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Article

Porifera Associated with Deep-Water Stylasterids (Cnidaria, Hydrozoa): New Species and Records from the Ross Sea (Antarctica)

1
Department of Life and Environmental Sciences (DISVA), Polytechnic University of Marche, Via Brecce Bianche, 60131 Ancona, Italy
2
National Biodiversity Future Center (NBFC), Piazza Marina 61, 90133 Palermo, Italy
3
Institute of Polar Sciences, Consiglio Nazionale delle Ricerche (CNR), Via Gobetti 101, 40129 Bologna, Italy
4
Stazione Zoologica Anton Dohrn, Villa Comunale, 80121 Napoli, Italy
5
Department of Geosciences, University of Padova, Via Gradenigo 6, 35131 Padova, Italy
6
Department of Earth, Environmental and Life Sciences (DISTAV), University of Genova, Corso Europa 26, 16132 Genova, Italy
*
Author to whom correspondence should be addressed.
J. Mar. Sci. Eng. 2024, 12(12), 2317; https://doi.org/10.3390/jmse12122317
Submission received: 31 October 2024 / Revised: 6 December 2024 / Accepted: 8 December 2024 / Published: 17 December 2024
(This article belongs to the Section Marine Biology)

Abstract

Stylasterid corals are known to be fundamental habitat-formers in both deep and shallow waters. Their tridimensional structure enhances habitat complexity by creating refuges for a variety of organisms and by acting as basibionts for many other invertebrates, including sponges. Porifera represent crucial components of marine benthic assemblages and, in Antarctica, they often dominate benthic communities. Here, we explore the sponge community associated with thanatocoenosis, mostly composed of dead stylasterid skeletons, collected along the Western and Northern edges of the Ross Sea continental shelf. Overall, 37 sponge species were identified from 278 fragments of the stylasterid Inferiolabiata labiata, of which 7 are first records for the Ross Sea, 1 is first record for Antarctic waters and 2 are proposed as new species. Despite the high biodiversity recorded in this and previous studies on Antarctic deep-sea communities, we are still far from capturing the true richness of Antarctic benthic assemblages. Long-term research programs designed to improve the knowledge of the deep-sea fauna inhabiting Antarctic waters are needed to support successful management and conservation plans, especially in this area, considered one of the main marine diversity hotspots worldwide.

1. Introduction

The family Stylasteridae (Cnidaria, Hydrozoa) represents the second largest group of calcified cnidarians after scleractinians in terms of number of species [1]. In the class Hydrozoa, Stylasteridae is the second most species-rich family distributed worldwide [2], except for the Arctic region, where no species have been described so far [2]. Conversely, stylasterid diversity in the South Ocean is comparatively higher than in other oceans [3] and, in the Antarctic region (southern of the 54° S), 14 genera and 33 species have been documented to date [4,5]. Here, these corals are predominantly found in deep waters, where they constitute an important component of Antarctic deep-water coral communities [2,6]. Despite their high occurrence in Antarctic collections, stylasterids remain a highly understudied group. A comprehensive synthesis of recent records is lacking, many collections remain unidentified beyond the family level, and knowledge of their ecology and distribution is limited, characterized by sporadic and inconsistent sampling across various depths [3].
Like their scleractinian counterparts—e.g., [7,8,9]—many stylasterid corals are considered fundamental habitat-formers contributing to the structuring of deep and shallow water habitats [10,11]. The tridimensional structure of their calcareous skeleton enhances the habitat complexity by creating refuges for a variety of mobile organisms [12] and by acting as basibionts for many other invertebrates, such as annelids, anthozoans, cirripeds, copepods, cyanobacteria, echinoderms, gastropods, hydroids and sponges [12,13,14,15].
Sponges represent crucial components of marine benthic assemblages, and in Antarctica, they often dominate the benthic communities [16]. A recent study documented a total of 397 sponge species across depths ranging from 0 to 7000 m throughout the Antarctic region, encompassing 139 genera within 70 families [16]. This notable diversity is likely related to factors such as geographic isolation, pronounced structural heterogeneity (particularly of the epibiotic communities), and large geographic extension [17,18,19]. These factors also explain the high level of sponge endemism, with 170 out of the 397 sponge species (43%) being endemic [16]. Nonetheless, knowledge about Antarctic deep-sea fauna and sponge diversity of deep-sea communities remains limited [20].
The first report of sponges associated with stylasterids dates back to 1998, when Alectona microspiculata Bavestrello, Calcinai, Cerrano & Sarà, 1998 was observed burrowing into the stem of Distichopora sp. colonies [21]. Subsequently, three sponge species, including one excavating, were found associated with Stylaster sanguineus (Milne Edwards & Haime, 1850) [15]. An unidentified hadromerid sponge was recorded inside the basal portion of a colony of Errina dabneyi (Pourtalès, 1871) [22], and other sponges were observed attached to both dead and living E. dabneyi skeletons [12]. A recent Italian campaign conducted in the Ross Sea (Antarctica) in 2017, using a Remotely Operated Vehicle (ROV), documented extensive areas covered with dead stylasterid colonies and provided initial findings on sponges associated with Antarctic hydrocorals [23]. The authors recorded 38 species, including 2 new to science, associated with 54 dead colonies of the two stylasterid species Errina fissurata Gray, 1872 and Inferiolabiata labiata (Moseley, 1879), revealing a very high diversity of sponges associated with the thanatocoenosis of this coral group.
In this context, the primary objective of the present study is to improve the knowledge of the taxonomy and ecology of sponges associated with the Antarctic stylasterid I. labiata from the Ross Sea. The output of this work not only confirms the great sponge diversity in Antarctic deep waters, but also provides crucial information to support the implementation of additional protection and conservation measures in a key area globally acknowledged as a critical biodiversity hotspot [24,25].

