Marine Ecology. ISSN 0173-9565
ORIGINAL ARTICLE
A facies of Kophobelemnon (Cnidaria, Octocorallia) from Santa Maria di Leuca coral province (Mediterranean Sea) Francesco Mastrototaro1, Porzia Maiorano1, Agostina Vertino2, Daniela Battista1, Antonella Indennidate1, Alessandra Savini2, Angelo Tursi1 & Gianfranco D’Onghia1 1 Department of Biology, LRU CoNISMa, University of Bari, Bari, Italy 2 Department of Geological Science and Geotechnology, LRU CoNISMa, University of Milano-Bicocca, Milan, Italy
Keywords Bathyal facies; biodiversity; cold-water corals; Kophobelemnon stelliferum; Mediterranean Sea. Correspondence Francesco Mastrototaro, Department of Biology, University of Bari, LRU CoNISMa Bari Campus, Via E. Orabona, 4,70125 Bari, Italy. E-mail:
[email protected] Accepted: 8 October 2012 doi: 10.1111/maec.12017
Abstract During a research cruise carried out in April 2010, aimed at updating the knowledge on the biodiversity of the Santa Maria di Leuca (SML) cold-water coral province (Mediterranean Sea), a facies of the sea pen Kophobelemnon stelliferum (Muller, 1776) was found on mud-dominated bottoms. This finding represents a new species and a new habitat record from the SML coral province as well as a new bathyal facies in the whole Central Mediterranean Sea. The colonies were collected using an epi-benthic sledge, at depths between 400 and 470 m. A significant positive relationship between polyp number and colony length was detected. Density of the colonies ranged from 0.003 to 0.038 Nm 2. Differences and affinities between Mediterranean and Atlantic occurrences of K. stelliferum are discussed.
Introduction The Santa Maria di Leuca (SML) coral province is located along the Apulian margin, a few miles off Cape Santa Maria di Leuca in the Northern Ionian Sea (Central Mediterranean; Fig. 1), where a core of oxygenated waters (12.92 °C; 38.6 psu) of Adriatic origin comes from the Otranto channel moving in geostrophic balance along the isobaths at 500–1000 m in depth (Budillon et al. 2010) and carrying particulate matter and nutrients that provide a crucial source for biological activity in deep waters (Civitarese et al. 1998; De Lazzari et al. 1999). During the APLABES project, (Corselli 2010) an area of 800 km2 was mapped by means of multibeam sonar, between 200 and 1300 m in depth. This area includes (i) a broad eastern sector, influenced by mass-transport deposition, with a very complex hummocky sea floor consisting of widespread mound-like reliefs, (ii) a central ridge where drift sedimentation has been recognised by documenting the action of contour currents from the northeast and (iii) prominent fault-scarps to the west, where widespread erosion processes are evident from the emergence of stiff and/or hardened older sediments (Savini & Corselli 2010; Marine Ecology 34 (2013) 313–320 ª 2013 Blackwell Verlag GmbH
Fig. 2). In the whole mapped area, 10 sites have been explored by means of underwater video recording systems (Etiope et al. 2010; Fig. 2; MS1–MS8). Two of these sites (MS04-Atlantis Mound and MS06-Yellow Chain) were accurately mapped and described in Vertino et al. (2010). In each of them, seven main benthic macrohabitats were identified: 1-CF: Coral Framework; 2-LCF: Loose Coral Framework; 3-CF/H: Coral Framework and Hardground; 4-H: Hardground crusts and boulders; 5-CR: Coral Rubble; 6-BR: Buried Rubble; 7-BS: Bioturbated finegrained Sediment. Information on the biodiversity of the SML coral province has been recently updated by Mastrototaro et al. (2010), who report a list of 222 taxa (202 at species level). The greatest number of benthic taxa belong to Porifera (36), Mollusca (35) and Cnidaria (31). Annelida, Crustacea and Bryozoa were found with 24, 23 and 19 species, respectively. A total of 40 benthopelagic fish species were collected. Other faunal taxa, such brachiopods and echinoderms, were recorded with smaller numbers of species. In all, 135 species were new for the SML coral province, 31 of which represented new records for the Northern Ionian Sea (Mastrototaro et al. 2010). Refuge effects from fishing were also detected in 313
A facies of Kophobelemnon in the Mediterranean Sea
Mastrototaro, Maiorano, Vertino, Battista, Indennidate, Savini, Tursi & D’Onghia
Fig. 1. Study area. (a): Location within the Mediterranean Sea. (b): Bathymetric map of the area with details of the SML Coral Province.
