808833 3,155811667
113–146 R S Mar. - Apr. 2011
11 40,03830167 3,166181667
110–116 CBP H June - July 2012
12 40,03035 3,172840957
103–115 MS H June - July 2012
13 40,0279648 3,1802042
130–200 MS S June - July 2012
14 40,060532 3,175133
233–341 R S Mar. - Apr. 2011
15 40,0835 3,2675
165–251 MS H June - July 2012
16 40,06435 3,293333
289–346 R V Mar. - Apr. 2011
17 40,07633167 3,310833333
174–180 MS H June - July 2012
18 40,04120598 3,35010009
177–301 MS H Mar. - Apr. 2011
19 40,0515 3,433333333
140–244 MS S Mar. - Apr. 2011
20 40,076 3,637866667
154–201 MS H Mar. - Apr. 2011
21 40,09543833 3,658056667
94–111 M H June - July 2012
22 39,98788991 3,309564234
102–104 M H July- Aug. 2014
23 39,98914776 3,307714193
July- Aug. 2014
24 39,99284405 3,309567126 97–110 M H
July- Aug. 2014
25 40,01289173 3,3356009 103–113 M H
July- Aug. 2014
26 40,01404871 3,332346633 103–108 M H
102–117 M H July- Aug. 2014
27 40,01393766 3,33284778
149–146 MS H June - July 2012
28 40,00305776 3,523648568
134–141 MS H June - July 2012
29 39,94863034 3,470327944
53 R H Aug. - Sep. 2011
30 39,99821667 3,702966667
52 R H Aug. - Sep. 2011
31 39,98046667 3,684466667
54 R H Aug. - Sep. 2011
32 39,9757 3,724783333
66 M H Aug. - Sep. 2011
33 39,96253333 3,648366667
July - Aug. 2014
34 39,93320938 3,695487424 63–66 MS H
67 M H Aug.–Sep. 2011
35 39,92841667 3,69125
July- Aug. 2014
36 39,92008841 3,640199905 73 M H
74–91 M H Mar. - Apr. 2011
37 39,91497333 3,636185
102 MS H Aug.–Sep. 2011
38 39,91643333 3,590333333
97–137 MS H June - July 2012
39 39,900746 3,366027205
60 MS H Aug. - Sep. 2011
40 39,86739574 3,342702259
77–81 M H Aug. - Sep. 2011
41 39,88445 3,443766667
80 M H Aug. - Sep. 2011
42 39,88035 3,520566667
 July- Aug. 2014
43 39,87854496 3,527235103 73–79 M H
88 M H Aug. - Sep. 2011
44 39,88016667 3,569083333
July- Aug. 2014
45 39,87855867 3,569505262 85–90 M H
57 CSP H Aug. - Sep. 2011
46 39,85833333 3,415216667
65 M H Aug. - Sep. 2011
47 39,8649 3,491733333
68 CSP H Aug. - Sep. 2011
48 39,8582 3,4839
67–72 M H Mar. - Apr. 2011
49 39,85369333 3,541128333
73 M H Aug. - Sep. 2011
50 39,85168333 3,5511
58 R H Aug. - Sep. 2011
51 39,82818333 3,491316667
60 M H Aug. - Sep. 2011
52 39,8185 3,5569
68 M H Aug. - Sep. 2011
53 39,80588333 3,583816667
55 R H Aug. - Sep. 2011
54 39,78226667 3,514783333
56 R H Aug. - Sep. 2011
55 39,7521 3,528766667
61 M H Aug. - Sep. 2011
56 39,748 3,589
76 CSP H Aug. - Sep. 2011
57 39,74411667 3,653333333
67 CSP H Aug. - Sep. 2011
58 39,59411667 3,5096
82 MS H Aug. - Sep. 2011
59 39,66 3,628733333
80 R H Aug. - Sep. 2011
60 39,66641667 3,628166667
131–290 MS H Sep. 2010
61 39,649415 3,70606
140–212 MS H Sep. 2010
62 39,67501667 3,761941667
230–239 R S Sep. 2010
63 39,68433833 3,775805
93–102 CSP H June - July 2012
64 39,7949521 3,724034128
122–137 MS H June - July 2012
65 39,81266363 3,738941789
217–232 MS H June - July 2012
66 39,81055568 3,748022401
115–187 MS H June - July 2012
67 39,82351287 3,754702939
155–268 MS H June - July 2012
68 39,81564233 3,774984831
July - Aug. 2014
69 39,85930828 3,786797717 76–86 CSP H
July - Aug. 2014
70 39,85518202 3,808042273 96–100 CSP H
July - Aug. 2014
71 39,88705665 3,823465422 73–79 M H
July - Aug. 2014
72 39,87789726 3,845174118 77–81 M H
115–151 MS H June - July 2012
73 39,86048046 3,866007765
319–347 R S Sep. 2010
74 39,81484667 3,927195
100–175 CSP H June - July 2012
75 39,86153344 4,012877317
92–132 R S Sep. 2010
76 39,87006833 4,018556667
102–211 R S Sep. 2010
77 39,87213333 4,027653333
July - Aug. 2014
78 39,8766896 4,03129652 95–112 R S
108–184 R S Sept. 2010
79 39,87266 4,029665
95–202 CSP H June - July 2012
80 39,86702127 4,037647151
171–182 CSP H June - July 2012
81 39,86295402 4,037271377
100–217 MS H June - July 2012
82 39,868649 4,021917
90–188 MS H June - July 2012
83 39,83936831 4,045295132
123–272 MS H June - July 2012
84 39,80608355 4,131433358
135–244 R S Sept. 2010
85 39,802075 4,150125