2. Materials and Methods

The samples investigated in the present study were collected in January and February 2017 in the Ross Sea (Antarctica) during the XXXII Antarctic campaign onboard the R/V Italica in the framework of the PNRA research program GRACEFUL (PNRA16_00069). Three sites on the western and northern edge of the Ross Sea continental shelf were explored, including Iselin Bank (IB), Cape Hallett Canyon (CHC), Hallett Ridge (HR), and several unknown stations (US), numbered from 1 to 9 (Table 1; Figure 1).
Samples were collected between 670 and 1022 m depth with an epibenthic sled (width = 150 cm; depth = 120 cm; height = 50 cm), equipped with a two-mesh net of different sizes (a 20 mm square mesh on the front, and a 5 mm on the back) at CHC, IB and HR, and with a 52 L box-corer in US 1–9. Video footage and images of the seabed were acquired with a deep camera attached to a Rosette system. Each video showed a similar seascape, characterized by extensive deposits of skeletons of dead stylasterids lying on a muddy bottom, and by a few living colonies (see Figure 1c in [23]). All collected samples were photographed and allowed to dry. At the end of the campaign, samples were routed to Italy and are now permanently stored at the Italian National Antarctic Museum (MNA, Genova Section, Italy), where they have been cataloged and labeled with the codes GRC “number” + MNA “number”. Part of the material has been previously studied at the University of Genova (see [23]), while the rest has been sent to the Polytechnic University of Marche (UNIVPM) for further analyses. The collected stylasterids have also been examined taxonomically, and an updated morphological description is currently being developed. The present study presents only the results of the sponge species associated with the stylasterid specimens identified as Inferiolabiata labiata.
All I. labiata specimens were observed under a stereomicroscope to detect the presence of sponges, following the method applied by [23]. Often, the limited amount of available material prevented the application of the classical procedure for the preparation of the samples for a Scanning Electron Microscope (SEM) (see [23,26]). In those cases, we adopted and modified the method proposed by [27] for diatoms to obtain clear images. Spicules were transferred onto stubs, coated with platinum or gold, and observed under a Philips XL20 SEM (Eindhoven, The Netherlands) at UNIVPM. For species identification, the length and width of at least 15 spicules per type were measured, and the minimum, mean (with standard deviation), and maximum spicule sizes were reported.
To assess the size, percentage cover and density of sponge specimens (individuals/cm2 of Stylasteridae), the surface of each stylasterid sample, along with its associated epibiontic sponges, was photographed. Due to their roughly fan-shaped growth habitus, both sides of coral specimens were photographed and measured using ImageJ software version 2.0.0 [28]. This methodology was also applied to measure the size of each sponge sample.
For newly described sponge species, their holotypes and associated coral colonies were deposited at the MNA Genova Section.

3. Results

The presence of sponges was investigated on 278 fragments (39.1 ± 48.6 cm2 in area) of Inferiolabiata labiata. Overall, 169 sponge specimens were found, belonging to 2 classes (Demospongiae and Hexactinellida), 7 orders and 19 families, for a total of 37 species. Among the identified species, seven are considered a first record for the Ross Sea, one is a first record for Antarctic waters and two are proposed as new species (Table 2).
The most abundant species, in terms of number of specimens, was Iophon radiatum (n = 47), followed by Clathrochone cf. clathroclada (n = 21) and Hymeniacidon fragilis comb. nov. (n = 20) (Table 2; Figure 2A). Most sponges displayed an encrusting growth habit (74% of species and 78% of specimens, respectively), while a smaller component exhibited a massive erect growth (26% of species and 22% of specimens, respectively) (Table 2; Figure 2B). No boring sponges were found, while, in various fragments, the boring barnacle Australophialus tomlinsoni (Newman & Ross, 1971) was recorded as the only boring organism.
The largest specimen analyzed belonged to the species Iophon unicorne, measuring 3.611 cm2 (Figure 3), while the smallest one was Isodictya setifera, with only 0.003 cm2 (Table 2). Overall, most specimens (131 out of 169) presented small sizes, ranging between 0.003 and 0.5 cm2 (Figure 4A).
The average sponge density on I. labiata fragments was 0.11 ± 0.10 individuals/cm2 (0.01–0.52 individuals/cm2). Among all species, Haliclona (Gellius) rudis had the highest percentage cover, reaching 2.06 ± 0.99%, followed by Haliclona sp.1 (1.57 ± 2.49%), Mycale (Anomycale) cf. titubans (1.53 ± 1.49%), Myxilla (Myxilla) elongata (1.06 ± 1.63%), Myxilla (Myxilla) mollis (1.05 ± 1.71%), C. cf. clathroclada (0.87 ± 1.48%), Hymeniacidon fragilis comb. nov. (0.79 ± 1.21%), Esperiopsis flagellata (0.65 ± 0.68%), I. radiatum (0.59 ± 1.00%) and Amphilectus rugosus (0.56 ± 0.18%) (Figure 4B).