this coral province (D’Onghia et al. 2010, 2011). Indeed, the sea floor roughness created by the presence of a close arrangement of coral-topped mounds and of a number of fault scarps, seems to protect the area from fishing (which is too risky for the safety of fishing gear), preserving the biocenoses of both mounds and also of the still poorly known intermound meso habitats. In this paper, a facies of the sea pen Kophobelemnon stelliferum (Mu¨ller, 1776), recently discovered on soft bottoms from the SML coral province, is described, updating knowledge on the biodiversity and benthic habitat typologies of the SML cold-water coral province. Furthermore, a detailed description of the morphotype of K. stelliferum is provided in order to comment on the taxonomic position of this Atlanto-Mediterranean species and the supposed endemic Mediterranean Kophobelemnon leuckartii Kölliker, 1872.
Fig. 2. Map of the study area with indication of the sites explored by means epi-benthic sledge (BT) and by underwater recording systems (MS). A mass transport deposit (MTD) is evident on track line of BT54.
Table 1. List of sampling stations in which colonies of Kophobelemnon stelliferum were found with indication of cruise code station (St), minimum and maximum depth, starting (S) and ending (E) sledge coordinates, swept area (m2), colony numbers (N) and density (Nm 2).
st
min–max depth
BT 47
429–451
BT 48 BT 54 BT 64 BT 65
423–467 454–457 412–446 404–446
Long E
swept area m2
N
density (Nm 2)
S 39°36.59′
18°22.42′
10,181
30
0.003
E 39°36.78′
18°20.40′ 11,999
241
0.020
9008
340
0.038
10,556
115
0.011
11,612
62
0.005
Lat N
S 39°36.06′
18°14.05′
E 39°35.57′
18°12.23′
S 39°35.85′
18°13.25′
E 39°35.38′
18°11.57′
S 39°36.58′
18°21.86′
E 39°36.73′
18°19.89′
S 39°36.58′
18°21.94′
E 39°36.73′
18°19.81′
Material and Methods During April 2010 a research cruise was carried out in the SML coral province, on board the R/V Universitatis, as part of the EU 7FP CoralFISH project. The sampling was carried out by means of an epi-benthic sledge with a fixed iron opening of 3.8 per 0.46 m and a mesh size of 0.5 cm. The sampling was carried out in five different stations at depths between 404 and 467 m on soft bottoms (BS habitat sensu Vertino et al. 2010) characterised by a gently sloping sea floor and a smooth surface, without mounds or any other seafloor irregularities (Fig. 2; Table 1). Each tow was carried out for about 30 min at a speed of about 3 knots. The steel cable of the sledge was 314
equipped with a dynamometer to take the real bearing of the sledge on the bottom. The area swept by the epi-benthic sledge was estimated for each sampling station according to the following formula: opening of the sledge 9 time of tow 9 speed. Subsequently, the density of Kophobelemnon stelliferum (number of colonies per m2) was computed for each station and an average density for the study area was calculated. All the species belonging to megafauna (>2 cm) were examined. The colonies of K. stelliferum were immediately anaesthetized with a saturated solution of menthol in seawater for 2–4 h according to the size of the colonies Marine Ecology 34 (2013) 313–320 ª 2013 Blackwell Verlag GmbH
Mastrototaro, Maiorano, Vertino, Battista, Indennidate, Savini, Tursi & D’Onghia
and contractions of polyps. After this procedure, the colonies were preserved in 5% seawater–formalin. All the colonies of Kophobelemnon were counted and 714 of them were measured (total length of the colony in mm). The colonies of Kophobelemnon consist of a primary polyp that forms the axis of the colonies called rachis and of a great number of secondary polyps divided into autozooids (polyps with well developed pinnular tentacles) and siphonozooids (polyps with a well developed siphonoglyph and without tentacles). For each colony the length of the peduncle (the lower part of the colony lacking polyps, including the terminal swelling