Introduction

It is well known that sponges host a variety of macrobenthic organisms, providing them with shelter and constant food supply (Fishelson 1966, Koukouras et al. 1985, Koukouras et al. 1996, Koukouras et al. 1992, Çinar and Ergen 1998, Ilan et al. 1994). The relationship between sponges and their associated macrofaunal species has been investigated by several scientists (e.g. Pearse 1932, Pearse 1950, Arndt 1933, Bacescu 1971, Frith 1976, Long 1968).

The sponge Sarcotragus foetidus Schmidt, 1862 (Fig. 1) belongs to the class of Demospongiae and, more specifically, to the subclass of Keratosa, i.e. sponges with skeleton comprised of spongin fibers (order Dictyoceratida, family Irciniidae).

Figure 1.

Photo of the demosponge Sarcotragus foetidus Schmidt, 1862, collected in the course of the study.

This particular sponge species has an extensive network of small and large channels and cavities, and thus allows a variety of benthic invertebrates to inhabit them. Its surface is characterized by conules of 2–3 mm height, which are 10–15 mm apart from one another. The main skeleton is composed by a reticulate network of primary (ca. 100–200 µm in diameter) and secondary (ca. 50–100 µm in diameter) fibres (Manconi et al. 2013). Interestingly, this species was even mentioned by Aristotle, who had named it "Aplysias", meaning that it cannot be cleaned and used as a bath sponge, although its external morphology resembles to that of common bath sponges (Voultsiadou and Vafidis 2007).

The macrofaunal assemblages associated with S. foetidus have been investigated by several authors (e.g. Çinar and Ergen 1998, Çinar et al. 2002, Koukouras et al. 1985, Pansini and Daglio 1981, Rützler 1976) and in different study sites (e.g. Aegean Sea, Ligurian Sea, Tunisian coasts), although they have not been studied yet in the Levantine Sea.

General description

Project description

Study area description: 

Samples were collected from one location at the coast of Halkidiki (Greece) and one location at the coast of Cyprus (Fig. 2). The two study sites can be distinguished based on their trophic state index. North Aegean Sea can be characterized as mesotrophic to eutrophic (Kyriakidis et al. 2015), in contrast to the oligotrophic Levantine Basin (Duineveld et al. 2000). Both study sites are rocky shores dominated by different species of photohilic algae, thus no obvious differences in the sponge associated fauna can be attributed to differences in the substrate type.

Figure 2.

Geographical location of the study area indicating the sampling stations.

Linaraki: The sampling site is located on the peninsula of Sithonia, in the North Aegean Sea (Fig. 3a). It is a moderately exposed rocky shore with a photophilic algal assemblage dominated by Ellisolandia elongata.

a

b

Figure 3.

Location of the sampling station in:

a: Greece.
b: Cyprus.

Milouria: The sampling site is located close to the town of Pafos (Southwest Cyprus) (Fig. 3b. It is an exposed rocky shore with a photophilic algal assemblage dominated by the the non-indigenous species Palisada perforata.

Sampling methods

Geographic coverage

Description: 

Includes one location in Cyprus (Milouria, Kissonerga, Pafos) and one location in Greece (Linaraki, Sithonia, Halkidiki) (Christodoulou et al. 2013). More information can be found in Table 1.

Table 1. Download as CSV 

Locality, geographical coordinates, depth (m) and physical characteristics of the sampling stations.

Locality	Coordinates	Depth (m)	Habitat
Linaraki Beach, Sykia, Chalkikidi, Greece	40° 2′ 15.003" N 23° 59′ 57.264" E	14 - 17	Rocky shore, moderately exposed. Photophilic algal assemblage dominated by Ellisolandia elongata (J.Ellis & Solander) K.R.Hind & G.W.Saunders, 2013
Synthiana’s bay, Milouria, Kissonerga, Paphos, Cyprus	34°′ 18.8148" N 32° 23′ 18.8808" E	5 - 10	Rocky shore, exposed. Photophilic algal assemblage dominated by Palisada perforata (Bory de Saint-Vincent) K.W. Nam, 2007

Taxonomic coverage

Description: 

The dataset comprises distribution information for 90 taxa, belonging in 48 families and 8 phyla. Detailed information is presented in Table 2. Of these, 8 species have been recorded for the first time in the Levantine Basin.

