Invasive Species Compendium

Detailed coverage of invasive species threatening livelihoods and the environment worldwide


Celtodoryx ciocalyptoides



Celtodoryx ciocalyptoides


  • Last modified
  • 13 November 2018
  • Datasheet Type(s)
  • Invasive Species
  • Preferred Scientific Name
  • Celtodoryx ciocalyptoides
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Porifera
  •       Subphylum: Cellularia
  •         Class: Demospongiae
  • Summary of Invasiveness
  • Celtodoryx ciocalyptoides is an encrusting, massive or globular sponge, usually yellow to pale brown, if alive, or beige in ethanol. It is found in semi-enclosed littoral environments, from 0.6 to 38 m of depth...

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Preferred Scientific Name

  • Celtodoryx ciocalyptoides Henkel & Janussen, 2011

Other Scientific Names

  • Celtodoryx girardae Pérez, et al. 2006
  • Coelosphaera physa Sim & Byeon, 1989
  • Cornulum ciocalyptoides Burton, 1935
  • Homoeodictya ciocalyptoides Koltun, 1971
  • Isodictya ciocalyptoides Henkel & Janussen, 2011

Summary of Invasiveness

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Celtodoryx ciocalyptoides is an encrusting, massive or globular sponge, usually yellow to pale brown, if alive, or beige in ethanol. It is found in semi-enclosed littoral environments, from 0.6 to 38 m of depth. It is native to the Northwest Pacific (Russian Far East, China, Japan and the Republic of Korea), and has been introduced to France and later to the Netherlands. Because of its ability to grow rapidly and cover large areas, the species poses a serious threat to native habitats, quickly becoming dominant within the benthic macrofauna of shallow waters (as has occurred in the Gulf of Morbihan, France, and in Dutch inshore waters). Aquaculture of the Pacific oyster has been assumed to be the probable introduction vector for the invasive sponge species to Europe.

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Porifera
  •             Subphylum: Cellularia
  •                 Class: Demospongiae
  •                     Subclass: Ceractinomorpha
  •                         Order: Poecilosclerida
  •                             Family: Coelosphaeridae
  •                                 Genus: Celtodoryx
  •                                     Species: Celtodoryx ciocalyptoides

Notes on Taxonomy and Nomenclature

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The higher classification of the phylum Porifera was revised in 2015 based on molecular sequence data (cf. Morrow and Cárdenas, 2015). 

Celtodoryx ciocalyptoides was first collected by Tarasov in 1926 from Posiet Bay, Sea of Japan, near the Russian port Vladivostok (spelt wrongly as “Tazasov” in Henkel and Janussen (2011)). Burton (1935) identified and described the material as Cornulum ciocalyptoides, which type series are present in the Zoological Institute of the Russian Academy of Sciences, Saint Petersburg (ZIN RAS). However, when the type material (plus material from other localities of the Northwest Pacific) was compared with a new invasive European sponge species Celtodoryx girardae, described by Pérez et al. (2006), it was found that C. girardae and Cornulum ciocalyptoides are in fact the same species. Therefore, Henkel and Janussen (2011) transferred the species from Cornulum (Acarnidae) to Celtodoryx (Coelosphaeridae) and consequently established Celtodoryx ciocalyptoides (Burton, 1935) as the senior synonym of Celtodoryx girardae.


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Sponges have a simple level of organisation: there are no tissues or organs, but specialized cells for various functions (Hooper and Van Soest, 2002; Van Soest et al., 2012). For example, T-shaped or flattened pinacocytes cover the outside of the sponge; flagella-bearing choanocytes line the internal system of canals and microscopic chambers and generate the water currents necessary for the filtering activity; individual cells of the mesohyl fill the space between canals and chambers and, together with skeleton structures, form a collagenous matrix that supports the animal. Elements of the sponge skeletons include discrete siliceous or calcareous spicules and/or an organic collagenous protein called spongin. Sponges with siliceous spicules (either monaxonic or tetraxonic, never triaxonic in shape) are united in the class Demospongiae, the largest class in the phylum Porifera. Species of the Demospongiae are predominantly leuconoid, having a complex structure, composed of a mass of flagellated chambers and water canals.

