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Coelosphaera (Coelosphaera) koltuni sp. nov. (Porifera: Demospongiae): a new species from South Orkney Islands, Antarctica

  • Laura SchejterEmail author
  • Javier Cristobo
  • Pilar Ríos
Original Paper
  • 62 Downloads

Abstract

In this work, we describe a new sponge species belonging to the genus Coelosphaera Thomson, 1873. It was collected during an Argentinean Summer Antarctic Research Cruise onboard the Research Vessel “Puerto Deseado” in March 2014, near South Orkney Islands (Antarctica), 60° 53′ S 42° 35′ W, 487-m depth. Coelosphaera (Coelosphaera) koltuni sp. nov. is a globular, fistular and hollow-shaped sponge, with significantly larger megascleres (almost twice the length) than other Antarctic and New Zealand Coelosphaera species. We provide comparative data of all the described Coelosphaera species, especially those from nearby regions.

Keywords

Sponges Poecilosclerida New species description Antarctic waters 

Abbreviations

MACN

Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”

MACN-IN

Invertebrates’ collection of MACN

IEO

Instituto Español de Oceanografía, Spain

INIDEP

Instituto Nacional de Investigación y Desarrollo Pesquero, Argentina

Introduction

The family Coelosphaeridae originally grouped poecilosclerid sponges with fistules, parchment-like surface membranes and hollow bladder–like morphologies. Van Soest (2002) revised Coelosphaeridae to contain species of Myxillina (this suborder has subsequently been abandoned (Morrow and Cardenas 2015)) possessing arcuate chelae, ectosomal smooth tornotes and if present, reticulate choanosomal skeleton. The fistular morphology was considered significant only at the genus level. Van Soest (2002) considered seven genera as valid: Chaetodoryx Topsent, 1927, Coelosphaera Thomson, 1873, Forcepia Carter, 1874 Histodermella Lundbeck, 1910 Inflatella Schmidt, 1875 Lepidosphaera Lévi & Lévi, 1979, Lissodendoryx Topsent, 1892. Subsequently, two other genera were described and added to Coelosphaeridae: Celtodoryx Perez, Perrin, Carteron, Vacelet & Boury-Esnault, 2006 and Myxillodoryx Aguilar-Camacho & Carballo, 2012.

The genus Coelosphaera contains 36 valid species and has a cosmopolitan distribution (van Soest 2002). Subsequently, van Soest et al. (2018) subdivided the genus into two sub-genera based in the possession of echinating acanthostyles (Histodermion) or its absence (Coelosphaera). Following this classification, there are 3 species in Coelosphaera (Histodermion) Topsent, 1927 and 33 in Coelosphaera (Coelosphaera) (van Soest et al. 2018). Only 2 Coelosphaera (Coelosphaera) species have been recorded from Antarctic waters: C. (C.) antarctica Koltun, 1976, and C. (C.) globosa Bergquist, 1961 (Koltun 1976).

Within the Antarctic region, the sponge fauna of the South Orkney Islands is probably one of the least studied areas. Sarà et al. (1992) reviewed the demosponge fauna of this region and listed only 8 species, most of the records were from the Soviet Antarctic Expeditions (in 1955–1958) reported by Koltun (1964). More recently, during Argentinean Antarctic Research Cruises carried out in the austral summers of 2012 and 2014 onboard R/V “Puerto Deseado”, 17 sites in the South Orkney Region were sampled using bottom trawls (see Schejter 2012; Gaitán et al. 2014; Schejter et al. 2016). Sponges were separated from catches and in 4 of the 17 sites they were the main component of the benthic community in terms of relative biomass (Schejter 2012; Gaitán et al. 2014). A preliminary list of the Demospongiae fauna (mainly Poecilosclerida) comprised 32 taxa (Schejter et al. 2017). In the current study, we describe a new species of Coelosphaera collected during these expeditions. In addition, we provide comparative information from all extant species of Coelosphaera.