Systematic Section

In this section, only newly described species and new records for Antarctica and/or the Ross Sea, as well as species for which only scant information is currently available, are described.
Order Poecilosclerida Topsent, 1928
Family Coelosphaeridae Dendy, 1922
Lissodendoryx (Lissodendoryx) stylosa sp. nov. Bertolino & Calcinai, 2024
Synonym: L. (L.) complicata (Hansen, 1885) sensu Boury-Esnault & Van Beveren (1982).
Zoobank: urn:lsid:zoobank.org:act:F7A3768A-C14E-42E4-B35B-6780983A6637.
Holotype: MNA 15999 (GRC-02-223 O1); paratypes: MNA 15960 (GRC-02-223 CA2, MNA 15961 (GRC-02-223 CB1).
Locality and material: Iselin Bank, 72°16.1196′ S, 176°36.2814′ W and 72°15.7728′ S, 176°35.5638′ W. Depth 670 m.
Description: The analyzed sponge samples are encrusting with a smooth surface and ochraceous in color. Their size varies from 0.027 to 0.132 cm2 (Figure 5A).
Skeleton: Not observed due to the small material available.
Spicules (holotype): Style I, curved, mainly in the third, upper part (Figure 6A), size 580 (644 ± 54) 710 × 20 μm. Style II, thin, straight or slightly curved (Figure 6B), 300 (364 ± 38) 425 × 10 (11 ± 2) 15 μm. Arcuate isochelae, with both tridentate ends (Figure 6C), 25 (35 ± 9) 42.5 × 15 μm. Sigmas in a single category, C- and S-shaped (Figure 6D), 25 (32 ± 2) 35 × 2.5 (4 ± 1) 5 μm.
Etymology: The new species is named after the presence of styles as dermal spicules.
Remarks: Our specimens fit well, in the size and shape of the spicules, with the specimen described by Boury-Esnault and Van Beveren [29] as Lissodendoryx (Lissodendoryx) complicata (Hansen, 1885) from Kerquelen Islands, except for the presence of a single category of sigma (Table 3). L. (L.) complicata was described in Southern Norway and later reported from the Arctic and several areas in the North Atlantic (Table 3). As a consequence, conspecificity between our specimen and the species described by Hansen [30] is unlikely considering the great geographic distance. The description by Hansen [30] is, in fact, inaccurate, and according to Lundbeck [31], who directly analyzed the samples, Hansen’s illustrations of the sponge spicules probably refer to foreign species. However, L. (L.) complicata differs from the species here described by the presence of smaller dermal styles (Table 3) and tylotes (replaced by styles in the new Antarctic species). Additionally, the smaller category of sigmas present in Hansen’s species is different in shape compared to the ones found in our samples. In fact, Lundbeck [31] (p. 168) reported the following description: “sigmata of the small form; these are rather characteristic; they are highly curved, often almost in a circular manner, the points, however, being generally curved a little more inward; they are plane”.
Lissodendoryx (Lissodendoryx) styloderma Hentschel, 1914
Locality and material: Iselin Bank, 72°16.1196′ S, 176°36.2814′ W and 72°15.7728′ S, 176°35.5638′ W. Depth 670 m. Cape Hallett Canyon, 71°58.8666′ S, 172°11.6298′ E and 71°59.2512′ S, 172°10.6020′ E. Depth 750 m. Two specimens: MNA 15966 (GRC-02-223 BV1), MNA 15967 (GRC-02-223 CX1).
Other material: MNA 13340 (GRC-02-223 (1) sp. 3), MNA 16008 (GRC-02-223 (1) sp. 5) (reported by Costa et al. [23] as Lissodendoryx (Lissodendoryx) cf. styloderma).
Description: The analyzed samples are encrusting with a smooth surface and a typical red-orange color; their size varies between 0.045 and 0.564 cm2 (Figure 5B).
Skeleton: Only the ectosomal skeleton has been observed, and it consists of loose bundles of tornotes and scattered chelae.
Spicules: Fusiform subtylostyles/tylostyles, slightly sinuous and with spined head and pointed tip (Figure 7A,B), 550 (673 ± 83) 810 × 15 (16 ± 2) 20 μm. Tornotes with one end slightly swollen, often mucronate, usually similar to styles; the other end very pointed, gradually tapering (Figure 7C,D), 300 (356 ± 39) 425 × 7.5 (9.5 ± 1) 10 μm. Arcuate isochelae with characteristic folded wings (Figure 7E), 22.5 (31 ± 4) 40.
Remarks: The sizes and shapes of the spicules of our specimens match the original description of the species by Hentschel [33]. The distinct shape of the chelae and the mucronate head of the small styles characterize this rare species, whose only sure record is for the East Antarctic Wilkes Land [33]. In 2013, Göcke and Janussen reported the presence of this species in the Weddel Sea, but their identification is doubtful, due to the presence of sigmas and tornotes in the spicule set [34]. Conversely, our specimens represent the first confirmed record of this species for the Ross Sea.
Family Crellidae Dendy, 1922