 

 

 
Nº Depth
Acronym Frequency Assemblage
Ind. range
Class CALCAREA

Order LEUCOSOLENIDA
Family SYCETTIDAE
Sycon sp. Syc_sp 25 1.6 87–104 H-A

Family CLATHRINIDAE
Clathrina clathrus (Schmidt, 1864) Cla_cl 3 <1 68–78 (M)
Ascandra contorta (Bowerbank, 1866) As_co 2 <1 74–78 (M)

Class DEMOSPONGIAE

Order AGELASIDA
Family AGELASIDAE
Agelas oroides (Schmidt, 1864) Ag_or 2 <1 110 (M)

Order AXINELLIDAE
Family AXINELLIDAE
Axinella sp.1 Ax_sp1 12 <1 99–124 (M; R)
Axinella damicornis (Esper, 1794) Ax_dam 1555 14.14 52–100 Ax*; H-A*

Axinella polypoides Schmidt, 1862 Ax_pol 69 2.3 66–112 Ax*

Axinella cf. pseudominuta Bibiloni, 1993 Ax_sp2 389 8.4 74–127 H-A*
Axinella vaceleti Pansini, 1984 Ax_vac 15 <1 68–98 (S)

Axinella verrucosa (Esper, 1794) Ax_ver 583 9.1 52–100 Ax; H-A
Phakellia hirondellei Topsent, 1890 Pha_hi 22 1.6 127–172 R-D
Phakellia robusta Bowerbank, 1866 Pha_ro 181 5.6 120–346 R-D; H-T

Family RASPAILIIDAE
Raspailia sp. Ras_sp 10 <1 79–90 (M)

Family STELLIGERIDAE
Stelligera sp. Ste_sp 1 <1 75 (M)

Order BIEMNIDA
Family RHABDEREMIIDAE
Rhabderemia sp. Rha_sp 6509 17.7 72–227 H-A*; R-D*; H-T

Order BUBARIDA
Family DICTYONELLIDAE
Acanthella acuta Schmidt, 1862 Ac_ac 9 <1 53–76 Ax

Dictyonella sp.1 Dic_sp1 1075 13.82 92–246 H-A; R-D*; H-T

Dictyonella sp.2 Dic_sp2 15 <1 68–82 (R)

Dictyonella sp.3 Dic_sp3 20 <1 78–80 (R)

Dictyonella sp.4 Dic_sp4 28 1.72 75–112 H-A

Dictyonella obtusa (Schmidt, 1862) Dic_ob 47 2.05 52–58 Ax

Order CHONDRILLIDA
 Family HALISARCIDAE
Halisarca sp. Hal_sp 4 <1 75–79 (R)

Order CHONDROSIDA
Family CHONDROSIIDAE
Chondrosia reniformis Nardo, 1847 Cho_re 74 2.26 53–81 H-A

Order CLIONAIDA
Family CLIONAIDAE
Cliona viridis (Schmidt, 1862) Cli_vi 7 <1 53–67 Ax

Family SPIRASTRELLIDAE
Diplastrella bistellata (Schmidt, 1862) Dip_bi 18 <1 183–199 R-D

Order DENDROCERATIDA
Family DARWINELLIDAE
Aplysilla sulfurea Schulze, 1878 Ap_su 3 <1 74 (R)
Dendrilla sp. Den_sp 1 <1 124 (R)