called ‘end bulb’) and the rachis (the polypiferous part of the colony) were also measured and the number of autozooids were counted according to Bayer et al. (1983). Some colonies were examined using a stereomicroscope to identify the main taxonomic characters of the species, such as the distribution of the polyps on the rachis or the morphology of the pinnular tentacles of the autozooids. Other colonies were stained with Masson colouring (haematein in a saturated solution of potassium alum) to examine the small parts of the colonies such as the siphonozooids. Microscope slides with calcareous sclerites were prepared by collecting small portions of tissue from six different parts along the colony (1: end bulb, 2: along peduncle, 3: between peduncle and rachis, 4: lower portion of rachis, 5: upper portion of rachis, 6: expanded autozooid). Each portion of tissue was dissolved in sodium hypochlorite. After digestion, the sclerites were washed in distilled water and dehydrated with a sequence of alcohols (70%, 80%, 95% and absolute ethanol). Part of the sclerites were transferred to slides and observed using an optical microscope. Moreover, parts of the intact colonies were directly observed by SEM to emphasize the structure and distribution of the sclerites on the rachis. The sclerites, found in the different portions of the colonies, were split into the different types and measured (50 spicules of each type) using an image analyzer program (Nikon ACT-2U). The relationship between polyp number (autozooids) and total length of the colony was computed by means of linear regression analysis.
A facies of Kophobelemnon in the Mediterranean Sea
autozooids. The peduncle length ranged from 16% to 57% of the total length of the colony. The sclerites were distributed throughout the colony (Fig. 4). The sclerites, found at different sites of the colonies, were manly represented by three-flanged spindles with tuberculated ends varying in length from 70 to 350 lm. Shorter sclerites (about 50 lm) with tubercules throughout the entire length were also found at the base of the peduncle in the end bulb. The estimated density of the colonies at each station ranged from 0.003 to 0.038 Nm 2 (Table 1) with an average density of 0.0154 ± 0.014 Nm 2. The total length of the whole colonies varied from 40 to 181 mm (Fig. 5). A great number of colonies (65% of the total) showed lengths between 60 and 90 mm with a number of autozooids ranging from 8 to 12. The following significant relationship between polyp (autozooid) number (PN) and colony length (CL) was obtained: PN = 0.1118CL + 1.6569 (R2 = 0.629; P < 0.01; Fig. 6). With regard to the associated megafauna in the samples, a total of 26 species (one Cnidaria Pennatulacea, seven Mollusca, four Annelida Polychaeta, nine Crustacea Decapoda, four Echinodermata and one Pisces) were collected (Table 2). Some of them are characteristic species of bathyal mud, such as the gastropods Aporrhais serresianus and Ranella olearium, the bivalves Abra longicallus and Cuspidaria cuspidata, the cephalopod Scaeurgus unicirrhus, the crustacean decapods Aristaeomorpha foliacea, Chlorotocus crassicornis, Paromola cuvieri, Plesionika martia, Polycheles typhlops and Rochinia rissoana; some others are mud-loving species, such as the annelid Dasybranchus caducus, the crustacean decapods Nephrops norvegicus and Goneplax rhomboides and the echinoderms Amphiura chiajei, Eostichopus regalis and Molpadia musculus. Some eurybathic species were also found, such as the mollusc Galeodea echinophora and Euspira fusca and the echinoderm Cidaris cidaris. The species of sea pen belonging to the genus Distichoptilum could represent the first finding of this genus in the Mediterranean Sea. It is worth noting that Dasybranchus caducus, a non-selective deposit feeder polychaete, is the only species always present in the samples in which Kophobelemnon was collected.