Material and methods

The specimens were collected during the second stage of the Argentinean Summer Antarctic Cruise (Campaña Antártica de Verano–CAV) onboard the Research Vessel “Puerto Deseado” in March 2014, near South Orkney Islands (Antarctica) (60° 53′ S, 42° 35′ W) at 487-m depth (Fig. 1). Samples were collected with a bottom otter trawl (each trawl was approximately 15 min long). Sponge specimens were sorted from the total catch then photographed and frozen onboard. Live colour and complementary data on the faunal assemblage were noted. Later, in the laboratory, the samples were preserved in a 4% formaldehyde solution or dried for spicule preparations.
Fig. 1

Type locality for Coelosphaera (Coelosphaera) koltuni sp. nov. in South Orkney Islands, Antarctica

In order to study the spicules, the organic matter was digested with a sodium hypochlorite solution, and consecutively washed with water and ethanol according to the known standard procedures (Cristobo et al. 1993; Hajdu et al. 2011). Spicules were measured and examined with a Leica DM 1000 stereomicroscope at the Laboratorio de Bentos (INIDEP), Argentina. The data for spicule sizes are based on about 25–30 measurements for each spicule category, comprising minimum, maximum and average lengths in micrometres (μm). The skeletal architecture was studied by light microscopy using a Nikon Eclipse 50i. Scanning electron microscope (SEM) images were obtained at the Mar del Plata University, Argentina, using a SEM microscope JEOL JSM 6460 LV and at the University of Oviedo, Spain, using a JEOL–6100 SEM.

General classification and the names of class, subclass, order and suborders follow the classification proposed by Morrow and Cardenas (2015) and the World Porifera Database (http://www.marinespecies.org/porifera/).

Results

Systematics

Phylum Porifera Grant, 1836

Class Demospongiae Sollas, 1885

Subclass Heteroscleromorpha Cárdenas, Pérez & Boury-Esnault, 2012

Order Poecilosclerida Topsent, 1928

Family Coelosphaeridae Dendy, 1922

Diagnosis: Fistular-hollow, branching, massive or encrusting sponges. Surface smooth in fistular forms. Non-fistular representatives have their surface often irregularly pitted and punctate, but areolated pore fields are absent. Skeleton reticulate, in fistular forms frequently vestigial. Ectosomal tornotes diactinal (often tylote). Choanosomal megascleres smooth or acanthose styles, occasionally oxeas or strongyles. In fistular forms, choanosomal megascleres may be lost. Next to arcuate isochelae, microscleres include sigmas and raphides (van Soest 2002).

Genus Coelosphaera Thomson, 1873

Synonymy: Coelosphaera Thomson 1873: 484. [Histoderma] Carter 1874: 220 (preocc.). Histioderma Carter, 1886: 452. [Sideroderma] Ridley & Dendy, 1886 348 (preocc.). Siderodermella Dendy, 1922 105. Histodermion Topsent, 1927: 9. Xytopsoocha de Laubenfels, 1936: 55. Naauna de Laubenfels, 1950: 78. Coelosphaericon Bakus, 1966: 33 (van Soest 2002).

Type species: Coelosphaera tubifex Thomson, 1873 represented as Coelosphaera (Coelosphaera) tubifex Thomson, 1873 (type by monotypy) (van Soest et al. 2018).

Diagnosis: Massive, bladder-like, or encrusting, burrowing growth forms with erect fistules; smooth ectosomal tylotes form a compact tangential crust; choanosomal skeleton consists of poorly developed tracts and scattered smooth spicules; megascleres are smooth tylotes or strongyles of one size only, to which acanthostyles may be added; microscleres are arcuate isochelae, sigmas and raphides (often in trichodragmata), some of which may be absent.

Subgenus Coelosphaera (Coelosphaera) Thomson, 1873

Coelosphaera (Coelosphaera) koltuni sp. nov.

(Figures 2, 3, 4, 5)
Fig. 2

Coelosphaera (Coelosphaera) koltuni sp. nov.: a Holotype; bf Paratypes

Fig. 3

Coelosphaera (Coelosphaera) koltuni sp. nov. Spicules SEM images: a Anisostrongyle; b, c Anisostrongyle ends; d Sigmas; e Isochela

Fig. 4

Coelosphaera (Coelosphaera) koltuni sp. nov. Schematic drawing of spicules: a Anisostrongyle; b Sigmas; c Isochela

Fig. 5

Coelosphaera (Coelosphaera) koltuni sp. nov.: ac View of the skeleton by SEM images. Arrow-pointed microscleres

Synonymy: Coelosphaera (Coelosphaera) globosa Koltun, 1976: 178.

Type material

Holotype MACN-IN 41376, CAV 2014, Stage 2, L11, east of South Orkney Islands, 60° 33.0′ S; 42° 35.5′ W, 487 m, collected by L. Schejter using bottom otter trawl (Fig. 2a).

