Zootaxa 2812: 41–62 (2011)
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Copyright © 2011 · Magnolia Press
ISSN 1175-5326 (print edition)
Article
ZOOTAXA
ISSN 1175-5334 (online edition)
Revised description of a poorly known Mediterranean Dictyoceratid
bath sponge, Spongia (Spongia) zimocca (Schmidt, 1862)
(Porifera: Demospongiae: Dictyoceratida)
JEANNE CASTRITSI-CATHARIOS1,4, ROB W. M. VAN SOEST2,
EFTHIMIOS KEFALAS1 & JEAN VACELET3
1
Department of Zoology-Marine Biology, Faculty of Biology, National and Kapodistrian University of Athens, Panepistimiopolis, Athens 157 84, Greece. E-mail: cathario@biol.uoa.gr
2
Zoological Museum of the University of Amsterdam, P.O. Box 94766, 1090 GT Amsterdam, The Netherlands.
E-mail: R.W.M.vanSoest@uva.nl
3
Centre d’Océanologie de Marseille, Aix-Marseille Université, CNRS UMR 6540 DIMAR, Station Marine d’Endoume, Rue de la Batterie des Lions, 13007 Marseille, France. E-mail: jean.vacelet@univmed.fr
4
Corresponding author. E-mail: cathario@biol.uoa.gr
Abstract
Spongia (Spongia) zimocca (Schmidt, 1862) is a real problem for taxonomists. This is due to the fact that it exhibits a wide
diversity of forms as well as similarities with other species of the genus. Nevertheless, professional sponge fishermen are
able to recognize this species easily based on their own empirical criteria. The lack of easily recognized taxonomic characters in conjunction with the ambiguity concerning the original description makes necessary an updated description of
the species. We tried to solve the uncertainty existing among the taxonomists who have been involved with the description
of this Mediterranean sponge, basing our description on the Schmidt type specimen and on material deposited in the Amsterdam Museum. In this paper we have summoned up not only the known morphological characters used for the identification of other species belonging to this genus (color, size, shape, skeleton structure, size of the fibers) but also additional
morphological characters, used for a first time, aiming to obtain a clearer picture. These new characteristics are: tensile
strength, skeletal density, morphology of the conules and the organic content of the fiber mass.
Key words: Demospongiae, Dictyoceratida, Leather sponge, identification, lectotype, polymorphism, commercial value,
biogeography
Introduction
Representatives of the Order Dictyoceratida Minchin (1900), Class Demospongiae, are lacking the mineral skeletal
element, which is substituted by spongin fibers, resulting in enhanced fine structure and flexibility. To this order
belongs the family Spongiidae Gray (1867), the skeleton of which consists exclusively of unstratified collagen
fibers, composed of spongin B, (Gross et al. 1956; Garrone et al. 1973). All species of the genus Spongia Linnaeus
(1759), which are widely known in the Mediterranean as bath sponges since ancient times (Voultsiadou 2007), still
“hold secrets” to this day regarding their taxonomy. The overall quality of a bath sponge is determined by several
factors, including tensile strength, elasticity, absorbency, water retention, quantity of foreign particles included in
the spongin fibers (Moore 1908; Castritsi-Catharios et al. 2007; Louden et al. 2007). Quantities of foreign particles
differ from one species to another but such differences can also be found within the same species in individuals of
different quality and/or from different habitat.
The commercial sponges harvested in the Mediterranean include the “Tsimoucha” (Greek) or “Chimousse”
(French), or “Leather sponge” (English), or “Zimoukha” (Arabic/Tunisian) i.e. Spongia (Spongia) zimocca which
still raises a real problem for taxonomists (Pronzato & Manconi 2008). This species is poorly studied, notwithstanding its substantial share of the total production of the sponge fishery, about 7% of the Mediterranean produc-
Accepted by E. Hajdu: 3 Mar. 2011; published: 8 Apr. 2011
41
tion (Castritsi-Catharios, 1995). It appears that the “Tsimoucha” is well known to many users and not only to those
involved in the sponge fishery (stakeholders), but it is poorly defined from a zoological point of view. This is due to
two basic reasons. The first is the polymorphism which this species exhibits, with a wide macro-morphological
variation depending mainly on the geographical harvesting area and on the depth of harvesting (Castritsi-Catharios
1995; Castritsi-Catharios et al 2011). The second is the fact that most of the descriptions have been based on commercially treated samples only, i.e. reduced to the spongin skeleton.
Aristotle describes in his zoological works “Ton Peri ta zoa istorion D, E, Z” that sponges living in deep and
calm waters are more soft (Όλως δ’ οί εν τοίς βαθέσι και ευδιεινοίς μαλακώτατοι εισίν). Concerning the geographical location, he writes that “the sponges harvested at Ellispontos are rough and dense and, in general, those harvested beyond Cape Maleas (South Peloponnese) appear to have varying softness and elasticity compared to those
harvested from the interior side”. In ancient years (according to Aristotle) this species was known as “very dense
sponge and tragos (goat)” referring to the dense structure of its fibers and to its roughness compared to other
health-care sponges. However, after the short description by Schmidt (1862) of the external morphology of S.
zimocca, with values of the diameter of the secondary fibers forming its skeleton, international bibliographic references to scientific studies are minimal and confused (Burton 1934; de Laubenfels 1948; Vacelet 1959, 1987; Rützler 1976; Hooper 1994; Castritsi-Catharios 1998, 2005; Ben Mustapha et al. 2003; Kefalas et al. 2003b; Pronzato
& Manconi 2008).