Results A total of 788 colonies of Kophobelemnon stelliferum were sampled at depths between 404 and 467 m. The colonies showed a rose and greyish colour and a slender rachis with clavate upper end (Fig. 3a). The end bulb (a basal swelling) was present (Fig. 3b). Autozooids with pinnular tentacles (Fig. 3c) and very spiculated siphonozooids (Fig. 3d–g) were distributed on the rachis. The former were distributed in longitudinal rows, the latter were numerous in the area above the peduncle and around the Marine Ecology 34 (2013) 313–320 ª 2013 Blackwell Verlag GmbH
Discussion and Conclusions Kophobelemnon stelliferum is a sea pen widely distributed along the continental slope of the Atlantic and Pacific Ocean at depths from 40 to 2500 m (Williams 1990; Rice et al. 1992; Lo´pez-Gonza´lez et al. 2001; Moen & Svensen 2004). Facies of Umbellula and Kophobelemnon were described by Le Danois (1948) along the Atlantic coast of France and a facies of Kophobelemnon was described by Carpine (1970) on deep-sea muddy bottoms along the 315
A facies of Kophobelemnon in the Mediterranean Sea
a
c
b
d
Mastrototaro, Maiorano, Vertino, Battista, Indennidate, Savini, Tursi & D’Onghia
e
f g Fig. 3. (a): Kophobelemnon stelliferum colony sampled in the SML coral province. (b): Base of colony with end bulb. (c): Detail of autozooid with pinnular tentacles on the upper zone of the colony. (d): Siphonozooids distributed on the rachis. (e): Detail of siphonozooids under stereomicroscope. (f,g): SEM images of siphonozooids.
Catalan coast. More recently, Rice et al. (1992) described this facies in the Porcupine Seabight (Northeast Atlantic) near the coast of Ireland at depths from 365 to 1600 m. This sea pen is considered a characteristic species of bathyal mud as well as a canyon species (Carpine 1970). To date, this sea pen has only been recorded in the Mediterranean Sea in the western basin from Gibraltar to the North and Middle Tyrrhenian Sea (Le Danois 1948; Gili & Page`s 1987; Morri et al. 1991). Recently, both Hebbeln et al. (2009) and Pardo et al. (2011) have reported Kophobelemnon species (Kophobelemnon c.f. leuckartii Hebbeln et al. (2009) and K. stelliferum Pardo et al. (2011)) as common elements of the bathyal mud facies from the Alboran Sea. The finding of Kophobelemnon in the SML coral province therefore represents the easternmost record of the species in the Mediterranean Sea and 316
a new bathyal facies in the whole Ionian Sea. Most probably, the presence of this particular facies of the bathyal mud in the SML coral province is due to the protection of hard substrates and coral mounds from the trawl fishing which occurs around this area. Indeed, hard substrates interspersed with coral mounds and muddy bottoms are less accessible to trawl fishing activities and thereby can provide a natural refuge not only for mobile fauna, as reported by D’Onghia et al. (2011), but also for a vulnerable facies such as that of Kophobelemnon. It is well known that the Mediterranean bathyal mud is often covered by octocoral facies, such as the ones of the sea pen Funiculina quadrangularis and of the gorgonian Isidella elongata. The former represents an essential habitat for some commercial crustaceans, such as Parapenaeus longirostris and Nephrops norvegicus, the latter, in deeper Marine Ecology 34 (2013) 313–320 ª 2013 Blackwell Verlag GmbH
Mastrototaro, Maiorano, Vertino, Battista, Indennidate, Savini, Tursi & D’Onghia
A facies of Kophobelemnon in the Mediterranean Sea
Fig. 4. Distribution of the different types of sclerite in the Kophobelemnon stelliferum colony.
Fig. 5. Length–frequency distribution of the colonies of Kophobelemnon stelliferum collected in the SML coral province.
Fig. 6. Relationship between polyp number (PN) and colony total length (CL, mm) for the colonies of Kophobelemnon stelliferum collected in the SML coral province.