The siliceous spicules of the Demospongiae are divided into megascleres, which strengthen the framework of the sponges, and microscleres, which possibly have defensive or supportive functions. Morphology of the spicules is an important taxonomic character. The order Poecilosclerida includes sponges with the ‘chelae’ microscleres type. The family Coelosphaeridae is diagnosed by a plumose to plumoreticulate choanosomal skeleton of ascending tracts consisting of megascleres of two types: tylotes (thin, usually short, straight with well-defined equal tyles) and anisostrongyles (straight, or slightly curved, with less and stronger spines on extremities and generally longer and thicker than the tylote type), both with terminal spines fanning out towards the surface, loosely connected (Pérez et al., 2006; Henkel and Janussen, 2011). The ectosomal (outer) skeleton has a loose tangential arrangement of scattered anisostrongyles/tylotes and microscleres. Microscleres (arcuate isochelae of two distinct size categories and oxychaetes (thin spines)) of one size category are distributed randomly within the choanosome.

Detailed spicules morphology and dimensions characteristic of C. ciocalyptoides can be found in Henkel and Janussen (2011). The authors state that length and diameter of spicule categories do not vary between specimens from two different oceans, or within the representatives of the respective areas. However, the differences found between the authors' material and the type series are explained by the fact that spiculogenesis and skeleton growth may be affected by fluctuations of both dissolved silica concentrations in seawater and water temperature.

Living specimens show encrusting, massive or globular form with irregularly lobate to honeycombed surface. In China, most specimens are quince-yellow to golden yellow (colour turning to whitish grey or brownish after fixation), with very soft, non-elastic texture, easy to cut or tear. Surface is smooth, producing large amounts of mucus after cut off. The sponge has a rugose, thickly incrusting base (mean thickness less than 3 cm) and fistulas on the surface measure from a few millimetres to 1 cm in length. A number of freshly collected specimens had conspicuous brown spots on the surface (Henkel and Janussen, 2011).

The size of the sponge varies significantly between representatives from different oceans (Henkel and Janussen, 2011). C. ciocalyptoides from the Northwest Pacific tends to be encrusting, with a limited spatial extension: type material is represented by rather small pieces of the sponge (1.5-2.5 cm) and observed living specimens had an area of less than 20 cm2. On the other hand, the Northeast Atlantic specimens are often massive or thickly encrusting, and of large individual size, covering areas up to 25 m2 (recorded from Oosterschelde, Netherlands; Pérez et al., 2006; Van Soest et al., 2007). Thickness ranges from a few centimetres (for all localities) to a maximum of 50 cm in populations from the Atlantic.

Foreign material such as sediment particles, diatoms and rhodophytes are usually abundant throughout a sponge’s body, which is often the cause of darkly coloured layers and/or patches. Thus, according to the description by Burton (1935), C. ciocalyptoides exhibits a blackish external layer in preserved conditions. Several specimens from the Chinese Yellow Sea also have patches of brownish crust (Henkel and Janussen, 2011).


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C. ciocalyptoides is native to the Northwest Pacific, more specifically from Russian Far East, China, Japan and Republic of Korea. Outside its native range, C. ciocalyptoides was recorded (as Celtodoryx girardae) in 1996 in a well-studied area in the Ria of Etel, Brittany, France and later, over several years successively, in the Gulf of Morbihan, France (Pérez et al., 2006). These French observations were confirmed in 2000 and 2001 and, within two years, growth forms changed from globular to massive and easily recognisable specimens. The species has also been recorded from the Oosterschelde estuary, Netherlands. In the shallow waters of the Gulf of Morbihan and in Dutch inshore waters, the species has been regarded as being among the dominant benthic macrofauna (Pérez et al., 2006).

C. ciocalyptoides may have been introduced and be present in a much wider area than what has been recorded so far (Henkel and Janussen, 2011).