This paper is focused on the detailed description of S. zimocca (external morphology and internal structure)
accompanied by environmental data, and comparisons are made with other commercial species of sponges by using
taxonomic characters, some of them being presented for the first time. After examining a large number of S.
zimocca specimens and based on the knowledge of boat owners and sponge merchants, we can assume that mainly
the geographic area, and to a lesser extent depth, contribute significantly to the macro-morphological polymorphism which is exhibited by this species. Morphological peculiarities which have not been described on one side
and the common morphometric characters on the other (which do not allow for the separation of the species),
induced us to make a more detailed and updated description of this species including all the forms of external morphology which are frequently found.
Material and methods
Sampling methods. The main methods for sponge sampling are: lagamna (Kefalas et al. 2003a), gagava (or gangava) (Maldura 1931; Castritsi-Catharios 1998) and nargiles (Castritsi-Catharios 1998; Pronzato 1999; Voultsiadou
et al. 2008).
Most of the samples used for the investigation of the diversity of forms of S. zimocca were sampled from Greek
waters (1993–1994), from Egypt (1995) and from Libya (1993). Sponges were collected through two methods
(gagava and nargile) for Greece, while in Egypt and Libya we used exclusively Kalymnian divers. The sampling
techniques of gagava and nargile are described below.
1. Fishing gear gagava. Τhis is old traditional gear, essentially a dredge. It hauls an iron or wooden frame at the
four corners of which iron shafts or chains are adapted through rings. All four shafts are joined to an outer ring
on which a rope or metal chain is tied and which is fastened to the boat. On the other side of the metal frame a
mesh is applied, 3–4 meters in diameter and 8 meters long. The mesh has an opening of 150–170 mm. An
improved modification of the gagava consists of placing wheels (30 cm diameter) at the lower shaft of the
frame so that the gear is not drawn directly onto the substratum. Furthermore, the opening of the mesh has been
increased. The speed of the boat does not normally exceed 6oo meters per hour, i.e. 0.324 knots. Gagava is normally used on smooth substrata in both shallow and deep waters. In the case of Posidonia beds a thin chain is
used, fastened at the lower part of the metal frame so that only sponges are removed but not the Posidonia
leaves. This chain cannot be used for other types of substrata.
2. Divers (nargile method). Τhey dive to a maximum depth of 65 m. Each dive, especially at great depth, lasts
two hours approximately. The first hour of diving is dedicated to fishing sponges while the second to decompression so that diver’s disease is avoided. This timing is valid for depths greater than 40–50 m, while for 20–
25 m the whole dive may last 70–80 min (40–50 min for fishing and 30–35 for decompression).
42 · Zootaxa 2812 © 2011 Magnolia Press
CASTRITSI-CATHARIOS ET AL.
For the sampling performed in the Aegean, Cretan and Ionian Seas, divers were given a data sheet on which
they recorded their findings after each dive, assisted by a biologist on the boat. The following data were recorded:
shape (two types for S. zimocca), size (small, medium or large), presence or absence of local currents and type of
substratum on which the sponges were found.
FIGURES 1–3. Spongia zimocca. 1, Map showing in red the species’ distribution in the Mediterranean Sea (Greece: Aegean
Sea, Dodekanissos, Cyclades, Crete, Ionian Sea, Laconia / Egypt / Libya / Tunisia). 2, Compact sponge from Libya, with flat to
slightly lobose upper surface. 3, Specimen harvested from Tunisia.
Sampling area. The sampling area (Fig 1) spanned across the eastern Aegean Sea towards the south-central
Aegean, Ionian and Cretan Seas (Cape Sounion 37.6067o N, 23.9937o E) Aegina Island (37.7079o N, 23.5851o E)
(Saronikos Gulf)Crete Gaidouronissi – Chrissi, 34.8773o N, 25.7011o E), Dodekanissa (Kalymnos, in Emporio
REVISED DESCRIPTION OF SPONGIA ZIMOCCA
Zootaxa 2812 © 2011 Magnolia Press ·
43
37.0355o N, 26.9334o E, Telendos 36.9973o N, 26.9173o E), Samos (Vathi 37.7708o N, 26.9490o E and Aghia
Paraskevi 37.3981o N, 26.9907o E), Cyclades (Tilos-Antitilos narrows 36.3661o N, 27.4537o E), Naxos 36.9027o N,
25.4332o E and Ios 36.7586o N, 25.3464o E ). Libya (Tobruk 32.5002o N 24.1467o E, Darnah 33.4534o N 22.6816o
E, Benghazi, 32.0527o N 20.0713o E, Marsa Misrata 32.4040o N 15.1186o E, Sirt 31.1153o N 16.4657o E, Western of
Tarabulus 32.8552o N 13.2479o E and close to the borders with Tunisia) and Egypt (Boyat, Ras Alarm el Hum,
Agiba, Mersa Matruh) (Castritsi-Catharios 1994; Castritsi-Catharios et al. 2005). An additional specimen from
Tunisia was collected through Tunisian fishermen (Monastir).
Transmission electron microscopy. Just after collection of the sponges, untreated subsamples were immediately cut in small pieces and fixed in glutaraldehyde (25% glutaraldehyde, 1 part; 0.4M cacodylate buffer, 4 parts;
filtered seawater, 5 parts). Since the sponges were collected in the open sea they remained in this fixative for several weeks. They were then washed in seawater, postfixed in 2% osmium tetroxide in microfiltered sea water,
washed in seawater and dehydrated in graded alcohols and propylene oxide. The pieces were then embedded in a
mixture of epon/araldite.
Sections for both light microscopy and electron microscopy were cut with a Sorvall MT-2B ultramicrotome
and stained according to standard procedures utilizing toluidine blue for the light microscopy sections and uranyl
acetate, followed by lead citrate for the electron microscopy sections.
EM grids were viewed under a Phillips EM 200 electron microscope and images were recorded on 35 mm
Kodak film.
Scanning electron microscopy. The tissue was prepared in a similar way up to the dehydration stage, followed by critical point drying and/or cryofracturing to reveal internal substructures. Metal coating was performed
with gold/palladium and the samples were examined under a Cambridge stereoscan microscope.