Marine Ecology 34 (2013) 313–320 ª 2013 Blackwell Verlag GmbH
317
A facies of Kophobelemnon in the Mediterranean Sea
Mastrototaro, Maiorano, Vertino, Battista, Indennidate, Savini, Tursi & D’Onghia
Table 2. List of the species associated with facies of Kophobelemnon found in the SML coral province (Ionian Sea) (number of colonies or specimens collected are indicated). species Cnidiaria Pennatulacea Kophobelemnon stelliferum (Mu¨ller, 1776) Distichoptilum sp. Ceriantharia Cerianthus sp. Mollusca Gastropoda Aporrhais serresianus (Michaud, 1828) Euspira fusca (Blainville, 1825) Galeodea echinophora (Linnaeus, 1758) Ranella olearium (Linnaeus, 1758) Bivalvia Abra longicallus (Scacchi, 1834) Cuspidaria cuspidata (Olivi, 1792) Cephalopoda Scaerurgus unicirrhus Brachipoda Griphus vitreus (Born, 1778) Annelida Polychaeta Dasybranchus caducus (Grube, 1846) Lumbrineridae sp. Maldanidae sp. Nephthys hystricis McIntosh, 1900 Crustacea Decapoda Aristaeomorpha foliacea (Risso, 1827) Chlorotocus crassicornis (Costa, 1871) Dardanus arrosor (Herbst, 1796) Goneplax rhomboides (Linnaeus, 1758) Munida intermedia A. Milde-Edwards & Bouvier, 1899 Nephrops norvegicus (Linnaeus, 1758) Paromola cuvieri (Risso, 1816)
318
BT 47
BT 48
BT 54
BT 64
BT 65
30
241
340
115
62
2
1 1
2
1
1
1 1
1
1
4 1
1
1
6
1
Table 2. Continued species
BT 47
Plesionika martia (A. Milne Edwards, 1883) Polycheles typhlops Heller, 1862 Rochinia rissoana (Roux, 1828) Echinodermata Ophiuroidea Amphiura chiajei Forbes, 1843 Echinoidea Cidaris cidaris (Linnaeus, 1758) Holothuroidea Eostichopus regalis (Cuvier, 1817) Molpadia musculus Risso, 1826 Pisces Osteichthyes Lophius budegassa Spinola, 1807
BT 48
BT 54
BT 64
BT 65
1
1
1
2
1
1
1
1
1
1
1
3
1 1
1
1
1
1
1
2
1
3
2
2
2
2
1 1 1
1
1 1 1 1 1
3 2
1
1
grounds, constitutes a selected habitat for the deep water shrimps Aristeus antennatus and Aristaeomorpha foliacea (Pe´re`s & Picard 1964). Both F. quadrangularis and I. elongata facies have almost completely disappeared due to trawl fishing in many Mediterranean areas (D’Onghia et al. 2003; Sarda` et al. 2004). This could also be true for K. stelliferum, although the actual distribution and abundance of this species in the Mediterranean is not known. The facies of Kophobelemnon found in the SML coral province showed lower colony densities than those estimated in Atlantic areas at the same depth (up to 2.6 colonies per m2 in Atlantic Porcupine Seabight; Rice et al. 1992). However, the densities computed in the present study could be underestimated due to the gear used, which is not the most suitable sampler for this species in the surveyed soft bottoms. In addition, a greater number of samples (200) were taken in the Atlantic Porcupine Seabight (Rice et al. 1992) than in the present study. Finally, the different densities estimated in the five stations of SML coral province could be related to the highly patchy distribution of the species. Video analyses performed on BS habitats of neighbouring sites (MS05 and MS06; Fig. 2) have indeed shown that pennatulaceans are heterogeneously distributed in the SML intermound areas (Vertino et al. 2010). In particular, in the MS05 site, the pennatulacean density, computed by dividing the total number of recognised specimens by the total area investigated during the dive, is around 0.015 Nm 2 Marine Ecology 34 (2013) 313–320 ª 2013 Blackwell Verlag GmbH
Mastrototaro, Maiorano, Vertino, Battista, Indennidate, Savini, Tursi & D’Onghia
and therefore comparable with the density estimated at station BT 64. The SML K. stelliferum colonies were smaller than those collected in the Atlantic (up to about 250 mm in total length; Rice et al. 1992). In addition, the sizes of sclerites were smaller than those reported for Atlantic colonies (210–570 lm in the rachis and 100–210 lm in the end part of the peduncle; Lo´pez-Gonza´lez et al. 2001). As known for other Mediterranean species, the particular size of colonies and sclerites of K. stelliferum collected in the Ionian Sea could represent a case of Mediterranean dwarfism due to the particular hydrological conditions of this basin with respect to those present in the Atlantic Ocean (Sanfilippo 2003; Massuti et al. 2004; Brunetti 2009). Although Ko¨lliker (1872) and subsequently Cecchini (1917) considered the Mediterranean colonies of Kophobelemnon to belong to an endemic species called K. leuckartii (in relation to the differences in size of the colonies and differences in size