Distribution Table

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The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes

Sea Areas

Atlantic, NortheastWidespreadIntroduced Invasive Pérez et al., 2006; Van Soest et al., 2007As Celtodoryx girardae. In French and Dutch waters
Pacific, NorthwestWidespreadNativeKhodakovskaya, 2003; 2005; Koltun, 1959; 1971; Burton, 1935; Hoshino, 1987; Sim and Byeon, 1989; Henkel and Janussen, 2011Russian Far East, Japan, Korean and Chinese waters


ChinaPresentNativePresent based on regional distribution
-LiaoningWidespreadNativeHenkel and Janussen, 2011Chinese Yellow Sea, around Dalian (Lv Shun, Dalian Wan Bay, Er Tuo Islands and Fujizhuang Beach)
JapanPresentNativeHoshino, 1987
Korea, Republic ofPresentNativeSim and Byeon, 1989Near Anhung


FrancePresentIntroduced1996 Invasive Pérez et al., 2006As Celtodoryx girardae. Ria of Etel, Brittany
NetherlandsPresentIntroduced2005 Invasive Van Soest et al., 2007; Henkel and Janussen, 2011Well established in the Oosterschelde estuary, where it was first reported
Russian FederationPresentNativePresent based on regional distribution
-Russian Far EastPresentNative Not invasive Khodakovskaya, 2003; 2005; Burton, 1935

History of Introduction and Spread

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It has been suggested that C. ciocalyptoides was introduced to the Northeast Atlantic from the Northwest Pacific via aquaculture, with the Pacific oyster Crassostrea gigas as the probable vector (Pérez et al., 2006; Henkel and Janussen, 2011). The invasion may have started after the 1960s, when a batch of C. gigas was imported from British Columbia and Japan to the Oosterschelde estuary, Netherlands and the Gulf of Morbihan, France.

Risk of Introduction

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Further spread of C. ciocalyptoides in the European North Atlantic shores has been suggested (Henkel and Janussen, 2011).


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The species is usually found in semi-enclosed environments on rocky substrates, mussel shells and on soft-bottoms, being abundant on gentle slopes and rare on vertical cliffs (Pérez et al., 2006; Henkel and Janussen, 2011). Water circulation in such localities is usually quite intense and primarily driven by a semi-diurnal tide.

According to Henkel and Janussen (2011), in the native range the species is found at depths between 2.5 and 16 m. In Peter the Great Bay, Sea of Japan, C. ciocalyptoides has been found  at depths of 0.6-16 m, in a variety of substrata, such as stone, sand, pebble, mud, shells of the living bivalve molluscs Arca boucardi and Crenomytilus grayanus, and in roots of Zostera (Khodakovskaya, 2005).

Records from the Northeast Atlantic (North Sea, Oosterschelde, Netherlands) and the Gulf of Morbihan (Brittany, France) have been found at depths of 2.5-38 m (Pérez et al., 2006; Van Soest et al., 2007). Maximum density and size of the sponge in the Atlantic were observed between 8 and 10 m before 2004, but later in areas deeper than 15 m.

Habitat List

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Biology and Ecology

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Reproductive Biology

As the majority of Demospongiae, C. ciocalyptoides develops through a larval stage. Embryos of the species, as found in Chinese specimens, are round, flattened, 195-370 mm wide, slightly orange, containing close-packed cells and distributed abundantly within the choanosome (Henkel and Janussen, 2011). After hatching, sponge larvae become part of the plankton, drifting in the water column for a limited time period before settling on the seafloor to become sessile juvenile sponges.

Population Size and Density  

Specimens from the native range in the Pacific Ocean usually cover areas of less than 20 cm2 (Henkel and Janussen, 2011), whereas in the Northeast Atlantic Ocean large representatives, with a maximum thickness of 50 cm, have been observed covering an area of 25 m2 (Van Soest et al., 2007). Records by Pérez et al. (2006) in the Gulf of Morbihan (France) indicate that larger forms are restricted to shallow waters, whereas thinly encrusting forms occur in deeper waters.


Nutrition of C. ciocalyptoides is typical for sponges, namely by filtering microscopic-size food particles from water.


As shown by Henkel and Janussen (2011), in Chinese waters C. ciocalyptoides is often associated with rhodophytes (red algae), which are in some cases completely incorporated into the sponge tissue. Van Soest et al. (2012) note a great diversity of symbiotic organisms that often thrive inside, or on the body of, a sponge, from microscopic prokaryotes to macroscopic organisms such as shrimps, polychaetes, hydrozoans and fishes. In France, ophiurids may temporary lie on the sponges and the hairy crab Pilumnus hirtellus often burrows at the base of the sponge (Pérez et al., 2006)). A full list of the flora and fauna associated with the species (in the Atlantic) can be found in Pérez et al. (2006).