Table 2. Download as CSV 

Taxa identified to the lowest taxonomic level possible and included in the dataset. *: non-indigenous species (ÖZTOPRAK et al. 2014, Pancucci-Papadopoulou et al. 2005). **: identification questionable for individuals from Cyprus (may be in fact Phascolosoma (Phascolosoma) stephensoni) (Açik 2014).

Phylum	Class	Scientific Name	New record for the Levantine Basin
Mollusca	Polyplacophora	Acanthochitona fascicularis
Arthropoda	Malacostraca	Alpheus dentipes
Annelida	Polychaeta	Amphitrite rubra
Annelida	Polychaeta	Amphitrite variabilis
Annelida	Polychaeta	Arabella iricolor
Sipuncula	Phascolosomatidea	Aspidosiphon (Aspidosiphon) muelleri muelleri
Arthropoda	Malacostraca	Athanas nitescens
Mollusca	Gastropoda	Bittium reticulatum
Annelida	Polychaeta	Branchiomma bombyx
Annelida	Polychaeta	Branchiosyllis exilis
Porifera	Calcarea	Calcarea
Mollusca	Polyplacophora	Callochiton septemvalvis
Annelida	Polychaeta	Capitella capitata
Arthropoda	Malacostraca	Ceradocus (Ceradocus) orchestiipes	+
Annelida	Polychaeta	Ceratonereis (Composetia) costae
Annelida	Polychaeta	Ceratonereis (Composetia) hircinicola
Arthropoda	Malacostraca	Cestopagurus timidus
Arthropoda	Malacostraca	Colomastix pusilla
Mollusca	Gastropoda	Columbella rustica
Arthropoda	Malacostraca	Cymodoce spinosa	+
Arthropoda	Malacostraca	Dexamine spinosa
Annelida	Polychaeta	Dipolydora armata	+
Annelida	Polychaeta	Dorvillea rubrovittata
Arthropoda	Malacostraca	Elasmopus pocillimanus
Annelida	Polychaeta	Eunice vittata
Arthropoda	Malacostraca	Eurydice affinis	+
Arthropoda	Malacostraca	Galathea intermedia
Arthropoda	Malacostraca	Gammaropsis crenulata
Annelida	Polychaeta	Glycera tesselata
Chordata	Actinopteri	Gobius geniporus
Annelida	Polychaeta	Harmothoe spinifera
Mollusca	Bivalvia	Hiatella arctica
Arthropoda	Malacostraca	Hippolyte leptocerus
Annelida	Polychaeta	Hydroides niger
Annelida	Polychaeta	Hydroides pseudouncinatus
Arthropoda	Malacostraca	Janira maculosa	+
Annelida	Polychaeta	Lepidasthenia elegans
Arthropoda	Malacostraca	Leucothoe spinicarpa
Arthropoda	Malacostraca	Liljeborgia dellavallei
Mollusca	Bivalvia	Lima lima
Mollusca	Bivalvia	Lithophaga lithophaga
Annelida	Polychaeta	Lumbrineris coccinea
Annelida	Polychaeta	Lumbrineris latreilli
Annelida	Polychaeta	Lysidice collaris
Annelida	Polychaeta	Lysidice ninetta
Arthropoda	Malacostraca	Lysmata seticaudata
Annelida	Polychaeta	Marphysa sanguinea
Arthropoda	Malacostraca	Microdeutopus bifidus
Mollusca	Gastropoda	Mitra cornicula
Nemertea	Nemertea	Nemertea spp.
Annelida	Polychaeta	Nereis pelagica
Mollusca	Gastropoda	Ocinebrina aciculata
Echinodermata	Ophiuroidea	Ophiactis savignyi*
Arthropoda	Malacostraca	Pagurus anachoretus
Annelida	Polychaeta	Palola siciliensis
Arthropoda	Malacostraca	Paractaea monodi
Arthropoda	Malacostraca	Paradoxapseudes intermedius	+
Annelida	Polychaeta	Perinereis cultrifera
Sipuncula	Phascolosomatidea	Phascolosoma (Phascolosoma) granulatum**
Annelida	Polychaeta	Pholoe minuta	+
Arthropoda	Malacostraca	Pilumnus spinifer
Annelida	Polychaeta	Platynereis dumerilii
Annelida	Polychaeta	Pontogenia chrysocoma
Annelida	Polychaeta	Psamathe fusca
Annelida	Polychaeta	Pseudopotamilla reniformis
Arthropoda	Malacostraca	Quadrimaera inaequipes
Annelida	Polychaeta	Serpula concharum
Annelida	Polychaeta	Serpula vermicularis
Annelida	Polychaeta	Spirobranchus polytrema
Annelida	Polychaeta	Spirobranchus triqueter
Mollusca	Bivalvia	Striarca lactea
Annelida	Polychaeta	Subadyte pellucida
Annelida	Polychaeta	Syllis armillaris
Annelida	Polychaeta	Syllis columbretensis
Annelida	Polychaeta	Syllis gerlachi
Annelida	Polychaeta	Syllis gracilis
Annelida	Polychaeta	Syllis hyalina
Annelida	Polychaeta	Syllis krohni
Annelida	Polychaeta	Syllis variegata
Arthropoda	Malacostraca	Synalpheus gambarelloides
Arthropoda	Malacostraca	Tanais dulongii	+
Annelida	Polychaeta	Trypanosyllis zebra
Annelida	Polychaeta	Vermiliopsis infundibulum
Annelida	Polychaeta	Vermiliopsis monodiscus
Annelida	Polychaeta	Vermiliopsis striaticeps