Crella (Crella) tubifex (Hentschel, 1914)
Locality and material: Iselin Bank, 72°16.1196′ S, 176°36.2814′ W and 72°15.7728′ S, 176°35.5638′ W. Depth 670 m. Two specimens: MNA 15968 (GRC-02-223 BO1), MNA 15969 (GRC-02-223 BO2).
Description: This sponge is encrusting, with an irregular, microspined surface. The size is between 0.084 cm2 and 0.103 cm2. The color is off-white to grey (Figure 5C).
Skeleton: Not observed.
Spicules: Slightly curved acanthostyles with conical and sharp extremities, and spines along their entire length (Figure 8A), 170 (221 ± 50) 310 × 15 (17 ± 2) 20 μm. Straight anisostrongyles (Figure 8B), 390 (538 ± 78) 660 × 10 (15 ± 5) 20 μm, smooth and curved. Acanthostrongyles are fusiform, slightly and irregularly curved, and with different ends (Figure 8C,D), 340 (385 ± 33) 450 × 20 (22 ± 4) 30 μm; they have spines along their entire shaft but with a higher concentration in the central portion (Figure 8D). No microscleres were observed.
Remarks: The sizes and shapes of the spicules in our specimens match the original description of the species by Hentschel [33]. This is a new record for the Ross Sea.
Family Esperiopsidae Hentschel, 1923
Esperiopsis flagellata sp. nov. Bertolino & Calcinai, 2024
Synonym: Esperiopsis villosa (Carter, 1874) sensu Kirkpatrick, 1908.
Zoobank: urn:lsid:zoobank.org:act:397BC0FC-A2B8-4920-80FB-803DE07DB803.
Holotype: MNA 15962 (GRC-02-223 (8) sp. 6); paratype: MNA 15963 (GRC-02-223 AV1).
Locality and material: Iselin Bank, 72°16.1196′ S, 176°36.2814′ W and 72°15.7728′ S, 176°35.5638′ W. Depth 670 m. Other material: MNA 15964 (GRC-02-223 AO1), MNA 15965 (GRC-02-223 AV)4.
Description: The sponge is encrusting and greyish and ranges from 0.037 to 0.464 cm2 (Figure 5D).
Skeleton: Not observed.
Spicules: Fusiform styles, mycalostyle-like with a slightly constricted neck and a central expansion, often with a mucronate end (Figure 9A,B). Some are straight, others curved and flexuous. Their size is 490 (599 ± 49) 670 × 10 (16 ± 5) 20 μm. Isochelae are divided into two classes: isochelae I with straight shaft (Figure 9C), 33.8 (54.84) 80.6, and isochelae II with arcuate shaft (Figure 9D), 18.2 (21.5) 23.4 μm. Large C-shaped sigmas of large size (Figure 9E) 85 (100 ± 13) 127.5. Some sigmas may remind of ’flagellated’ sigmas, but as their end have the same length and there is no strong asymmetry in the endings, they cannot be defined as such (Figure 9F).
Etymology: This new species is named after the presence of sigmas reminiscent of flagellated sigma.
Remarks: This new species here described is very similar to Esperiopsis villosa Carter, 1874 in the shape of the spicules (see page 660 Figure 3A,B in [35]) and in size. Carter [36] described the species from the Faroe Islands and later collected it in the North Atlantic Ocean and Arctic area [37]. Kirkpatrick recorded this species in the Ross Sea and discussed the affinities with Carter’s species, concluding that “the only difference worthy of mention that I can find between the Northern and Antarctic specimens is the absence of the placocheles or isochelae palmatae with broad shafts. I can only discover two kinds of isochelae palmate” (see page 35 in [38]).
Our samples fit well with that of Kirkpatrick [38], especially in the description of styles both in size and shape (671 × 18 μm; larger palmate isochelae 43 μm, smaller isochelae, 18 μm); considering the great geographical separation, and the absence of a third category of isochelae, we assume that the conspecificity between our specimen and the species described by Carter [36] is unlikely, leading us to consider our samples as a new species, previously erroneously considered by Kirkpatrick [38] as E. villosa.
Family Microcionidae Carter, 1875
Subfamily Ophlitaspongiinae
Artemisina plumosa Hentschel, 1914
Locality and material: Iselin Bank, 72°16.1196′ S, 176°36.2814′ W and 72°15.7728′ S, 176°35.5638′ W. Depth 670 m. Hallett Ridge, 72°23.0340′ S, 176°06.1020′ E and 72°23.3868′ S, 176°06.2094′ E. Depth 910 m. Five specimens: MNA 15989 (GRC-02-223 AN2), MNA 15990 (GRC-02-223 Q1), MNA 15991 (GRC-02-223 BC3), MNA 15992 (GRC-02-223 BC4), MNA 15993 (GRC-02-223 CV1).
Description: The sponge is massive, slightly bushy and characterized by a bristly surface. The color is light grey. The sponge varied in size from 0.031 cm2 to 0.188 cm2 (Figure 5E).
Skeleton: Not observed.