Image pro-image analysis. Images were scanned using a Hewlett PackardTM scanjet II ex CanonTM scanner.
Acquisition of the images was made using Adobe PhotoshopTM 3.0 software on a Macintosh Quadra 700TM. The
image characteristics were examined and modulated using Image Pro IITM software on a 486 IBMTM PC. Measurements: spatial calibrating operation was used.
Biometry. The biometrical approach was performed in situ on sponges just harvested. The dimensions were
measured based on the following: as length, we took the max length of the axis parallel to the substratum, as width
the max length perpendicular to the previous axis and as height the max distance between the base (point of attachment to the substratum) and the highest point of the sponge.
Organic content of the fiber mass and skeletal density. All our samples were mechanically treated on the
boat freeing them of the tissues and of most of the foreign bodies. This mechanical treatment includes the following steps: immediately after the sponges are transferred onto the boat, the divers trample on them, wash them and
then immerse them in the sea for about two hours, again trampling on them and rewashing. After this process, the
sponges are beaten by using branches from palm trees. This way foreign structures enclosed within the sponge are
smashed and removed. In the evening the sponges are placed into a net and immersed in the sea, and in the morning
they have become “white” (in the case of S. zimocca the color turns to light grey-brown), i.e. the external membrane and a considerable quantity of the tissue have been destroyed. The whole process is then repeated until only
the fibers have been left. The sponges are then dried, losing in total 20–40% of their initial weight (water, tissues
and foreign structures are removed).
For the determination of the organic content of the fiber mass of S. zimocca we selected three pieces (samples)
from each individual sponge, as representative as possible of the species, which had been treated as above
described, i.e. they were free of foreign structures and tissue. The samples were placed in an oven, at 70 oC for 24
h, to reach a stable weight (Lovegrove 1962; Bamstedt 1974), by losing any remaining water. The dry samples
were placed in a dessicator for 30 minutes until equilibrium was achieved thus obtaining the “dry weight”. Samples
of known dry weight were placed in an oven, at 500 oC, for 4 hours, in porcelain crucibles. The final weight of the
ash is the inorganic content. By subtracting from the dry weight the weight of the inorganic material (ash) the
organic material of each sample was determined.
For the determination of the skeletal density we used samples taken from homogenous parts of the initial
sponge, free of foreign structures, and cut in the form of cuboid parallelepipeds (volume = length x width x height
of the cuboid sample). Prior to the measurement of the dimensions the samples were placed in an oven at 70 oC for
24 h. The volume was expressed in mm3 and the density in gr/cm3.
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CASTRITSI-CATHARIOS ET AL.
Tensile strength. The measurements were performed in the chemical laboratory of the Hellenic Navy Yard in
Salamina Island, Greece. The tensile strength meter ‘Zwick Material/Profung 1485, certification number 40983,
certification date 08/04/04, asset no. NS DNX 0001’ was used, connected to a computer with corresponding software.
The results were automatically printed as % maximum elongation, which is a measure of the elasticity, elongation in mm and the maximum load in kg, which is a measure of the resistance of the sample. The initial distance
between the jaws was 10 cm (the minimum possible). The upper jaw is stable while the other moves with a constant speed. All the sponges were not chemically treated to avoid any disturbance in the physical properties due to
this treatment. The sponges were washed with deionized water and were dried for 48 h at room temperature. Elongated strips were cut according to the specifications of the instrument for the ‘sample preparation’. The results are
comparable since these are expressed in kg cm-2 (specific maximum load).
Results
Taxonomy
Phylum Porifera Grant in Todd, 1836
Class Demospongiae Sollas, 1885
Order Dictyoceratida Minchin, 1900
Family Spongiidae Gray, 1867
Genus Spongia Linnaeus, 1759
Subgenus Spongia Linnaeus, 1759
Species Spongia (Spongia) zimocca Schmidt, 1862
LECTOTYPE: Schmidt’s specimen coded LMJG 15470 Cyprus, in the Graz Museum (Austria).
Material examined. Representative individuals of the material studied are deposited in the Amsterdam
museum (ZMA) except one specimen, which is kept in the Athens University, and the lectotype..
Spongia (Spongia) zimocca (between Cape Sounion and Aegina Island) ZMA POR 21430
Spongia (Spongia) zimocca (Dodekanisos) ZMA POR 21429
Spongia (Spongia) zimocca (Egypt) ZMA POR 21431
Spongia (Spongia) zimocca (Libya) ZMA POR 21428
Spongia (Spongia) zimocca (Tunisia) ZMA POR 21717
Diagnosis. Massive commercial sponge of medium size, up to 25–30 cm in width. Shape highly variable
depending on depth and fishing ground, generally irregularly lobate, with more or less developed lobes. Surface
non armored, finely conulose, with conules usually 200–300 µm high. Oscules simple and irregularly dispersed.
Color in life black to dark gray on surface, gray inside. Texture of the skeleton fibrous, rougher than in the other
Mediterranean commercial sponges. Choanosome dense, devoid of large cavities. Skeleton made of three types of
spongin fibers. Primary fibers 40–50 µm in diameter, formed by the anastomoses of secondary fibers mostly near
the surface conules, without pith, uncored or with a very few amount of foreign spicules. Secondary fibers in two
sizes, the larger 23–40 µm in diameter forming a dense reticulation, the smaller (“pseudo-tertiary fibers”) 4.7–18.4
µm in diameter forming a more or less developed reticulation.
Geographical distribution. Eastern Mediterranean Sea, possibly also in the south-west basin.
Depth. 25 to 85 m
Substratum.: Mostly on hard substrata in the coralligenous biocoenosis. The sponge occurs also on Posidonia
beds and on sandy soft bottoms.