Environmental Requirements

C. ciocalyptoides generally requires high water temperature, although it seems to withstand noticeable seasonal variation (4-25°C). Koltun (1971) assumes that high summer temperatures (up to 26°C in lagoons and bays) allow this typically subtropical sponge species to live in Far East waters (Posiet Bay).

In the native range, seasonal temperature differences may be more pronounced than in Western European waters (Henkel and Janussen, 2011). For example, the climate in Dalian, China, is monsoon-influenced, humid and continental, characterized by humid summers and cold, windy, dry winters, with regular icing. It has been suggested that cold winter periods may cause an adverse effect on C. ciocalyptoides growth  (Henkel and Janussen, 2011). Similarly, although the climate in the Northeast Atlantic is moderate, with relatively cool summers and mild winters with infrequent icing, low winter temperatures may have been the cause of a mass mortality event of C. ciocalyptoides in the Gulf of Morbihan in 2003 (Pérez et al., 2006). After this, the sponge population recovered, but its growth forms are now mainly encrusting, rather than massive globular.

All Chinese localities where the species is present are isohaline (31-32.5 ppm), eutrophic and with low concentrations of silica (0.01 mmol/m3). Similarly, in the Atlantic, salinity ranges from 32 to 32.5 ppm (Pérez et al., 2006). All locations where C. ciocalyptoides is found are known for eutrophication-induced nutrient enrichment, indicating that the sponge seems to tolerate strongly eutrophicated waters (GISD, 2015).

Latitude/Altitude Ranges

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Latitude North (°N)Latitude South (°S)Altitude Lower (m)Altitude Upper (m)

Water Tolerances

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ParameterMinimum ValueMaximum ValueTypical ValueStatusLife StageNotes
Depth (m b.s.l.) 0.6-38 m tolerated
Salinity (part per thousand) 31-32.5 ppt tolerated; Henkel and Janussen (2011)
Water temperature (ºC temperature) 4-25 °C tolerated; Henkel and Janussen (2011)

Means of Movement and Dispersal

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Natural Dispersal

Swimming capabilities of sponge larvae ensure short distance dispersal.

Accidental Introduction

The invasion of C. ciocalyptoides in Europe is considered to be directly related to the commercial transfer of the pacific oyster Crassostrea gigas to aquaculture farms in lagoons along the French and Dutch coasts (Henkel and Janussen, 2011). Although invasion pathways of C. ciocalyptoides cannot be easily determined, the fact that the invasive sponge is often found on oyster shells speaks in favour of this hypothesis.

Pathway Causes

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CauseNotesLong DistanceLocalReferences
AquacultureAccidentally from the North West Pacific through aquaculture of the Pacific oyster Crassostrea gigas Yes Pérez et al., 2006; Henkel and Janussen, 2011
HitchhikerAccidentally from the North West Pacific on Pacific oysters Yes Pérez et al., 2006; Henkel and Janussen, 2011

Pathway Vectors

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VectorNotesLong DistanceLocalReferences
Aquaculture stockAdult living sponges as epibionts on shells of Crassostrea gigas Yes Pérez et al., 2006; Henkel and Janussen, 2011
Host and vector organismsOn shells of Crassostrea gigas Yes Pérez et al., 2006; Henkel and Janussen, 2011

Vectors and Intermediate Hosts

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Crassostrea gigasPérez et al., 2006. MolluscPacific, Northwest

Environmental Impact

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Impact on Habitats

As noted by Pérez et al. (2006), С. ciocalyptoides, as other suspension-feeders, may impact the environment in different ways (see also Darrigran, 2002Pérez et al., 2005). The sponge may act as a biofilter, being able to collect and concentrate, or degrade, a wide range of pollutants, thus contributing to purification of colonized areas. On the other hand, sponges are able to release a large amount of inorganic matter (mainly nitrogen and phosphorus compounds) that may be a source of pollution to the invaded habitats. More recent research indicates an ability of sponges to take up dissolved organic matter, thus influencing dissolved organic carbon (DOC) concentration in marine waters (De Goeij et al., 2008). Therefore, encrusting sponges may be of importance in the removal of DOC in marine communities, such as coral reefs.