Paratypes: MACN-IN 41377 (4 specimens) and Collection “Porifera” of the Oceanographic Centre of Gijón (IEO), Spain (1 specimen) CAV 2014, Stage 2, L11, east of South Orkney Islands, 60° 33′ S; 42° 35.5′ W, 487 m, collected by L. Schejter using bottom otter trawl (Fig. 2b–f).

Description

External morphology: Globular sponge attached to hard substrata by a short and narrow peduncle that can include small stones (Fig. 2). Surface rough with hollow fistules (oscules) of different lengths and widths, most of them are open with a cribose structure. Size: the holotype is 53 mm high, 34 mm in diameter at equatorial level and 0.5 mm thick. Paratypes are smaller. Measurements were taken after fixation process. Consistency tough, parchment-like. Inside it is hollow. Colour light beige (alive), white in preserved and dried specimens.

Skeleton: Ectosome two or three layers of tangential crisscrossed anisostrongyles without distinctive arrangement (Fig. 3a, b). Scattered microscleres are sometimes evident (Fig. 3c). Reduced choanosome in brushes, brown when dried.

Spicules: Megascleres: anisostrongyles, asymmetrically ended without any swollen end and smooth shaft; 850 (1093.4) 1300 μm by 33–38 μm. Microscleres: arcuate isochelae tridentate: 25 (29.8) 35 μm. Sigmas comprised classical and contorted in S and C shape, slightly hooked in the ends: 35 (46.1) 55 μm (Figs. 4 and 5).

Ecology: A catch of 17.72 kg was obtained in the type locality, and was dominated by large holothurians (Stychopus sp.), fishes and sea stars. Pycnogonids, small crustaceans, mollusks, ascidiaceans, sea urchins, polychaetes and bryozoans were also recorded. Studies on scleractinid corals belonging to the species Caryophyllia (Caryophyllia) antarctica (Marenzeller, 1904), Paraconotrochus antarcticus (Gardiner, 1929), Javania antarctica (Gravier, 1914), Flabellum (Flabellum) flexuosum Cairns, 1982 and F. (F.) impensum Squires, 1962 were carried out on specimens collected in the same haul (Schejter et al. 2016), while sea pens, sea anemones and primnoid corals were also recorded (Gaitán et al. 2014). In addition, 15 sponge taxa were identified (Schejter et al. 2017). The sampling methodology employed did not allow better characterisation of the seabed.

Etymology: Derivatio nominis. Dedicated to the memory of the late Professor Vladimir M. Koltun (1921–2005), prominent Russian spongiologist. In honour of his contribution to the knowledge of Coelosphaera species and the Antarctic sponge fauna.

Remarks: Coelosphaera (C.) koltuni sp. nov. fits with the definition of the Coelopshaera given by van Soest (2002) in terms of its shape and consistency, with leathery hollow fistules and the type of spicules, but in this case there we found no raphides or trichodragmata. The new species differs from most Coelosphaera species in the spiculation. Unlike most species of this genus, it lacks tylotes as megascleres. Instead, there are anisostrongyles. It was assigned to the subgenus Coelosphaera (Coelosphaera) due to the absence of acanthostyles. Only two other Coelosphaera species were previously recorded from Antarctic waters: C. (C.) antarctica Koltun, 1976, and C. (C.) globosa Bergquist, 1961 (Koltun 1976). There are only drawings of the type species of C. (C.) antarctica, originally described as C. appendiculata antarctica, a sub-species of C. appendiculata from the northern hemisphere. C. appendiculata is currently considered to be a junior synonym of C. (C.) tubifex Thomson, 1873 (van Soest et al. 2018). The new species can be distinguished from the later by the size (the new species has bigger megascleres) and shape of spicules, which are tylotes in C (C.) antarctica, whilst in C. (C.) koltuni sp. nov. they are anisostrongyles (Table 1).
Table 1

Comparison of megascleres and microscleres dimensions (in μm) of Antarctic, Australian and New Zealand Coelosphaera (Coelosphaera) species

Species

Anisostrongyles

Tylotes

Isochelae

Sigmas/toxas

Locality

Depth (m)

Reference

C. (C.) koltuni sp. nov.

850–1300

25–35

35–55 sigmas

Antarctica S. Orkney Islands

487

This work

Koltun’s “C. globosa

920–1200

_

33–38

44–55 sigmas

Antarctica (MacRobertson Land?)