and distribution of the sclerites), Kukental (1915) considered the two species to be synonymous. In our opinion, according to Kukental (1915), the exclusive variability in size of both colonies and sclerites does not justify the existence of an endemic species of Kophobelemnon in the Mediterranean Sea. However, as above reported, the actual distribution of K. stelliferum in the Mediterranean Sea is still scarcely known; in particular, it is not known whether there is a genetic flux from Atlantic populations towards the Mediterranean. For these reasons, genetic studies are necessary to evaluate any molecular differences between the Atlantic and Mediterranean populations and, if there are, to consider the Mediterranean Kophobelemnon as a distinct species (sibling species). Currently, the taxonomic position of Kophobelemnon species is not clear; in fact, the WoRMS (World Register of Marine Species) reports K. leukartii Ko¨lliker 1872 whereas the ERMS (European Register of Marine species) considers K. leuckarti Cecchini 1917 to be an accepted species. Acknowledgements This study was supported by EU 7FP CoralFISH project (www.eu-fp7-coralfish.net). The authors wish to thank S. Desantis and G. Accogli (Department of Animal Production, Faculty of Veterinary Medicine Bari, Italy) for the SEM photograph, and D. Vafidis, G. Williams and C. Morri for their suggestions. References Bayer F.M., Grasshoff M., Verseveldt J. (1983) Illustrated trilingual glossary of morphological and anatomical terms
Marine Ecology 34 (2013) 313–320 ª 2013 Blackwell Verlag GmbH
A facies of Kophobelemnon in the Mediterranean Sea
applied to Octocorallia. ISBN 90 04 07139 3 printed in the Netherlands by Brill EJ. Brunetti R. (2009) Botryllid species (Tunicata, Ascidiacea) from the Mediterranean coast of Israel, with some considerations on the systematics of Botryllinae. Zootaxa, 2289, 18–32. Budillon G., Lo Bue N., Siena G., Spezie G. (2010) Hydrographic characteristics of the water masses and circulation in the Northern Ionian Sea. Deep-Sea Research II, 57, 441–457. Carpine C. (1970) E´cologie de l’e´tagebathyal dans la Me´diterrane´e. Bulletin Institut Oce´anographique (Monaco), 71, 1–140. Civitarese G., Gacic M., Vetrano A., Boldrin A., Bregant D., Rabitti S., Souvermezoglou E. (1998) Biogeochemical fluxes through the Strait of Otranto (Eastern Mediterranean). Continental Shelf Research, 18, 773–789. Corselli C. (2010) Introduction: cold-water coral communities in the Mediterranean Sea. Deep-Sea Research II, 57, 323–325. De Lazzari A., Boldrin A., Rabitti S., Turchetto M.M. (1999) Variability and downward fluxes of particulate matter in the Otranto Strait area. Journal of Marine Systems, 20, 399–413. D’Onghia G., Mastrototaro F., Matarrese A., Politou C-Y., Mytilineou Ch. (2003) Biodiversity of the upper slope demersal community in the eastern Mediterranean: preliminary comparison between two areas with and without trawl fishing. Journal of Northwest Atlantic Fishery Science, 31, 263–273. D’Onghia G., Maiorano P., Sion L., Giove A., Capezzuto F., Carlucci R., Tursi A. (2010) Effects of the deep-water coral banks on the abundance and size structure of the megafauna in the Mediterranean Sea. Deep-Sea Research II, 57, 397–411. D’Onghia G., Indennidate A., Giove A., Savini A., Capezzuto F., Sion L., Vertino A., Maiorano P. (2011) Distribution and behaviour of deep-sea benthopelagic fauna observed using towed cameras in the Santa Maria di Leuca cold-water coral province. Marine Ecology/Progress Series, 443, 95–110. Etiope G., Savini A., Lo Bue N., Favali P., Corselli C. (2010) Deep-sea survey for the detection of methane at the “Santa Maria di Leuca” cold-water coral mounds (Ionian Sea, South Italy). Deep-Sea Research II, 57, 431–440. Gili J.M., Page`s F. (1987) Pennatula´ceos (Cnidaria, Anthozoa) recolectados en la plataforma continental Catalana (Mediterra´neo Occidental). Miscellania Zoologica, 11, 25–39. Hebbeln D., Wienberg C., Beuck L., Freiwald A., Wintersteller P.; cruise participants (2009) Report and preliminary results of RV Poseidon cruise POS 385 ‘Cold-Water Corals of the Alboran Sea (western Mediterranean Sea)’, May–June 2009, Faro–Toulon. Berichte, Fachbereich Geowissenschaften, Universita¨t Bremen, No. 273, 79 pp (available at http:// www.marum.de/en/Page9183.html). Le Danois E. (1948) Les Profondeurs de la Mer trente ans de recherches sur la faune sous-marine au large des côtes de France. Payot, Paris: 303 pp.