Spicules: Style I, large, mainly straight or slightly curved styles (Figure 10A), 600 (1471 ± 501) 2600 × 20 (25 ± 4) 30 μm. Style II, small, strongly curved (Figure 10B), 380 (476 ± 50) 530 × 15 (16 ± 2) 20 μm. Tylotes with round, slightly spined heads (Figure 10C,D), 260 (372 ± 56) 480 × 5 (6 ± 2) 10 μm. Isochelae (Figure 10E), 7.5 (13 ± 2) 15 μm. Toxas are characterized by a very evident, angular central curve. The size and relative angle of curves vary greatly among these spicules, which are also, but not always, characterized by spined and slightly curved ends (Figure 10F), 90 (216 ± 95) 600 μm.
Remarks: The megascleres of the present samples showed a wider size range than that reported by Hentschel [33] in his original description. According to the author, the large styles were up to 1232 μm long, while the small ones were up to 456 μm long. The size of the isochelae is similar, while the toxas in our samples are longer than those measured by Hentschel [33] (96–144 μm). This is a new record for the Ross Sea.
Family Mycalidae Lundbeck, 1905
Mycale (Anomomycale) cf. titubans (Schmidt, 1870)
Locality and material: Iselin Bank, 72°16.1196′ S, 176°36.2814′ W and 72°15.7728′ S, 176°35.5638′ W. Depth 670 m. Four specimens: MNA 15994 (GRC-02-223 AH1), MNA 15995 (GRC-02-223 AU4), MNA 15996 (GRC-08-065 CM1), MNA 15997 (GRC-02-223 (2) sp. 1), MNA 15998 (GRC-02-223 (2) sp. 4).
Description: The sponge is encrusting, has a size ranging from 0.070 to 1.615 cm2, and displays a coloration from beige to greyish (Figure 5F).
Skeleton: Not observed.
Spicules: Large slightly curved and fusiform mycalostyles (Figure 11A) of 600 (852 ± 62) 940 × 20 (25 ± 5) 30 μm. Thinner, straight mycalostyles (Figure 11B) of 380 (441 ± 21) 480 × 10 μm. Curved and contorted anomochelae (Figure 11C) with basal ala of the foot expanded and characterized by a serrated margin, 27.5 (44 ± 7) 50 μm. Highly curved C-shaped sigmas (Figure 11D), 65 (80 ± 5) 142 × 2.5 μm.
Remarks: The characteristics of our specimens match those of Mycale (Anomomycale) titubans (Schmidt, 1870) recorded in the North Atlantic Ocean, with no evident differences found in the spicule set (Table 4). Nevertheless, the conspecificity between our specimens and the species described by Schmidt [39] is unlikely considering the great geographic distance of the collected samples. Boury-Esnault and Van Beveren [29] reported the presence of M. (A.) titubans with very large sigmas and anomochelae in the Kerguelen Archipelagos (Antarctic Ocean) (Table 4), but these characters are not shared by our specimens and the ones collected in the north hemisphere. As a consequence, Boury-Esnault and Van Beveren’s sample is a probable new species. New information for morphological and molecular studies could help to clarify the differences among the Antarctic, Kerguelen and North Atlantic specimens and to demonstrate the presence of two new species of Mycale (Anomomycale) in the Antarctic area.
Order Suberitida Chombard & Boury-Esnault, 1999
Family Suberitidae Schmidt, 1870
Hymeniacidon fragilis Koltun, 1964 comb. nov.
Synonym: Plicatellopsis fragilis Koltuna, 1964
Locality and material: Iselin Bank. 72°16.1196′ S, 176°36.2814′ W and 72°15.7728′ S, 176°35.5638′ W. Depth 670 m. Eighteen specimens: MNA 16003 (GRC-02-223 BZ2), MNA 15970 (GRC-02-223 A4), MNA 15971 (GRC-02-223 A5), MNA 15972 (GRC-02-223 C1), MNA 15973 (GRC-02-223 C2), MNA 15974 (GRC-02-223 C3), MNA 15975 (GRC-02-223 E3), MNA 15976 (GRC-02-223 O2), MNA 15977 (GRC-02-223 S3), MNA 15978 (GRC-02-223 AL), MNA 15979 (GRC-08-065 CM3), MNA 15980 (GRC-02-091 CU10, MNA 15981 (GRC-02-133 CT3), MNA 15982 (GRC-02-223 BB5), MNA 15983 (GRC-02-223 BB1), MNA 15984 (GRC-02-223 BF1), MNA 15985 (GRC-02-093 CS1), MNA 15986 (GRC-02-093 CS5).
Other material examined: MNA 13377 (GRC-02-223 (22) sp. 1) (Figure 3B), MNA 16004 (GRC-02-223 (37) D) reported by Costa et al. [23] as Plicatellopsis cf. fragilis (Figure 12A,B), and MNA 15987 (GRC-02-223 (17) sp. 1), MNA 15988 (GRC-02-223 (15) sp. 1) reported by Costa et al. [23] as Clathria (Clathria) paucispicula.
Description: Some specimens are massively encrusting (Figure 12A) or globular; the color is cream and ochre, with sizes of about 0.010 cm2 to 6.75 cm2. Other specimens are laminar in shape, and their size is up to 3.4 × 2 cm large and 0.5 cm thick (Figure 12B). Dry specimens are friable.
Skeleton: Specimens show a choanosomal skeleton of bundles of styles I and II running towards the sponge surface (Figure 12C,D), making it slightly hispid. The ascending bundles are connected by transversal bundles and free spicules, creating an irregular network (Figure 12C,D). Style II styles are also scattered among the mashes. The ectosomal skeleton consists of a thin layer of tangential, intercrossing bundles and single style II styles (Figure 12E).