REVISED DESCRIPTION OF SPONGIA ZIMOCCA
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45
Description of Schmidt’s lectotype. Schmidt’s sample deposited in the Graz Museum and coded LMJG
15470 Cypern has been examined. It is a massive, irregularly lobate sponge (Fig. 6 see also Pronzato & Manconi,
2008). According to our observation, the primary fibers are devoid of inclusions. They are usually difficult to follow in the inner part of the skeleton. Near the conules, they are 40–50 µm in diameter. Their diameter is more irregular than that of the secondary fibers. They do not have a pith, but they appear less homogeneous than the
secondary. The secondary fibers are 28–30 µm in maximum diameter. There are a few well-characterised thinner
secondary fibers (“pseudo-tertiary fibers”), 10–15 µm in diameter, possibly with intermediaries with the secondary.
Description of our specimens.
External morphology. S. zimocca or Leather sponge or Tsimoucha is present in two forms, A from Egypt,
Libya and Tunisia mainly, B1 and B2 from Greek seas mainly.
A (Figs. 2–6). Compact sponge, with flat to slightly lobose upper surface (directly opposite to the base), where
large oscula are observed (size range 4.2–5.4 mm). These oscula appear on the top of lobes, the latter located
closely together without any empty spaces in between. Conversely on all other surfaces, and more specifically at
the circumference of the base and up to the upper surface, filamentous projections (Fig.28) are observed. These
protrusions connect to the main body and each has a large osculum at their upper surface. This particular form is
found in Tunisia, Libya, and Egypt.
B This form includes two sub-forms. The first one B1, (Figs.7–8), which can be found in shallow depths,
exhibits a compact form with numerous protrusions connected at their base while the upper extremities are free. In
comparison to form A, form B has smaller lobes and oscula. In this form we can easily observe two categories of
oscula, i.e. small (size range 1.7–2.5 mm) and large (size range 3.5–4.5 mm). This form, especially that of ZMA
POR 21430 (Fig.7), is morphologically identical to Schmidt’s lectotype from Cyprus (Fig. 8).
The second sub-form B2 (Fig.9–10) can be found in deeper waters and is usually fished by dredge-trawlers.
The lobed formations are well developed with 1/4 or 1/5 of their length remaining attached to the sponge body
(Fig. 11). This form is in good accordance with the description of Euspongia zimocca var. adjimensis Topsent
(1924), who mentions that according to Duboscq “this sponge is often loboid with lobes ending to one or different
levels, with big gaps between them in a part or along all their length”. Their free surface is bilateral, symmetrical or
not. Oscula are not necessarily found on the summits of the lobes but more or less on the side, a little underneath
the round apex. From the osculum starts a linear channel visible through naked eye in the dry samples, as described
by Schmidt 1862 (Fig. 12).
In Table 1 all forms of S. zimocca are shown (Α, Β1 & Β2) collected by the divers during the sampling trips as
well as the corresponding geographical distribution. Of 3,783 sponges harvested from Libya, 302 were S. zimocca
in depths between 20–45 m, and of these, 241 individuals were of form A and 61 of form B1.
TABLE 1. Frequency distribution of S. zimocca forms in different areas of harvesting.
AREA
FORM A
FORM B 1
FORM B 2
Crete
-
43.8% (117)
56.2% (150)
Dodekanissos
-
36.8% (127)
63.2% (218)
Cyclades
-
52.0% (38)
48.0% (35)
N. Aegean
-
-
100.0% (37)
S. Evia
-
-
-
Ionion
-
-
-
Libya
79.8% (241)
20.2% (61)
-
The external color of S. zimocca in life depends on its habitat. Like many sponges, it is lighter coloured in
shaded habitats, and called “drossitis” by divers. In Posidonia beds, it is black, like the other commercial sponge
46 · Zootaxa 2812 © 2011 Magnolia Press
CASTRITSI-CATHARIOS ET AL.
species (Fig.13–14). As soon as the sponge is transferred to the boat and prior to any kind of treatment, a cut of the
sponge appears light gray or light brown (rusty brown), while the channels leading to the oscula are dark gray
(pers. comm., Manolis Saroukos).
The conules are thin, dense, arranged irregularly and their surface carries deep corrugations, parallel from the
base to the top of the conule (Fig 15). In samples taken from Greek waters the conules have the following mean
values (preliminary results): height = 200–300 μm, diameter of the base = 150–250 μm, diameter of the top = 50–
100 μm. (Table 2).
TABLE 2. Characteristics of the conules of the commercial species of sponges (preliminary results).
Species
Height in μm
Nr. of specimens / nr. of
replicates
Diameter of the
base in μm
Diameter of the top Surface of the conule
in μm
(External morphology)
S. zimocca 5/3
200–300
150–250
50–100
Deep corrugations, parallel, from the
base to the top of the conule
S. o. mollissima 4/3
150–200
200–250
50–100
Leaf like structure
S. o. officinalis 3/3
300–600
400–600
100–200
Thin grained structure or smooth
S. lamella 3/3
200–250
200–2500
120
Pinacoderm-like with many surfaces.
H. communis 4/3
800–1200
700–800
200–300
Pinacoderm-like, which is not differentiated up to the top. Thin, irregular
corrugations
Several extraneous structures are attached to the pinacodermal membrane. Primary fibers form conules on the
surface, which they frequently pierce (Fig. 28), surrounded by a dense mesh of secondary fibers (Figs. 16, 17).
Internal structure. Primary fibers (Figs 18, 19) are abundant near the surface, with very rare foreign inclusions, with a diameter of about 40–50 μm. The secondary fibers (Figs 20–23) are smooth and uncored. Two size
classes were found (values of their diameter are shown in Table 3). The secondary fibers with a significantly
smaller diameter, that we call here “pseudo-tertiary fibers” (Figs 24–27) exist in abundance in all our samples.