The contribution of the different phyla found in the samples is shown in the figures below (Figs 4, 5, 6, 7, 8, 9).

Figure 4.

Percentages of main phyla in the summer samples of Cyprus, as calculated based on the individual count of species per total sponge volume.

Figure 5.

Percentages of main phyla in the winter samples of Cyprus, as calculated based on the individual count of species per total sponge volume.

Figure 6.

Percentages of main phyla in all the samples of Cyprus, as calculated based on the individual count of species per total sponge volume.

Figure 7.

Percentages of main phyla in the summer samples of Greece, as calculated based on the individual count of species per total sponge volume.

Figure 8.

Percentages of main phyla in the winter samples of Greece, as calculated based on the individual count of species per total sponge volume.

Figure 9.

Percentages of main phyla in all the samples of Greece, as calculated based on the individual count of species per total sponge volume.

In Cyprus samples, Arthropoda was the phylum with the higher representation, both in summer (Fig. 4) and winter samples (Fig. 5). This was evident also when all the samples were included in the analysis, independent of season (Fig. 6). Interestingly, the increased abundance of Echinodermata in the summer (Fig. 4) was substantially reduced in the sponges collected during winter (Fig. 5).

However, the macrofaunal assemblage associated with the sponges collected from Greece was different; Arthropoda were still highly abundant, especially in the summer samples (Fig. 7), but other taxa such as Annelida and Mollusca showed also very high abundances and dominated the winter assemblages (Fig. 8). Overall, independently of season, the importance of the latter two phyla differentiated the sponges collected from the two locations (Fig. 9).

The aforementioned differences in the sponge associated macrofauna are also depicted on the MDS plot (Fig. 10), where it is apparent that the sponge samples cluster per country and season (ANOSIM: R = 0.811, p < 0.01).

Figure 10.

Multidimensional scaling of the sponge samples, based on the individual count of the associated species per sponge volume. Data labels according to the location of the sampling station and the sampling season.

Only 9 macrofaunal species out of the 83 were found to exist in both winter and summer sponge samples from Cyprus. On the contrary, sponges from Greece had 26 species in common when the two seasons were compared. Overall, when the species lists from the sponges collected from Greece and Cyprus were compared, there were 21 species common for both locations (Fig. 11; Table 3).

Table 3. Download as CSV 

Species found in common between the samples. *: non-indigenous species (ÖZTOPRAK et al. 2014, Pancucci-Papadopoulou et al. 2005). **: identification questionable for individuals from Cyprus (may be in fact Phascolosoma (Phascolosoma) stephensoni) (Açik 2014).