529–540

Koltun 1976

C. (C.) antarctica

240–580

28–44

50–60 sigmas

Antarctica

(Enderby Land)

220

Koltun 1976

C. (C.) globosa

650–700

600–725

25–28

25–40

New Zealand

530

Bergquist 1961

C. (C.) transiens

264–350

20–29.5

28–72

New Zealand

0

Bergquist and Fromont 1988

C. (C.) verrucosa

423

21

38

Australia

?

Carter 1886

C. (C.) polymasteides

381

33

Australia

?

Carter 1886

Koltun (1976) described some specimens that he named Coelosphaera globosa, considering them to be very close to the New Zealand species described by Bergquist (1961) from the Chatham Rise, to the east of the South Island of New Zealand. He commented that two specimens of C. globosa were larger than the holotype in size and spiculation. In addition, he also mentioned that the containing jar had 3 different labels from 3 stations in Antarctica and New Zealand. It is possible that Koltun had found this species, but decided to keep the name of the New Zealand species. Koltun’s samples are probably at the Zoological Museum of the Zoological Institute of the Russian Academy of Sciences, Saint Petersburg. However, the material belonging to the BANZAR expedition remained uncataloged, unassembled by species and some remained with Koltun’s identifications in jars. This is a very large collection, several hundred jars, and it is not possible to find the sponges easily (Olga Bozhenova, pers. com.). Although we were not able to examine Koltun’s samples, we considered that Koltun’s description of his specimens could be consistent with the new species herein described, and for that reason we decided to name it in his honour.

The new species can be distinguished from its nearest relatives in New Zealand and Australian waters Coelosphaera (C.) transiens Bergquist and Fromont, 1988, C. (C.) verrucosa (Carter, 1886) and C. (C.) polymasteides (Carter, 1886) (see Table 1). Coelosphaera (C.) transiens has tylotes while the new species has anisostrongyles; the shape is also different, being encrusting in the Australian species and globular in the new one. Also, it is important to highlight here the different ecology of both species, Coelosphaera (C.) transiens living in mid-tidal pools whilst Coelosphaera (C.) koltuni sp. nov. is from deep sea. With reference to C. (C.) verrucosa and C. (C.) polymasteides, the differences are based in the type of megascleres that are also tylotes. Additionally, C. (C.) polymasteides lacks sigmas (Table 1). Megascleres are also different among the new species and C. (C.) globosa; the Australian species has slightly wavy tylotes that can be subtylote or can have a secondary expansion behind the end; strongyles occur as rare modifications of the tylotes (Bergquist and Fromont 1988).

Coelosphaera (C.) koltuni sp. nov. can be also distinguished from the rest of the Coelosphaera species (see Table 2) as follows: In the absence of raphides/trichodragmas the new species is different of C. (C.) barbadensis van Soest, 2017, C. (C.) biclavata (Priest, 1881) (originally as Polymastia), C. (C.) crumena Pulitzer-Finali, 1993, C. (C.) hechteli van Soest, 1984, C. (C.) macrosigma (Topsent, 1890), C. (C.) navicelligera (Ridley, 1885), C. (C.) peltata (Topsent, 1904), C. (C.) physa (Schmidt, 1875), C. (C.) ramosa (Dendy, 1922) and C. (C.) raphidifera (Topsent, 1889) (originally Fibularia raphidifera). In the absence of tylotes as megacleres, the new species is different from most species of subgenus: C. (C.) barbadensis, C. (C.) biclavata, C. (C.) calcifera (Burton, 1934), C. (C.) crumena, C. (C.) crusta Tanita and Hoshino, 1989, C. (C.) dichela (Hentschel, 1912), C. (C.) encrusta (Kumar, 1925), C. (C.) fistula Little, 1963, C. (C.) fucoides (Topsent, 1897), C. (C.) gracilis (Hentschel, 1912), C. (C.) hatchi (Bakus, 1966), C. (C.) hechteli van Soest, 1984, C. (C.) lissodendoryxoides van Soest, 2017, C. (C.) macrosigma, C. (C.) navicelligera, C. (C.) oglalai Lehnert, Stone & Heimler, 2006, C. (C.) pedicellata Lévi, 1993, C. (C.) ramosa, C. (C.) solenoidea (Lévi 1964), C. (C.) tubifex and C. (C.) tunicata (Schmidt, 1870). Coelosphaera (C.) koltuni sp. nov. has sigmas as microscleres and in this character it is different from: C. (C.) crumena, C. (C.) oglalai, C. (C.) peltata, C. (C.) physa and C. (C.) tunicata. In comparison to species having strongyles as megascleres the new species has anisostrongyles, a clear difference with: C. (C.) bullata Lévi, 1993 (also sigmas are thinner), C. (C.) calcifera (this one has also tylotes), C. (C.) chondroida Lévi, 1993 (has similar ends), C. (C.) oglalai, C. (C.) peltata, C. (C.) phlyctenodes (Carter, 1876) (has one of the ends pointed and clear different size of strongyles, isochelae and sigmas), C. (C.) picoensis Topsent, 1928 (that the author defines as substrongylotylotes and they are slightly polytylotes and with one of the end rounded and swollen) and C. (C.) raphidifera with strongyles smaller than 300. Finally, C. (C.) physa has oxeas and not strongyles.
Table 2