According to our knowledge this taxonomic character has not been recorded for this Mediterranean species of
Spongia.
ΤABLE 3. Spongia zimocca. Diameter of the secondary and pseudo-tertiary fibers on different positions. (samples from the
market and the specimen deposited in Graz Museum).
Diameter at the middle of Diameter close to the nodes Diameter at the mid- Diameter close to the nodes of
the secondary fibers
of the secondary fibers (μm) dle of the pseudo-ter- the pseudo-tertiary fibers (μm)
(μm)
tiary fibers (μm)
MIN
23.0
31.8
4.7
6.9
MAX
31.9
40.9
12.0
18.4
AVERAGE
27.4
37.1
9.0
12.7
STDEV
2.9
2.9
2.9
4.3
The choanocyte chambers are spherical with a diameter of 18–22 μm (Fig 29). The choanocytes are pear
shaped with a diameter of 5.6 μm. The cells carry on their upper outer side a collar formed by microvilli and a welldeveloped flagellum (approximate length 3 μm).
Distribution and biogeography. Data gathered from sampling trips: 1223 individual sponges were collected by the hookah method and 114 by the gagava method, 1337 individual sponges were collected in total, S.
zimocca representing 0.8% of this total and was found in all fishing grounds except Εvia Island and Ionian Sea, at
depths varying between 25 and 87.5 m,. The records of abiotic parameters, the depth and the type of substratum are
given in Table 4. In Table 5 we provide a summary of the results obtained from the measurements of our samples
(biometrical data correlated to the sampling stations and the depth of collection).
REVISED DESCRIPTION OF SPONGIA ZIMOCCA
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47
TABLE 4. Spongia zimocca, Sampling area and stations, abiotic parameters, depth of collection and substratum.
SN
AREA
TESU
o
C
SASU
/00
PHBO
DOSU
DOBO
TURB
m
DEPTH
m
SUB
0
SABO
0
/00
PHSU
o
30
70
a
55
b
TEBO
C
1
A
22.5
22.0
40
40
8.0
7.9
7.2
6.0
2
B
23.5
22.0
41
39
7.9
7.9
6.2
6.6
3
C
22.0
21.3
40
39
8.0
8.1
6.8
6.3
40
80
c
4
C
21.0
21.5
40
39
8.1
8.0
7.0
6.5
39
55
d
5
C
23.5
23.0
42
41
8.1
8.1
7.2
6.9
27
25
e
6
D
22.0
22.0
40
40
7.9
7.9
6.8
6.6
27
87
f
SN: serial number, AREA: geographical area, TESU: temperature of surface, TEBO: temperature of bottom, SASU: salinity of
surface, SABO: salinity of bottom, PHSU: Ph of surface. PHBO: Ph of bottom, DOSU: dissolved oxygen of surface, DOBO:
dissolved oxygen of bottom, TURB: turbidity, DEPTH: depth at which the samples were collected, SUB: type of substratum
(see below). A: Creta island, B: Cyclades complex of islands, C: Dodekanissos complex of islands, D: Samos island, I: Gaidouronissi island, II: Naxos – Ios islands, III: Kalymnos island (Emporio), IV: Kalymnos island (Vathi), V: Tilos – Antitilos
islands, VI: Vathi (Aghia Paraskevi ). a: littoral zone: hard substratum / infralittoral and circalittoral: among calcareous algae,
coralligenous biocoenosis and thin grain sand. b: infralittoral and circalittoral zone : calcareous algae, thick grain sand and coralligenous biocoenosis. c: infralittoral zone: biocoenosis of fine, well sorted, sand – anthropogenous interference / circalittoral
zone: hard substratum, coralligenous biocoenosis. d: infralittoral and circalittoral zone among calcareous algae and coralligenous biocoenosis. e: infralittoral zone: Posidonia beds and sandy soft bottoms, precoralligenous biocoenosis / circalittoral zone:
coralligenous biocoenosis. f: infralittoral and circalittoral zone : hard bottom, calcareous algae and coralligenous biocoenosis
TABLE 5. S. zimocca. Sampling stations, collection depths and dimensions of specimens.
SAMPLING
AREA
DEPTH
m
LENGTH
cm
WIDTH
cm
HEIGHT
cm
CIRCUMFERENCE
cm
CRETE
Gaidouronissi
70
60
60
25
160
70
28
19
10
78
70
17
10
10
50
55
15
11
13
37
55
16
10
8
42
KALYMNOS
Emporios-Telendos
80
5
10
7
24
80
13
10
10
31
KALYMNOS
Vathis
80
5
10
7
24
TILOS-ANTITILOS
25
14
11
10
40
25
14
11
9
34
SAMOS
Vathis-Agia Paraskevi
82
20
16
11
47
MIN
25
5
10
7
24
NAXOS-IOS
MAX
82
60
60
25
160
AVERAGE
62.9
18.8
16.2
10.9
51.5
STDEV
20.9
15.1
14.8
5
38.9
Data obtained through the fleet (diver-key forms): The divers collected 21570 individual bath sponges during the period mid-July to mid-October of 1993 and 1994 (15 boats and 45 divers). The distribution of S. zimocca
(5% of the total catch) is given in Table 6. The distribution by shape and size of S. zimocca per area and depth is
included in Table 7.
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CASTRITSI-CATHARIOS ET AL.
TABLE 6. Number of S. zimocca individuals harvested in the sampling areas and their % participation in the total catch (data
from the fishing fleet, key forms).
LIBYA
CRETE
DODEKANISSA
CYCLADES
LAKONIA
IONIAN SEA
REST AEGIAN
302
8.0%
267
2.4%
345
12.7%
127
7.1%
0
0%
0
0%
37
6.3%
Total catch = 1078 individuals, i.e. S. zimocca represents 5% of the total catch
TABLE 7. Frequency of appearance and depths at which S. zimocca was collected by the fishing fleet.