319
A facies of Kophobelemnon in the Mediterranean Sea
Mastrototaro, Maiorano, Vertino, Battista, Indennidate, Savini, Tursi & D’Onghia
Lo´pez-Gonza´lez P.J., Gili J.M., Williams G.C. (2001) New records of Pennatulacea (Anthozoa: Octocorallia) from the Atlantic coast, with description of a new species and a zoogeographic analysis. Scientia Marina, 65, 59–74. Massuti E., Gordon J.D.M., Moranta J., Swan S.C., Stefanescu C., Merrett N.R. (2004) Mediterranean and Atlantic deep-sea fish assemblages: differences in biomass composition and sizerelated structure. Scientia Marina, 68 (Suppl. 3), 101–115. Mastrototaro F., D’Onghia G., Corriero G., Matarrese A., Maiorano P., Panetta P., Gherardi M., Longo C., Rosso A., Sciuto F., Sanfilippo R., Gravili C., Boero F., Taviani M., Tursi A. (2010) Biodiversity of the white coral bank off Cape Santa Maria di Leuca (Mediterranean Sea): an update. Deep-Sea Research II, 57, 412–430. Moen F.E., Svensen E. (2004) Marine fish and invertebrates of Northern Europe. KOM publishing, Kristiansund: 608 pp. Morri C., Bavestrello G., Bianchi C.N. (1991) Faunal and ecological notes on some benthic cnidarians species from the Tuscan Archipelago and Eastern Ligurian Sea (Western Mediterranean). BMIB Bollettino Musei Istituti Biologici Universita di Genova, 54–55, 27–47. Pardo E., Aguilar R., Garcı´a S., de la Torriente A., Ubero J. (2011) Documentacio´n de arrecifes de corales de agua frı´a en el Mediterra´neo occidental (Mar de Albora´n). Chronica naturae, 1, 20–34. Pe´re`s J.M., Picard J. (1964) Nouveau Manuel de Bionomie benthique de la mer Me´diterrane´e. Recueil des Travaux de la Station Marine d’Endoume, 31, 1–137.
320
Rice A.L., Tyler P.A., Paterson G.J.L. (1992) The pennatulid Kophobelemnon stelliferum (Cnidaria: Octocorallia) in the Porcupine seabight (North-est Atlantic Ocean). Journal of the Marine Biological Association of the United Kingdom, 72, 417–434. Sanfilippo R. (2003) Climatic response in the genus Placostegus Philippi, 1844 from PlioPleistocene to recent Mediterranean, with description of a new species (Polychaeta, Serpulidae). Bollettino della Societa` Paleontologica Italiana, 42, 171–178. Sarda` F., Calafat A., Flexas M.M., Tselepides A., Canals M., Espino M., Tursi A. (2004) An introduction to Mediterranean deep-sea biology. Scientia Marina, 68(Suppl. 3), 7–38. Savini A., Corselli C. (2010) High-resolution bathymetry and acoustic geophysical data from Santa Maria di Leuca cold coral province (Northern Ionian Sea-Apulian continental slope). Deep-Sea Research II, 57, 326–344. Vertino A., Savini A., Rosso A., Di Geronimo I., Mastrototaro F., Sanfilippo R., Gay G., Etiope G. (2010) Benthic habitat characterization and distribution from two representative sites of the deep-water SML Coral Mound Province (Mediterranean). Deep-Sea Research II, 57, 380–396. Williams G.C. (1990) The Pennatulacea of southern Africa (Coelenterata, Anthozoa). Annals of the South African Museum, 99, 31–119.
Marine Ecology 34 (2013) 313–320 ª 2013 Blackwell Verlag GmbH