Spicules: Style I, large, thick, fusiform and slightly curved (Figure 12F), 640 (677 ± 34) 754 × 40 (45 ± 5) 50 μm. Style II, fusiform, slightly curved (sometimes sinuous) and thinner (Figure 12F), 193 (303 ± 14) 377 × 10 μm.
Remarks: The spicular set, shape and consistency (lamellar and particularly friable) of the examined specimens fit those of Plicatellopsis fragilis described by Koltun (see page 85 and Figure 22 in [42]; Table 5). The only difference is in the thickness of the style I styles, which is slightly thinner according to Koltun (i.e., 20–30 μm; [42]). However, the skeleton organization is typical of the genus Hymeniacidon (see page 807 in [43]); in fact, Koltun [42] described the ectosomal skeleton as “composed chiefly of tangential styles”. Therefore, we suggest transferring the species Plicatellopsis fragilis to the genus Hymeniacidon.
Order Tetractinellida Marshall, 1876
Family Tetillidae Sollas, 1886
Tetilla coronida Sollas, 1888
Locality and material: 71°38.4132′ S and 172°09.3048′ E. Depth 1022 m. One specimen: MNA 15999 (GRC-TR17-007 CP1).
Description: The sponge of about 0.3 cm2 is white and with a hispid surface due to the protruding long spicules (Figure 5G).
Skeleton: Not observed.
Spicules: Fusiform and very sharply pointed oxeas (Figure 13A), 610 (716 ± 46) 790 × 20 (24 ± 5) 30 μm. Very long protriaenes, with a thin rhabdome (Figure 13B), 1600 (2082 ± 383) 2800 × 10 μm, clads are 40 (100 ± 48.5) 175 μm. Anatriaenes (Figure 13C) with rhabdome of 3000 (3830 ± 279) 4300 × 15 (20 ± 4) 27.5 μm and clads 90 (108 ± 14.7) 130 × 10 (10.8 ± 2) 15 μm. Anamonaenes as modified protriaenes where two of the clads are lacking, and the remaining is curved to form a semicircle (Figure 13D), 210 (280 ± 38) 330 × 20 (23 ± 3) 25 μm. Sigmaspires, small C- or S-shaped with an irregular, rough surface (Figure 13E), 15 (20 ± 4) 27.5 μm.
Remarks: The spicule size reported by Sollas [44] in his original description is larger considering the length of the protiaenes (3.37 mm) and of the anatriaenes (7.14 mm). In our specimens, however, most of the spicules were broken, making them potentially much longer than observed. Nevertheless, the spicule set of our sample matches perfectly with that reported by Sollas [44]. The species is known from Heard Islands and later from Kerguelens [29], but this is the first report for the Antarctic continent.
Family Vulcanellidae Cárdenas, Xavier, Reveillaud, Schander & Rapp, 2011
Poecillastra antarctica Koltun, 1964 comb. nov.
Synonym: Poecillastra compressa antarctica Koltun, 1964.
Locality and material: Iselin Bank, 72°16.1196′ S, 176°36.2814′ W and 72°15.7728′ S, 176°35.5638′ W. Depth 670 m. Three specimens: MNA 16000 (GRC-02-223 N3), MNA 16001 (GRC-02-223 L3), MNA 16002 (GRC-02-223 AQ2). Other material: MNA 13302 (GRC-02-223 (26) sp.1) (reported by Costa et al. [23] as Poecillastra compressa antarctica).
Description: The sponge is massive, brown, with a stiff consistency; it ranges from 0.015 to 0.315 cm2 (Figure 5H).
Skeleton: Not observed.
Spicules: Fusiform oxeas with pointed extremities, sometimes presenting one blunt ending (Figure 14A), 730 (1260 ± 436) 2480 × 10 (22 ± 8) 35 μm. Calthrops and orthotrianes with very short rhabdomes and with clads bent in the same direction (Figure 14B), measuring 560 (909 ± 79) 1100 × 30 (39 ± 3) 40 μm. Plesiasters with two to five microspined rays (Figure 14C), 35 (56 ± 15) 90 × 5 (8 ± 3) 15 μm. Amphiasters with three to four thin rays (Figure 14D) and spirasters with thicker rays (Figure 14E), 10 (16 ± 3) 25 × 3 (4 ± 1) 5 μm. Microxeas absent.
Remarks: For the Southern Ocean, two species of the genus Poecillastra are currently known: P. compressa parvistellata Topsent, 1913 and P. compressa antarctica Koltun, 1964. The shape and size of the spicules in the analyzed sponge samples differ from P. compressa parvistellata, lacking microxeas and plesiasters, while spirasters are smaller (10–13 μm; pages 611–613 in [45]). Conversely, they match the description of the species by Koltun [42] for P. compressa antarctica, except for the oxeas, which are longer in Koltun’s species (3.2–3.7 mm in length). The author already proposed to erect this as a new subspecies of P. compressa (Bowerbank, 1866) due to its “larger size of tetractines, the different habitus and other minor features” (page 19 in [42]). In particular, Koltun [42] does not comment on the absence of the microxeas, which are also absent in our samples. Therefore, considering these differences, which have been also observed in our samples, and the disjunct geographic areas, the subspecies P. compressa antarctica should be considered as a new species, P. antarctica comb. nov. Koltun, 1964.