CRETE
1–20 m
21–40 m
41–60 m
1–20 m
21–40 m
41–60 m
61–80 m
81–100 m
BS
8
30
0
5
2
2
0
0
BM
86
0
0
26
57
23
13
3
Depth
Shape
And
Size
DODEKANISSOS
BL
26
0
0
32
43
7
0
5
TB
120
30
0
63
102
32
13
8
AS
10
0
0
0
19
0
0
0
AM
76
0
0
0
65
3
2
0
AL
31
0
0
0
63
2
0
0
TA
117
0
0
0
120
5
2
0
GT
237
30
0
63
222
37
15
8
continued.
CYCLADES
1–20 m
21–40 m
41–60 m
61–80 m
81–100 m
101–140 m
BS
0
0
0
0
0
0
BM
6
17
17
6
0
0
BL
18
7
12
0
0
0
TB
24
24
29
6
0
0
AS
5
0
2
0
0
2
AM
4
12
3
0
4
0
AL
2
5
5
0
0
0
TA
11
17
10
0
4
2
GT
35
41
39
6
4
2
Depth
Shape
And
Size
AS: shape A, size small / AM: shape A, size medium / AL: shape A, size large / TA: total A. BS: shape B, size small / BM:
shape B, size medium / BL: shape B, size large / TB: total B. GT: grand total.
The sponge fishing fleet did not find any S. zimocca in the Ionian Sea and in the Lakonia prefecture, at depths
1–40 m. In 2008, the Kalymnian divers harvested the following quantities of sponges in the areas previously mentioned:
-
H. communis: 2,000 kg, sizes 12.5–51 cm
S. o. officinalis: 1,700 kg, sizes 12.5–30.5 cm
S. o. mollissima: 300 kg, sizes 12.5–38 cm
S. zimocca and S. lamella were not harvested
Organic content of the fiber mass and skeletal density. Our results for organic content on samples from typical specimens of S. zimocca are given in Table 8. Comparative results for skeletal density of four other Mediterranean commercial sponges are presented in Table 9.
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TABLE 8. Calculation of the organic content for distinct commercial sponges.
Dry Weight/Wet Weight Average
0
/0
SD
S. zimocca
S. o. mollissima
S.o. officinalis S. lamella
H. communis
93.32
92.18
92.27
94.06
93.34
1.21
1.23
1.00
0.45
0.83
Inorganic Material/Dry
Weight 0/0
Average
9.17
14.23
11.38
30.10
16.15
SD
2.28
5.22
4.36
10.04
5.60
Organic Material/Dry
Weight 0/0
Average
90.83
85.77
88.62
69.90
83.85
SD
2.28
5.22
4.36
10.04
5.60
Inorganic Material/
Organic Material 0/0
Average
10.15
16.98
13.07
44.55
19.72
SD
2.79
7.54
5.85
20.77
8.11
6
7
5
2
7
Nr Of Samples
TABLE 9. Skeletal Density (gr cm-3), statistical data.
Species
N
Mean Value
Mean Deviation
Typical Deviation
S. zimocca
7
0.053707
0.006273
0.007902
S. o. offcinalis
4
0.055743
0.003183
0.004354
S. o. mollissima
9
0.049945
0.009146
0.011433
S. lamella
7
0.066438
0.015211
0.022291
H. communis
7
0.044999
0.022848
0.034300
Tensile strength. Castritsi-Catharios et al. (2007) suggest that both the tensile strength and the elasticity can be
used as two objective criteria for the evaluation of the quality and commercial value of commercial sponges. These
criteria could also provide additional taxonomic characters for the distinction of ill-defined species. Preliminary
results for the tensile strength and elongation of all five Mediterranean commercial species of sponges are presented in Table 10.
TABLE 10. Preliminary results for the tensile strength (in Kg cm-2) of commercial sponges.
Species
Nr. of specimens
Mean Value
Mean Deviation
Maximum Value Elongation (%)
Maximum Value
Elongation (%)
Hippospongia communis
7
1.3129
27.1000
0.3869
6.0286
Spongia lamella
6
4.0079
21.0167
2.0248
1.9889
Spongia. zimocca
6
1.8476
17.3000
0.2879
3.3333
Spongia o.officinalis
9
2.6802
20.7667
0.9609
3.6815
Spongia o. mollissima
5
2.2360
24.9400
0.5021
4.2080
Discussion
Sponges exhibit significant macro- and micro-morphological variations due to adaptive mechanisms to the environment (Gaino et al. 1995; Bell & Barnes 2000; McDonald et al. 2002; Meroz-Fine et al. 2005, Bell 2007). For
this reason, the shape of sponges, for a significant number of taxa, should not be considered as a reliable taxonomic
character. S. zimocca represents a typical example supporting this view. Its diversity of form is well known to both
the sponge divers and merchants. The structure of its populations and its wide distribution (from Tunisia to Egypt,
Cyprus, Cretan and Aegean Seas, absent from the Ionian Sea) suggest there probably is an interaction between habitat diversity and the diversity of forms in this species. In our study we wondered whether we should consider the
observed forms as intraspecific diversity or as evidence for the existence of subspecies. For this reason, we
searched for objective taxonomic characters which will be discussed below.
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FIGURES 4–6. Spongia zimocca. 4, Form A sponges harvested from Libya. 5, Characteristic specimen from Tunisia (upper
surface). 6, Characteristic specimen from Tunisia (surface of attachment).
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FIGURES 7–9. Spongia zimocca. 7, Compact form found between Cape Sounion and Aegina Island. 8, Specimen LMJG
15470 from the Graz Museum collection. 9, Sponges harvested by gagava trawler from Posidonia beds.