Impact on Biodiversity

When С. ciocalyptoides is dominant in the macrofauna (as in shallow waters of the Gulf of Morbihan and the Dutch coast), it can outcompete other macrobenthic organisms, overgrowing sessile invertebrates, such as other sponges and octocorals (Henkel and Janussen, 2011). In French waters, the species was found competing with the demosponge Esperiopsis fucorum [Amphilectus fucorum]  and the gorgonian Eunicella verrucosa (Pérez et al., 2006).

Risk and Impact Factors

Top of page Invasiveness
  • Proved invasive outside its native range
  • Has a broad native range
  • Abundant in its native range
  • Highly adaptable to different environments
  • Is a habitat generalist
  • Capable of securing and ingesting a wide range of food
  • Fast growing
  • Gregarious
  • Reproduces asexually
Impact outcomes
  • Damaged ecosystem services
  • Ecosystem change/ habitat alteration
  • Modification of natural benthic communities
  • Modification of nutrient regime
  • Reduced native biodiversity
  • Threat to/ loss of native species
Impact mechanisms
  • Competition - monopolizing resources
  • Competition
  • Filtration
  • Fouling
  • Rapid growth
Likelihood of entry/control
  • Highly likely to be transported internationally accidentally
  • Difficult to identify/detect in the field
  • Difficult/costly to control


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Social Benefit

Rashid et al. (2009) used exopolysaccharides (EPS) extracted from C. ciocalyptoides to study potential antiviral activity of the substance against the Herpes simplex virus.

Uses List

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  • Research model

Detection and Inspection

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In shallow water localities, living specimens can be collected via scuba diving, fixed in 6% formaldehyde and later transferred to 96% ethanol. Skeletal architecture may be observed in 200-400 mm thick sections under a light microscope (following Vacelet, 2006). Spicules can be obtained by dissolution of the sponge organic components in nitric acid and then examined and measured under light microscope, or by using Scanning Electron Microscopy (see Henkel and Janussen, 2011 for details).

Similarities to Other Species/Conditions

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C. ciocalyptoides may be confused with other species and genera of the family Coelosphaeridae (Henkel and Janussen, 2011). The genus Chaetodoryx is similar to Celtodoryx by the presence of oxychaetes, but in the former the skeleton is reticulate, consisting of choanosomal acanthostyles (spiny megascleres). A similar species from the family Coelosphaeridae, Lissodendoryx (Acanthodoryx) fibrosa is red and has a plumoreticulate skeleton with acanthostyles, rather than the plumose to plumoreticulate skeleton, lacks oxychaetes and has been exclusively recorded from tropical coral reefs in the Philippines.

Prevention and Control

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Because C. ciocalyptoides is a recent European invader, no specific prevention methods have been developed for the species. However, general methods of aquatic invasive species control and prevention, such as those reviewed by Gollasch (2002), can be used.

An European surveillance network and an appropriate adaptation of monitoring strategies are recommended (Nehring and Klingenstein, 2005), with the aim to assess risks, impacts and the invasiveness of the alien species, as well as the efficiency of eradication and control measures. Regulatory international measures against invasions in the European Seas are discussed in Gollasch (2006).

As sponges are commonly found in epibiotic fouling communities, recommendations regarding fouling treatment and control of the bay barnacle Amphibalanus improvisus (CABI, 2018) could be adapted to this species.

Methods to contain and mitigate the invasion by the Pacific oyster Crassostrea gigas (Herbert et al., 2012), which is thought to be the main vector responsible for introductions of C. ciocalyptoides, can be useful to not only control, but also monitor, populations of C. ciocalyptoides.

Gaps in Knowledge/Research Needs

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Because of the complexity of sponge taxonomy (Hooper and Van Soest, 2002), there is a lack of records of invasive sponge species, which results in deficiency of regular monitoring of sponge invasions. Related to this, there is also a lack of knowledge on sponge distributions, which prevents sound decisions on potential invading species. Thus, several sponge species suspected of invasive behaviour may in fact be non-invasive, not having been recognized or noticed previously as members of the local sponge fauna.