4. Discussion

In 2010, Post and colleagues reported the discovery of field-like aggregations of Stylasterid corals and a high abundance of associated fauna in eastern Antarctica, pointing out the high diversity of the Antarctic benthos [46]. Other aggregations of Stylasterids are patchily distributed throughout Antarctica, sub-Antarctica, Patagonia and New Zealand [3,47]. These regions are all identified as priority conservation areas, for the protection and preservation of their benthic biodiversity. These field-like aggregations support a highly diverse associated fauna—e.g., [46,48,49]—and provide crucial ecosystem services. They serve as secondary substrates for other organisms and offer critical habitats for feeding, spawning, nursery and refuge, potentially hosting economically important fish populations [3]. The role of the tridimensional structure created by these calcareous hydrozoans is confirmed by our result: a rich and diverse sponge fauna was, in fact, recorded in association with stylasterid beds in deep Antarctic waters, in line with the known levels of Antarctic diversity [16,23].
In the present study, 37 species associated with the stylasterid Inferiolabiata labiata were identified, representing about 10% of the sponge biodiversity of the entire Antarctic continent. Our results align with those of Costa et al. [23], who reported a rich and diversified sponge fauna of 37 sponge species associated with the stylasterids Errina fissurata and I. labiata, of which 16 were shared between the two studies. This count includes the revision of various samples from Costa et al. [23], specifically the ones erroneously named (i) Lyssodendoryx nobilis and L. cf. nobilis actually belonging to the species Myxilla (Ectyomyxilla) hentscheli Burton, 1929; (ii) Clathria (Clathria) paucispicula (Burton, 1932) and Plicatellopsis cf. fragilis Koltun, 1964, actually Hymeniacidon fragilis (Koltun, 1964) comb. nov.; and (iii) the ones identified as Clathria sp., instead being Clathria (Clathria) toxipraedita Topsent, 1913, a species previously known only for the Antarctic peninsula [37]. With the new records from this study, the number of sponge species associated with the two stylasterid corals rises to 58.
In line with Costa et al. [23], our results also confirmed that the most common associated species was Iophon radiatum, with a total of 47 specimens (28%). As a point of fact, the species of the genus Iophon seem to be very common epibionts also on other Antarctic taxa, such as the bivalves Adamussium colbecki (E. A. Smith, 1902) and the echinoid Ctenocidaris perrieri Koehler, 1912, which host the species Iophon unicorne Topsent, 1907 and I. radiatum, respectively [50], as well as the ophiuroids Ophiurolepis spp., found encrusted by I. radiatum [51]. The diversity of the Antarctic deep-sea fauna, especially regarding sponges, is still scarce even though the number of Antarctic species and endemism is expected to increase in the near future [20]. The high number of new and already known species recorded in this and earlier studies [16,23] highlights both the remarkable diversity of this often-overlooked group and the need to preserve three-dimensional reef-like aggregations. Such structures can enhance habitat complexity, even when created by clusters of dead corals. In deep environments, generally characterized by a scarcity of primary rocky substrates, coral beds represent fundamental secondary hard substrates available for benthic species, especially for those taxa presenting high phenotypic plasticity (e.g., sponges, other cnidarians, bryozoans) that allows them to adapt to any potentially available niche [52,53]. This phenotypic plasticity is particularly enhanced in the observed sponge samples, which were able to survive inside these large clusters of dead stylasterids thanks to their very small sizes, in most cases not exceeding 0.5 cm2.
According to Gutt and Schickan [51], filter-feeding epibionts generally prefer living on elevated substrates, and the large abundance of sponge species identified on I. labiata fragments is particularly interesting when compared to the results obtained by Cerrano et al. [50]: on average, we identified a total of 2 species per fragment of I. labiata, with a maximum of 8 species on a single fragment, in line with the estimations of Cerrano et al. [50], who recorded a maximum of 10 sponge species on a single A. colbecki scallop. However, apart from a few alive colonies, our stylasterid samples were neither elevated nor living substrates, but the framework of dead coral skeletons could have helped the creation of local microcurrents supporting the colonization and survival of sponges [54,55].
No boring sponges have been detected despite availability of dead calcareous material available, used by other excavating organisms such as the boring barnacle Australophialus tomlinsoni. It is known that cnidarians calcareous skeletons are typical substrates for boring sponge species—i.e., [56,57,58,59]; however, it has been observed that stylasterids are probably not suitable substrates for boring sponges [15]. Additionally, the absence of excavating sponges in our samples is not surprising considering that no boring sponge species have been ever found in the Antarctic continent so far, and macroborers are in general scarce in polar environments [60].
In conclusion, studies on other invertebrate groups show that Antarctic deep-sea communities can be as rich, or even richer, than those found in shallower environments [61]. Our results fully align with this pattern, yet they also suggest that we are still far from capturing the true richness of Antarctic benthic assemblages. There is, therefore, an urgent need for long-term research programs aimed at collecting enough samples to enhance our understanding of the deep-sea fauna inhabiting Antarctic waters. This fundamental knowledge of biodiversity is crucial for the development of effective management and conservation plans, especially in regions like Antarctica, considered one of the main marine diversity hotspots in the world.

Author Contributions

Conceptualization, B.C. and M.B.; resources, P.M., C.M. and S.C.; investigation, C.V., B.C., M.B. and S.P.; formal analysis, C.V., C.R. and T.M.; writing—original draft preparation, C.V., B.C., M.B. and C.R.; writing—review and editing, all authors; visualization, C.V., C.R., B.C. and M.B. All authors have read and agreed to the published version of the manuscript.

Funding

The project PNRA-GRACEFUL (Grant No. PNRA16_00069) contributed to the study by providing the stylasterid samples, video footage and still photographs of the seafloor of the Ross Sea. The study was also partially funded under the National Recovery and Resilience Plan (NRRP), Mission 4, Component 2, Investment 1.4—Call for tender No. 3138 of 16 December 2021, rectified by Decree n.3175 of 18 December 2021, of the Italian Ministry of University and Research funded by the European Union—NextGenerationEU; Award Number: Project code CN_00000033, Concession Decree No. 1034 of 17 June 2022 adopted by the Italian Ministry of University and Research, CUP D33C22000960007 and D31B21008270007, Project title “National Biodiversity Future Center—NBFC”.

Data Availability Statement

All relevant data are presented within the manuscript.

Conflicts of Interest

The authors declare no conflicts of interest.