Order DYCTIOCERATIDA
Family DYSIDEIDAE
Dysidea sp. Dys_sp 3 <1 53–61 (R)
Dysidea avara (Schmidt, 1862) 21 1 53–65 Ax
Pleraplysilla spinifera (Schulze, 1879) Par_spi 7 <1 77–123 (E; R)

Family IRCINIIDAE
Ircinia spp. Irc_spp 8 <1 53–79 (R)

Family SPONGIIDAE
Hippospongia communis (Lamarck, 1814) Hip_co 12 1.18 53–82 Ax
Spongia (Spongia) lamella (Schulze, 1879) Spo_la 12 1 53–58 (R)

Family THORECTIDAE
Cacospongia sp. Cac_sp 7 <1 55 (R)

Order HAPLOSCLERIDA
Family CALLYSPONGIIDAE
Siphonochalina sp.1 Sip_sp1 85 2.15 74–106 H-A
Siphonochalina sp.2 Sip_sp2 46 2.37 60–79 H-m*
Siphonochalina sp.3 Sip_sp3 17 1.07 63–80 H-m

Family CARALLISTIDAE
Neophrissospongia nolitangere (Schmidt, 1870) Neo_no 51 2.15 120–329 R-D; H-T

Family CHALINIDAE
Haliclona sp.1 Hal_sp1 1149 7.1 75–113 H-A*
Haliclona sp.2 Hal_sp2 19 <1 64–68 Ax

Haliclona sp.3 Hal_sp3 38 <1 78–80 (M; R)

Haliclona sp.4 Hal_sp4 23 1 73–114 (M; R)

Haliclona sp.5 Hal_sp5 2 <1 61 (R)

 Haliclona sp.6 Hal_sp6 1 <1 75 (R)

Haliclona sp.7 Hal_sp7 44 <1 98–104 H-A

Haliclona sp.8 Hal_sp8 42 1 75–104 H-A

Haliclona sp.9 Hal_sp9 34 1.8 75–112 H-A

Haliclona sp.10 Hal_sp10 547 5.18 73–105 H-A*

Haliclona sp.11 Hal_sp11 24 <1 76–103 (M; R)

Haliclona sp.12 Hal_sp12 89 1.6 75–111 H-A

Haliclona (Reniera) sp.1 Hal_re1 6 <1 61 (S)

Haliclona (Reniera) sp.2 Hal_re2 19 <1 68–82 (R)
Haliclona (Reniera) mediterranea Griessinger, 1971 Hal_me 1799 12.85 53–112 Ax; H-m*; H-A*
Haliclona (Halichoclona) fistulosa (Bowerbank, 1866) Hal_fis 38 1.8 138–298 R-D; H-T
Haliclona (Halichoclona) fulva (Topsent, 1893) Hal_fu 297 5.18 53–105 Ax; H-A*

Haliclona (Halichoclona) magna (Vacelet, 1969) Hal_ma 2 <1 100 (R)

Haliclona (Soastrella) implexa (Schmidt, 1868) Hal_imp 1414 7.1 73–110 H-A*
Haliclona (Soastrella) mucosa (Griessinger, 1971) Hal_mu 66 3.13 148–326 R-D; H-T*
Haliclona cf. elegans (Bowerbank, 1866) Hal_el 6481 19 61–116 H-m; H-A*
Haliclona poecillastroides (Vacelet, 1969) Hal_po 262 6.5 74–114 H-A*

Family PETROSIIDAE
Petrosia (Petrosia) ficiformis (Poeiret, 1789) Pet_fi 9 <1 79–110 (M)
Petrosia sp. Pet_sp 25 1.3 106–297 R-D

Order MERLIIDA
Family HAMACANTHIDAE
Hamacantha sp. Ham_sp 1204 5.03 95–174 H-A*
Hamacantha (Vomerula) falcula (Bowerbank, 1874) Ham_fa 1557 9 102–346 R-D*; H-T*

Order POECILOSCLERIDA
Family CRAMBEIDAE
Crambe sp. Cra_sp 26 1.1 59–67 Ax
Crambe crambe (Schmidt, 1862) Cra_cr 34 2 56–80 Ax

Family CRELLIDAE
Crella (Crella) sp. Cre_sp 64 1.3 73–112 H-A
Crella (Crella) elegans (Schmidt, 1862) Cre_el 21 1 53–80 (M; R)

Family ESPERIOPSIDAE
Amphilectus fucorum (Esper, 1794) Amp_fu 37 1.5 53–81 H-A

Family HYMEDESMIIDAE
Hemimycale sp. Hem_sp 7 <1 79–104 (E)
Hemimycale columnella (Bowerbank, 1934) Hem_co 18 <1 53–80 (E; M; R)
Hymedesmia sp. Hym_sp 2 <1 104 (R)