Further research should also involve the study of the impact of C. ciocalyptoides on native benthic communities and the probability of its proliferation on coastlines of Northeast Europe due to global warming (Henkel and Janussen, 2011).


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Burton M, 1935. Some sponges from the Okhotsk Sea and the Sea of Japan. Exploration des Mers de l’URSS, 22, 61-79.

CABI, 2018. Amphibalanus improvisus [original text by Ekaterina Shalaeva]. In: Invasive Species Compendium. Wallingford, UK: CAB International.

Darrigran, G. A., 2002. Potential impact of filter-feeding invaders on temperate inland freshwater environments. Biological Invasions, 4, 145-156.

De Goeij JM, Van den Berg H, Van Oostveen MM, Epping EHG, Van Duyl FC, 2008. Major bulk dissolved organic carbon (DOC) removal by encrusting coral reef cavity sponges. Marine Ecology Progress Series, 357, 139-151.

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Gollasch S, 2002. Hazard analysis of aquatic species invasions. In: Invasive aquatic species in Europe. Distribution, impacts and management, [ed. by Leppakoski E, Gollasch S, Olenin S]. Dordrecht, Netherlands: Kluwer Academic Publishers. 447-455.

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Henkel, D., Janussen, D., 2011. Redescription and new records of Celtodoryx ciocalyptoides (Demospongiae: Poecilosclerida) - a sponge invader in the north east Atlantic Ocean of Asian origin?. Journal of the Marine Biological Association of the United Kingdom, 91(2), 347-355. doi: 10.1017/S0025315410001487

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Koltun VM, 1959. Kremnerogovye gubki severnykh i dal’nevostochnykh morei SSSR (Otryad Cornacuspongida) (Cornacuspongidan sponges of the Northern and Far Eastern Seas of the USSR (Order Cornacuspongida)). Identification Books of the USSR Fauna, 67. Moscow, Leningrad, USSR: Nauka, Zoological Institute of the USSR Academy of Sciences.235 pp.

Koltun VM, 1971. On the knowledge on the fauna of sponges of Pos’eta Bay, Sea of Japan. (Fauna i flora zaliva Pos’et Yaponskogo morya). Issledovaniya Fauny Morei (Explorations of Marine Fauna), 8(16), 22-30.

Morroz C, Cárdenas P, 2015. Proposal for a revised classification of the Demospongiae (Porifera). Frontiers in Zoology, 12(7), 1-27.

Nehring S, Klingenstein F, 2005. Alien species in the Wadden Sea - a challenge to act. Wadden Sea Newsletter, 31, 13-16.

Pérez T, Longet D, Schembri T, Rebouillon P, Vacelet J, 2005. Effects of 12 years' operation of a sewage treatment plant on trace metal occurrence within a Mediterranean commercial sponge (Spongia officinalis, Demospongiae). Marine Pollution Bulletin, 50, 301-309.

Pérez, T., Perrin, B., Carteron, S., Vacelet, J., Boury-Esnault, N., 2006. Celtodoryx girardae gen. nov. sp. nov., a new sponge species (Poecilosclerida: Demospongiae) invading the Gulf of Morbihan (North East Atlantic, France). Cahiers de Biologie Marine, 47(2), 205-214.

Rashid ZM, Lahaye E, Defer D, Douzenel P, Perrin B, Bourgougnon N, Sire O, 2009. Isolation of a sulphated polysaccharide from a recently discovered sponge species (Celtodoryx girardae) and determination of its anti-herpetic activity. International Journal of Biological Macromolecules, 44(3), 286-293.

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Van Soest RWM, de Kluijver MJ, van Bragt PH, Faasse M, Nijland R, Beglinger EJ, de Weerdt WH, de Voogd NJ, 2007. Sponge invaders in Dutch coastal waters. Journal of the Marine Biological Association of the United Kingdom, 87, 1733-1748.

Principal Source

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14/09/15 Original text by:

Ekaterina Shalaeva, Consultant, UK

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