Cyprus	Greece	Cyprus - Greece
Summer - Winter	Summer - Winter
Cestopagurus timidus	Alpheus dentipes	Alpheus dentipes
Cymodoce spinosa	Amphitrite variabilis	Aspidosiphon (Aspidosiphon) muelleri muelleri
Elasmopus pocillimanus	Athanas nitescens	Bittium reticulatum
Leucothoe spinicarpa	Bittium reticulatum	Cestopagurus timidus
Microdeutopus bifidus	Ceratonereis (Composetia) costae	Colomastix pusilla
Ophiactis savignyi*	Ceratonereis (Composetia) hircinicola	Dipolydora armata
Phascolosoma (Phascolosoma) granulatum**	Cestopagurus timidus	Janira maculosa
Quadrimaera inaequipes	Colomastix pusilla	Leucothoe spinicarpa
Synalpheus gambarelloides	Harmothoe spinifera	Liljeborgia dellavallei
	Hiatella arctica	Lysidice collaris
	Hippolyte leptocerus	Nemertea spp.
	Janira maculosa	Nereis pelagica
	Lepidasthenia elegans	Palola siciliensis
	Leucothoe spinicarpa	Paradoxapseudes intermedius
	Liljeborgia dellavallei	Phascolosoma (Phascolosoma) granulatum**
	Lumbrineris coccinea	Pseudopotamilla reniformis
	Lumbrineris latreilli	Quadrimaera inaequipes
	Lysidice collaris	Serpula vermicularis
	Nemertea spp.	Striarca lactea
	Palola siciliensis	Synalpheus gambarelloides
	Phascolosoma (Phascolosoma) granulatum	Vermiliopsis striaticeps
	Spirobranchus triqueter
	Subadyte pellucida
	Syllis gracilis
	Synalpheus gambarelloides
	Vermiliopsis monodiscus

Figure 11.

Total number of species (in blue) and common number of species (in red) for both locations and both sampling seasons.