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FIGURES 10–13. Spongia zimocca. 10, Sponge from deep waters of Dodekanissos. 11, Well developed lobes, ¾ of their
length being free. 12, Linear channel starting from osculum, as described by Schmidt (1862). 13, Sponge just harvested from
Agathonissi Island (ca. 60m depth.).
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FIGURES 14–16. Spongia zimocca. 14, Sponge just harvested from Astipalea Island (ca. 18m depth.). 15, SEM of conules
with pinacodermal membrane carrying many foreign structures. 16, Primary fiber surrounded by a dense network of secondary
fibers in a specimen from Libya.
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FIGURES 17–20. Spongia zimocca.17, End of primary fibers in a Tunisian specimen. 18, Primary and secondary fibers from a
Libyan sponge. 19, Primary and secondary fibers from a Tunisian specimen. 20, Secondary fiber from a sponge collected near
Agathonissi, at 60m depth.
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FIGURES 21–26. Spongia zimocca. 21, Secondary fibers of a sample collected from Egypt. 22, Secondary fibers from a specimen obtained from shallow depths between Cape Sounion and Aegina Island. 23, Secondary fibers of a sample collected at
great depth from the Aegean Sea. 24, Pseudo-tertiary fibers of a specimen collected from Libya. 25, Pseudo-tertiary fibers from
a North African specimen. 26, Pseudo-tertiary fibers of a sample collected at great depth from the Aegean Sea.
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FIGURES 27–28. Spongia zimocca. 27, Secondary and pseudo-tertiary fibers of a specimen from Aegean Sea deep waters
(60–70 m). 28, Primary fiber protruding from the conule of a sponge harvested at Anafi Island in 2009.
The external macromorphology of S. zimocca exhibits differences which are probably correlated to its geographical distribution. We tried to group all these morphotypes under the forms Α, Β1 & Β2, but we have occasionally collected intermediate forms during our samplings. According to the view of Kalymnian divers and by
examining all the taxonomic characters previously mentioned we conclude that all these forms belong to the same
species. In the future it would be worthwhile to perform a genetic approach to the problem.
Cook & Bergquist, 2002 p. 1054 report that “Spongia species are very difficult, if not impossible, to distinguish
solely on skeletal morphology. However, there are some skeletal characteristics which are consistent across groups
of species”.
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FIGURE 29. Spongia zimocca (collected from Naxos Island at 60m depth). Spherical choanocyte chamber 20 μm in diameter,
choanocyte cells with characteristic shape and the collar of microvilli surrounding the flagellum.
The diameter (thickness) of the primary fibers and the presence or absence of foreign materials, as well as the
diameter and the texture of the secondary fibers, constitute taxonomic characters of specific importance for the
Spongia species, which have already been used for the characterization of S. zimocca.
Schmidt (1862) reported the thickness of secondary fibers to be equal to 20.4–33.8 μm in the dry samples he
examined, while in the depiction of the mesh of primary and secondary fibers, the texture of the primary fiber is
smooth, similar to that of the secondary fibers. Burton (1934) found the thickness of secondary fibers to equal
13.4–30.2 μm and de Laubenfels (1948) equal to 40 μm. The latter author reported the diameter of primary fibers to
be 55 μm without mentioning inclusion of any foreign particles. Vacelet (1959) pointed out that Schulze (1879),
Topsent (1925) and Arndt (1937) had observed the presence of primary fibers in S. zimocca without referring to the
inclusion of foreign particles, while they had found diameters for the secondary fibers of 30–45 μm. Later, Vacelet
(1987) reported inclusion-free primary fibers with 50–70 μm in diameter, while the secondary ones were 30–40 μm
thick. These data are summarized in Table 11.
TABLE 11. Thickness of secondary fibers according to different authors (data from the bibliography).
Author
Thickness of secondary fibers (μm)
Schmidt (1862)
20.4–33.8
Schulze (1879)
30–45
Topsent (1925)
30–45
Burton (1934)
13.4–30.2
Arndt (1937)
30–34
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In our effort to understand the differences observed in the measurements of the diameter of the secondary fibers in different specimens and by different authors, we investigated whether thickness varies relative to the position
of the sectioned sample within the sponge. We divided the sponge into three parts: base (close to the substratum),
middle and upper surface. ANOVA was applied on the diameter values of a relatively small number of specimens
from all the commercial sponge species. Our results showed no significant differences for S. zimocca, S. o. officinalis and H. communis (p > 0.05), indicating a homogeneous distribution of the diameter values within the same
sponge and that the aforementioned differences are due either to sampling errors or to the spread of the measurements. On the contrary, significant differences were found among the samples taken from the three parts for S.
lamella and S. o.. mollissima (p < 0.05), showing a non-homogeneous distribution of the diameter values within the
same sponge. The measurements also indicated that even in the case of homogeneity within the same sponge there
is a tendency in secondary fibers to be thinner near their base (ends nearer to the substratum). A statistically significant difference (p < 0.05) was found between the maximum and minimum diameter values of the secondary fibers,
which cannot be credited to the position of the sample within the sponge. However, it is also important to highlight
that a single fiber within a single mesh do not display the same diameter along its entire length, being thinner in the
middle and thicker in the nodes. All the above observations lead us to the conclusion that a protocol must be established for the measurements in which the origin of the sponge, the depth, the position of the sectioned sample
within the sponge (base, middle, upper part) and, mainly, the position on the fiber where the measurement was
taken should all be recorded.
In nearly all the samples of S. zimocca examined (Greece, Egypt, Libya, Tunisia), including the lectotype, we
observed pseudo-tertiary fibers, the number of which varies from one sponge to another, with diameter at the middle of the fibers (half way between both nodes) of 4.7–11.9 μm, and of 6.9–18.4 µm near the nodes (Table 3). More
specifically, in all the samples originating from greater depth in the Aegean Sea we observed a dense reticulation
consisting of these pseudo-tertiary fibers.