Comparison of megascleres and microscleres dimensions (in μm) of all the Coelosphaera (Coelosphaera) species (except Antarctic, Australian and New Zealand species)

Species

Strongyles

Tylotes

Isochelae

Sigmas/toxas

Tricodragmata/raphide

Locality

Depth (m)

Reference

C. (C.) koltuni sp. nov.

850–1300

25–35

35–55

S. Orkney Islands

487

This work

C. (C.) barbadensis

231–384–488

19.9–24.7–28.2

42.4–49.6–61.2

20.0–24.4–40.0

294–431–473 (R)

Barbados

3–5

van Soest 2017

C. (C.) biclavata

508

317.5

12.7–16.9

50.8

254 (T)

Honduras

25–31

Priest 1881

C. (C.) bullata

750–1100

32–38

40–50

New Caledonia

965

Lévi 1993

C. (C.) calcifera

350

350

20.26

28.70

GBR Australia

51

Burton 1934

C. chondroida

1000–1400

500–750

35–45

50–60

 

New Caledonia

650–680

Lévi 1993

C. (C.) crumena

250–280

40–44

250 (R)

Kenya

160

Pulitzer-Finali 1993

C. (C.) crusta

300–350

< 20

25

30–60

Japan

61–87

Tanita and Hoshino 1989

C. (C.) dichela

456–640

288–320

39–43

15–19

17–19

Indonesia

18

Hentschel 1912

C. (C.) encrusta

280–290

210–220

23–26

30–40

India

?

Kumar 1925

C. (C.) fistula

107–211–240

9–10.5–12

25–40.8–53

Florida Gulf

12–14

Little 1963

C. (C.) fucoides

300–500

16–40

50

India

0–10

Topsent 1897

C. (C.) gracilis

304–376

176–224

15–19

12–13

37–40

Indonesia

10–14

Hentschel 1912

C. (C.) hatchi

535–770–970

32–38–42

111–134–162 (T)

California

221

Bakus 1966

C. (C.) hechteli

268–556.7–533

22–26.7–30

38–48.4–60

285–306.5–330 (T)

Curaçao

90

van Soest 1984

C. (C.) lissodendoryxoides

378–413–456

236–260–336

28–32.3–36

16–18.8–21

34–40.4–48

18–29

Guyana Shelf

25–29

van Soest 2017

C. (C.) macrosigma

700

415

30

15–65 (T)

Azores I.

736

Topsent 1890

C. (C.) navicelligera

280–595

20

10

60

19

450 (T)

New Guinea

?

Ridley 1885

C. (C.) oglalai

570–634

325–364

47–52

Alaska

100–155

Lehnert et al. 2006

C. (C.) pedicellata

550–840

340–400

25–30

32–40

New Caledonia

965–1005

Lévi 1993

C. (C.) peltata

500–530

50

130 (T)

Azores

599

Topsent 1904

C. (C.) phlyctenodes

592

50

84

Portugal

684

Carter 1876

C. (C.) physa

800 oxas

70

?

Baffin Bay

309–334

Schmidt 1875

C. (C.) picoensis

630 substrongylotylotes

48–50

69

Azores

736

Topsent 1928

C. (C.) ramosa

480

20

9

40

20

246 (T)

Seychelles

17

Dendy 1922

C. (C.) raphidifera

300

?

 

60

Campeche

?

Topsent 1889

C. (C.) solenoidea

275–325

18

30–35

Acantoxes 200–250

Mozambique

70

Lévi 1964

C. (C.) tubifex

500

900

25–30

?

_

North Atlantic, Arctic

?