Traits coverage

Temporal coverage

Collection data

Usage rights

Data resources

Additional information

Acknowledgements

Author contributions

References

Açik Ş (2014)
Checklist of Sipuncula from the coasts of Turkey
.
Turkish Journal of Zoology
38
:
723
‑
733
. https://doi.org/10.3906/zoo-1405-74
Arndt W (1933)
Die Biologischen Beziehungen zwischen Schwämmen und Krebsen
.
Mitteilungen aus dem Zoologischen Museum in Berlin
19
:
221
‑
305
.
Bacescu M (1971)
Les spongiaires; un des plus interessants biotopes benthiques marins
.
Rapport Commission international Mer Méditerranée
20
(
3
):
239
‑
241
.
Christodoulou M, Paraskevopoulou S, Syranidou E, Koukouras A (2013)
The amphipod (Crustacea: Peracarida) fauna of the Aegean Sea, and comparison with those of the neighbouring seas
.
Journal of the Marine Biological Association of the United Kingdom
93
(
5
):
1303
‑
1327
. https://doi.org/10.1017/s002531541200183x
Çinar ME, Ergen Z (1998)
Polychaetes associated with the sponge Sarcotragus muscarum Schmidt, 1864 from the Turkish Aegean coast
.
Ophelia
48
(
3
):
167
‑
183
. https://doi.org/10.1080/00785236.1998.10426964
Çinar ME, Katağan T, Ergen Z, Sezgin M (2002)
Zoobenthos-inhabiting Sarcotragus muscarum (Porifera: Demospongiae) from the Aegean Sea
.
Hydrobiologia
482
(
1
):
107
‑
117
. https://doi.org/10.1023/A:1021260314414
Duineveld GC, Tselepides A, Witbaard R, Bak RP, Berghuis EM, Nieuwland G, van der Weele J, Kok A (2000)
Benthic–pelagic coupling in the oligotrophic Cretan Sea
.
Progress in Oceanography
46
:
457
‑
481
. https://doi.org/10.1016/s0079-6611(00)00029-x
Fishelson L (1966)
Israel South Red Sea expedition, 1962, reports no. 20: Spirastrella inconstans Dendy (Porifera) as an ecological niche in the littoral zone of the Dahlak archipelago (Eritrea)
.
Bulletin of the Sea Fisheries Research Station, Israel
41
:
17
‑
25
.
Frith D (1976)
Animals associated with sponges at North Hayling
.
Zoological Journal of the Linnean Society
58
(
4
):
353
‑
362
. https://doi.org/10.1111/j.1096-3642.1976.tb01005.x
Ilan M, Ben-Eliahu MN, Galil B (1994)
Three deep water sponges from the Eastern Mediterranean and their associated fauna
.
Ophelia
39
(
1
):
45
‑
54
. https://doi.org/10.1080/00785326.1994.10429901
Koukouras A, Voultsiadou-Koukoura E, Chintiroglou H, Dounas C (1985)
Benthic bionomy of the North Aegean Sea III. A comparison of the macrobenthic animal assemblages associated with seven sponge species
.
Cahiers de Biologie Marine
26
:
301
‑
319
.
Koukouras A, Russo A, Voultsiadou-Koukoura E, Arvanitidis C, Stefanidou D (1996)
Macrofauna associated with sponge species of different morphology
.
Marine Ecology
17
(
4
):
569
‑
582
. https://doi.org/10.1111/j.1439-0485.1996.tb00418.x
Koukouras A, Russo A, Voultsiadou-Koukoura E, Dounas C, Chintiroglou C (1992)
Relationship of sponge macrofauna with the morphology of their hosts in the North Aegean Sea
.
Internationale Revue der gesamten Hydrobiologie und Hydrographie
77
(
4
):
609
‑
619
. https://doi.org/10.1002/iroh.19920770406
Kyriakidis P, Vasios G, Kitsiou D (2015)
Proc. SPIE 9535
.
Third International Conference on Remote Sensing and Geoinformation of the Environment (RSCy2015), 95351W
.
7
pp. URL: https://doi.org/10.1117/12.2192565 https://doi.org/10.1117/12.2192565
Long ER (1968)
The associates of four species of marine sponges of Oregon and Washington
.
Pacific Science
22
:
347
‑
351
.
Manconi R, Cadeddu B, Ledda F, Pronzato R (2013)
An overview of the Mediterranean cave-dwelling horny sponges (Porifera, Demospongiae)
.
ZooKeys
281
:
1
‑
68
. https://doi.org/10.3897/zookeys.281.4171
ÖZTOPRAK B, DOĞAN A, DAĞLI E (2014)
Checklist of Echinodermata from the coasts of Turkey
.
TURKISH JOURNAL OF ZOOLOGY
38
:
892
‑
900
. https://doi.org/10.3906/zoo-1405-82
Pancucci-Papadopoulou MA, Zenetos A, Corsini-Foka M, Politou CY (2005)
Update Of Marine Alien Species In Hellenic Waters
.
Mediterranean Marine Science
6
(
2
):
147
‑
158
. https://doi.org/10.12681/mms.188
Pansini M, Daglio S (1981)
Osservazioni sull’ inquilinismo di policheti erranti in alcune demospongie del litorale Ligure
.
Bollettino dei Musei e degli Istituti Biologici dell'Università di Genova
48-49
:
55
‑
60
.
Pearse AS (1932)
Inhabitants of certain sponges at Dry Tortugas
.
Papers from the Tortugas Laboratory of the Carnegie Institution of Washington
28
:
117
‑
124
.
Pearse AS (1950)
Notes on the inhabitants of certain sponges at Bimini
.
Ecology
31
(
1
):
149
‑
159
. https://doi.org/10.2307/1931369
Rützler K (1976)
Ecology of Tunisian commercial sponges.
Tethys
7
:
249
‑
264
. URL: https://repository.si.edu/bitstream/handle/10088/7850/iz_Ruetzler_1976.pdf?sequence=1&isAllowed=y
Voultsiadou E, Vafidis D (2007)
Marine invertebrate diversity in Aristotle’s zoology
.
Contributions to Zoology
76
(
2
):
103
‑
120
. URL: http://www.repository.naturalis.nl/document/49637

Supplementary material