Cook & Bergquist (2001; 2002 p. 1051, 1054 & 1055) argued that no true tertiary fibers occur in Spongia, but
only thinner secondary fibers or "pseudo-tertiary fibers” in some species from Australasia. According to their revision, such pseudo-tertiary fibers characterize a subgenus Heterofibria. In the type species, Spongia (Heterofibria)
cristata, the secondary fibers are 17–42 µm in diameter and the thinner ones (pseudo-tertiary) are 4–17 µm. These
diameters are similar to those found in S. zimocca. It might thus be considered that S. zimocca belongs to this subgenus. However, there are several reasons for maintaining it in Spongia subgenus Spongia:
1) The 7 species which were allocated to Heterofibria are from Australasia (6) and Brazil (1), very far from the
Mediterranean.
2) S. (Heterofibria) cristata has primary fibers cored, when S. zimocca has uncored primary fibers.
3) The distinction of subgenus Heterofibria does not appear sound, as thinner secondary fibers are sometimes
found in Spongia o. officinalis and in Spongia lamella, and are regularly present in Spongia virgultosa, (with two
diameters of secondary fibres, respectively 40–50 µm and 7–20 µm in diameter) (Vacelet 1959).
Another morphometric character, which might constitute a taxonomic criterion, is the conules. These are produced by the projections of the terminations of the primary fibres. Their size, density, shape and the frequency of
appearance, vary from one species to another. In order to compare the five commercial species of sponges, our
measurements for this character have been placed in Table 10, from where a clear morphological differentiation
arises. This taxonomic character must be further studied in connection with the diversity of forms, especially for S.
zimocca.
The ratio (100 x (wet weight minus dry weight) / wet weight) shows the percent moisture content of the sponge
(on wet basis). This is actually the equilibration moisture since the specimen was exposed at 21 oC and 63% relative humidity for 48 hours inside a conditioning chamber. S. lamella (5.9%) followed by H. communis (6.7%) and
then by S. zimocca (6.7%) appear to have lower absorbing ability from the atmosphere in comparison to S. o. mollissima (7.8%) and S. o. officinalis (7.7%).
The percent amount of inorganic material constitutes a safe qualitative criterion since the commercial value of
a sponge increases as the amount of foreign particles decreases. The ratio (100 x (dry weight minus organic material) / dry weight) shows the percentage (on a dry basis) of the inorganic material within the sponge thus defining
its pureness. As shown in Table 8, S. zimocca contains the smallest amount of inorganic material (9.2%) against all
the other commercial species of sponges, i.e. S. o. officinalis (11.4%), S. o.. mollissima (14.3%), H. communis
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(16.2%) and S. lamella (30.1%). Despite this fact, the commercial value of S. zimocca is relatively low, depending
on its shape and origin, due to the texture of its skeleton (how it feels when touched), as discussed below. We
remind that inclusions are found outside of the fibers, inside the meshes, and partially remain even after the
mechanical treatment.
Although preliminary, our findings on the tensile strength give an indication of the resistance and the elasticity
of the commercial sponges. H. communis exhibits higher elasticity followed by S. o mollissima while S. lamella
shows the higher resistance to draw followed by S. o officinalis and then by S. o. mollissima. S. zimocca shows the
lowest resistance to draw and lowest elasticity of all the commercial species of sponges. These findings are in full
accordance with the classification done by divers and sponge merchants concerning the resistance and the texture
of their products. Interest in these measurements for commercial sponges has already been demonstrated (Moore
1908; Castritsi-Catharios et al. 2007; Louden et al. 2007).
The geographical distribution of S. zimocca in the Mediterranean has been reported by several researchers: in
Greek Seas, mainly Crete and Dodekanissos (Schmidt 1862; Schulze 1879; Castritsi-Catharios 1995, 1998; Kefalas
et al. 2003a; 2003b; Voultsiadou 2005); in Egypt (Castritsi-Catharios et al. 2005); in Libya (Castritsi-Catharios
1995, Milanese et al. 2007) and in Morocco (Voultsiadou, 1994). Topsent (1925), Rützler (1976) and Ben Mustapha et al (2003) report that S. zimocca is found in Tunisia, together with H. communis and S. lamella. The species
is also possibly present in the Western Mediterranean, as it has been reported once by Guella et al. (1992) south of
Livorno and by Zubia et al. (1994) in the area of Mazara del Vallo (Channel of Sicily). However, these records are
given with reserves due to possible wrong identification. In summary, S. zimocca appears to be mainly localized in
the eastern Mediterranean basin with a possible presence in the western basin (Ligurian Sea).
We cannot find a possible explanation for the apparent complete absence of S. zimocca from the areas of Evia
Island, Laconia and the Ionian Sea; as well as for its presence as form B in the North Aegean Sea, and its abundance in the area of Dodekanissos. Since in both Libya and Egypt the commercial sponges are almost exclusively
H. communis and S. zimocca, we investigated the distribution of the first species in the other areas. We found that
in Crete 4,952 individuals of H. communis were harvested, in Dodekanissos 1,036, in Lakonia 501, in Cyclades
485 and in the North Aegean 243, while it was completely absent from Evia Island and the Ionian Sea. We
observed highest abundance and diversity of commercial species of sponges in Crete, Dodekanissos and Cyclades.
More specifically, for S. zimocca we found that the area of Dodekanissos represents its optimal habitat (based on
abundance).
Acknowledgements
We gratefully acknowledge Dr. Karl Adlbauer, the Graz Museum, for supplying the specimen coded LMJG 15470.
We also acknowledge the two reviewers and the editor for their constructive remarks. Finally, the authors thank the
diver Mr. Kalomoiros Sakellaris and the gagava owner and president of the divers union Mr. Manolis Saroukos for
the information supplied to them regarding the fresh material.
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