Hymedesmia (Hymedesmia) versicolor (Topsent,1893) Hym_ver 34 2.05 102–174 R-D

Hymedesmia (Stylopus) coriacea (Fristedt, 1885) Hym_cor 23 <1 53–104 (E; R)
Phorbas sp.1 Pho_sp1 3 <1 75–79 (R)

Phorbas sp.2 Pho_sp2 15 1 56–73 (R)

Phorbas tenacior (Topsent, 1925) Pho_te 48 1.4 53–82 H-m

Family LATRUNCULIIDAE
 Latrunculia sp.1 Lat_sp1 1 z1 97 (M)

Latrunculia sp.2 Lat_sp2 7 <1 136–329 (R)

Latrunculia sp.3 Lat_sp3 3 <1 67–82 (S)

Order POLYMASTIIDA
Family POLYMASTIIDAE
Polymastia sp. Pol_sp 1 <1 98 (S)
Quasillina brevis (Bowerbank, 1861) Qua_br 17 <1 74–104 (M; S)

Order SUBERITIDA
Family HALICHONDRIIDAE
Axinyssa digitata (Cabioch, 1968) Axi_dig 2 <1 55–110 (S)
Ciocalypta penicillus Bowerbank, 1862 Cio_pe 2 <1 79 (S)
Spongosorites sp.1 Spo_sp1 21 1 146–223 (R)
Spongosorites sp.2 Spo_sp2 11 <1 153–237 (R)
Topsentia calabrisellae Bertolino, Bo, Canese,
Top_cal 59 3.23 96–223 R-D
Bavestrello & Pansini, 2013

Family SUBERITIDAE
Aaptos aaptos (Schmidt, 1864) Aap_aa 351 7.2 77–181 H-A
Aaptos papillata (Keller, 1880) Aap_pa 7 <1 63–82 (S)
Protosuberites rugosus (Topsent, 1893) Pro_rug 19 <1 78–84 (M)
Rhizaxinella pyrifera (Delle Chiaje, 1828) Rhi_py 4 <1 174–209 (S)
Suberites carnosus (Johnston, 1842) Sub_ca 21 1.4 158–293 H-T
Suberites domuncula (Olivi, 1792) Sub_dom 13 1.18 53–82 (E)
Suberites ficus (Johnston, 1842) Sub_fi 1 <1 55 (S)
Suberites syringella (Schmidt, 1868) Sub_sy 10 <1 98–128 (S)

Order TETHYIDA
Family TETHYIDAE
Tethya sp. Tet_sp 14 1.07 56–80 (M)
Tethya aurantium (Pallas, 1766) Tet_au 48 2.9 53–98 H-m*

Order TETRACTINELLIDA
Family GEODIIDAE
Erylus sp. Ery_sp 43 2.26 79–112 H-A

Family PACHASTRELLIDAE
Pachastrella monilifera Schmidt, 1868 Pac_mo 67 3.5 114–295 R-D

Family TETILLIDAE
Craniella sp. Cra_sp 23 <1 174–183 (R)
Craniella cranium (Müller, 1776) Cra_cr 5 <1 67–80 (R)

Family THENEIDAE
Thenea muricata (Bowerbank, 1858) Th_mu 415 2.15 100–270 Th-m*

Family VULCANELLIDAE
Poecillastra compressa (Bowerbank, 1866) Poe_co 282 9.5 95–246 H-A; R-D*; H-T

Order VERONGIIDA
 Family APLYSINIDAE
Aplysina aerophoba Nardo, 1833 Apl_ae 1 <1 53 (R)

Aplysina cavernicola (Vacelet, 1959) Apl_cav 3878 11.33 72–179 H-A*; R-D

Family IANTHELLIDAE
Hexadella cf. dedritifera Topsent, 1913 Hex_ded 447 9.17 86–321 R-D*; H-T*
Hexadella topsenti Reveillaud, Allewaert, Pérez, Vacelet,
Hex_top 27 1.18 56–80 (R)
Banaigs & Vanreusel, 2012

Class HOMOSCLEROMORPHA

Order HOMOSCLEROPHORIDA
Family OSCARELLIDAE
Oscarella sp. Osc_sp 20 <1 56–58 (E; R)

Class HEXACTINELLIDA

Order HEXACTINOSIDA
Family TRETODICTYIDAE
Tretodictyum reiswigi, Boury-Esnault, Vacelet &
Tre_tub 143 3.7 142–329 H-T*
Chevaldonné, 2017