Thomson 1873

C. (C.) tunicata

323

69

Florida

?

Schmidt 1870

Discussion

In the study of material of the B.A.N.Z. Antarctic Research Expedition, Koltun (1976) reported two specimens of Coelosphaera, which he identified as C. globosa, a New Zealand species. However, Koltun (1976) noted some differences between his material and that of the holotype. The size of the specimens and spicules was larger in Antarctic specimens, and particularly the megascleres were described as “Strongyla fusiform and mostly asymmetrically ended”. Koltun did not notice that the megascleres of the New Zealand species were tylotes and significantly smaller. The present description of Coelosphaera (Coelosphaera) koltuni sp. nov. clarifies the status of Koltun’s specimens of C. globosa collected almost a century ago. We suggest that C. (C.) globosa is restricted to New Zealand waters, while the only two Coelosphaera species occurring in Antarctic waters are C. (C.) antarctica and the new species herein described.

Sponges have been reported as a dominant group in terms of the biomass of benthic communities in South Orkney (Ramos 1999; Schejter 2012; Gaitán et al. 2014; Brasier et al. 2018) reaching up to 90% of the benthic catches in some sites. This region is part of the Scotia Arc (Thomson 2004), a large area consisting of several archipelagos that could potentially play an important role in the distribution of species between Antarctica and Magellan regions. The present study has the potential to help fill gaps in our understanding of the biogeography of the Southern Ocean and whether it may have been less isolated over geological time than once thought (Arntz et al. 2005). Sponges were also employed as Indicator Taxa to detect Vulnerable Marine Ecosystems (Lockhart and Jones 2008; Jones and Lockhart 2011; Brasier et al. 2018). However, these studies did not provide a species list and only mentioned the group “Porifera” or classes “Demospongiae” and “Hexactinellida”. A quick search through the OBIS database for this marine Ecoregion showed about 300 unpublished records of sponges, most of them not identified to species level, and with many synonyms (R. Downey, pers. comm.). The fact that new species are currently being described in these vulnerable ecosystem highlights its conservation value.

Sará et al. (1992) reviewed the presence of eight species in the South Orkney Islands, while later van Soest et al. (2012) mentioned nine species, emphasising the need to get appropriate data in under sampled regions of the world. The present description of a new sponge species is another step to elucidate species richness and biodiversity of a scarcely studied region in Antarctica. We estimate that the number of sponge species on the South Orkney Islands sponge inventory will be at least doubled after the in-progress studies (i.e. Schejter et al. 2017).

Notes

Acknowledgements

The authors would like to thank Esteban Gaitán and Hugo Merlo for their valuable effort during sampling onboard de RV “Puerto Deseado”, as well as the rest of the crew, and to Mónica Oppedisano (Mar del Plata University) and Alfredo Quintana (Universidad Oviedo) for their technical assistance during SEM image acquisition. In addition, we would we like to acknowledge the kind assistance of Olga Bozhenova for the information pertaining to Koltun’s specimens in the Zoological Institute of the Russian Academy of Sciences, Saint Petersburg. The authors would like to thank Elena Isla and Christine Morrow for the careful revision of English. This is INIDEP Contribution N°2167.

Funding

This study was partially funded by Instituto Nacional de Investigación y Desarrollo Pesquero, Dirección Nacional del Antártico (Argentina) and PICT 2013-0629 to LS. LS is a reseacher at Consejo Nacional de Investigaciones Científicas y Técnicas, Argentina. JC and PR are researchers at Instituto Español de Oceanografía, Spain.

Compliance with ethical standards

Conflict of interest

The authors declare that they have no conflict of interest.

Ethical approval

All applicable international, national and/or institutional guidelines for the care and use of animals were followed by the authors.

Sampling and field studies

All necessary permits for sampling and observational field studies have been obtained for the Madibenthos Survey and the authors from the competent authorities and are mentioned in the Acknowledgements.

Data availability

All data generated or analysed during this study are included in this published article.

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© Senckenberg Gesellschaft für Naturforschung 2019

Authors and Affiliations

  1. 1.Instituto de Investigaciones Marinas y Costeras (IIMyC-CONICET) and Instituto Nacional de Investigación y Desarrollo Pesquero (INIDEP)Mar del PlataArgentina
  2. 2.Centro Oceanográfico de GijónInstituto Español de OceanografíaGijónSpain
  3. 3.Departamento de Zoología y Antropología FísicaUniversidad de AlcaláMadridSpain

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