References

  1. Cairns, S.D. Species Richness of Recent Scleractinia. Atoll Res. Bull. 1999, 459, 1–46. [Google Scholar] [CrossRef]
  2. Cairns, S.D. Global Diversity of the Stylasteridae (Cnidaria: Hydrozoa: Athecatae). PLoS ONE 2011, 6, e21670. [Google Scholar] [CrossRef]
  3. Bax, N.; Cairns, S. Chapter 5.7. Stylasteridae (Cnidaria: Hydrozoa). In Biogeographic Atlas of the Southern Ocean; De Broyer, C., Koubbi, P., Griffiths, H.J., Raymond, B., Udekem d’Acoz, C., Van de Putte, A.P., Danis, B., David, B., Grant, S., Gutt, J., et al., Eds.; Scientific Committee on Antarctic Research: Cambridge, UK, 2014; pp. 107–112. [Google Scholar]
  4. Cairns, S.D. Antarctic and Subantarctic Stylasterina (Coelenterata: Hydrozoa). In Biology of the Antarctic Seas XIII; American Geophysical Union (AGU): Washington, DC, USA, 1983; pp. 61–164. ISBN 978-1-118-66672-2. [Google Scholar]
  5. Cairns, S.D. New Records of Stylasteridae (Hydrozoa: Hydroida) from the Galapagos and Cocos Islands. Proc. Biol. Soc. Wash. 1991, 104, 209–228. [Google Scholar]
  6. Heifetz, J. Coral in Alaska: Distribution, Abundance, and Species Associations. Hydrobiologia 2002, 471, 19–28. [Google Scholar] [CrossRef]
  7. Patton, W.K. Distribution and Ecology of Animals Associated with Branching Corals (Acropora spp.) from the Great Barrier Reef, Australia. Bull. Mar. Sci. 1994, 55, 193–211. [Google Scholar]
  8. Stella, J.S.; Pratchett, M.S.; Jones, P.A.H.G.P. Coral-Associated Invertebrates: Diversity, Ecological Importance and Vulnerability to Disturbance. In Oceanography and Marine Biology; CRC Press: Boca Raton, FL, USA, 2011; ISBN 978-0-429-10992-8. [Google Scholar]
  9. Hoeksema, B.W.; Van der Meij, S.E.T.; Fransen, C.H.J.M. The Mushroom Coral as a Habitat. J. Mar. Biol. Assoc. U. K. 2012, 92, 647–663. [Google Scholar] [CrossRef]
  10. Roberts, J.M.; Wheeler, A.J.; Freiwald, A. Reefs of the Deep: The Biology and Geology of Cold-Water Coral Ecosystems. Science 2006, 312, 543–547. [Google Scholar] [CrossRef] [PubMed]
  11. Häussermann, V.; Försterra, G. Extraordinary Abundance of Hydrocorals (Cnidaria, Hydrozoa, Stylasteridae) in Shallow Water of the Patagonian Fjord Region. Polar Biol. 2007, 30, 487–492. [Google Scholar] [CrossRef]
  12. Braga-Henriques, A.; Carreiro-Silva, M.; Porteiro, F.M.; de Matos, V.; Sampaio, Í.; Ocaña, O.; Ávila, S.P. The Association between a Deep-Sea Gastropod Pedicularia Sicula (Caenogastropoda: Pediculariidae) and Its Coral Host Errina dabneyi (Hydrozoa: Stylasteridae) in the Azores. ICES J. Mar. Sci. 2011, 68, 399–407. [Google Scholar] [CrossRef]
  13. Zibrowius, H. Associations of Hydrocorallia Stylasterina with Gall-Inhabiting Copepoda Siphonostomatoidea from the South-West Pacific. Part I. On the Stylasterine Hosts, Including Two New Species, Stylaster papuensis and Crypthelia cryptotrema. Bijdr. Tot Dierkd. 1981, 51, 268–281. [Google Scholar]
  14. Goud, J.; Hoeksema, B. Pedicularia Vanderlandi Spec. Nov., a Symbiotic Snail (Caenogastropoda: Ovulidae) on the Hydrocoral Distichopora Vervoorti Cairns & Hoeksema, 1998 (Hydrozoa: Stylasteridae), from Bali, Indonesia. Zool. Verh. 2001, 334, 77–97. [Google Scholar]
  15. Pica, D.; Bertolino, M.; Calcinai, B.; Puce, S.; Bavestrello, G. Boring and Cryptic Sponges in Stylasterids (Cnidaria: Hydrozoa). Ital. J. Zool. 2012, 79, 266–272. [Google Scholar] [CrossRef]
  16. Downey, R.V.; Griffiths, H.J.; Linse, K.; Janussen, D. Diversity and Distribution Patterns in High Southern Latitude Sponges. PLoS ONE 2012, 7, e41672. [Google Scholar] [CrossRef] [PubMed]
  17. Brey, T.; Klages, M.; Dahm, C.; Gorny, M.; Gutt, J.; Hain, S.; Stiller, M.; Arntz, W.E.; Wägele, J.W.; Zimmermann, A. Antarctic Benthic Diversity. Nature 1994, 368, 297. [Google Scholar] [CrossRef]
  18. Gray, J.S. Antarctic Marine Benthic Biodiversity in a World-Wide Latitudinal Context. Polar Biol. 2001, 24, 633–641. [Google Scholar] [CrossRef]
  19. Starmans, A.; Gutt, J. Mega-Epibenthic Diversity: A Polar Comparison. Mar. Ecol. Prog. Ser. 2002, 225, 45–52. [Google Scholar] [CrossRef]
  20. Janussen, D.; Tendal, O.S. Diversity and Distribution of Porifera in the Bathyal and Abyssal Weddell Sea and Adjacent Areas. Deep Sea Res. Part II Top. Stud. Oceanogr. 2007, 54, 1864–1875. [Google Scholar] [CrossRef]
  21. Bavestrello, G.; Calcinai, B.; Cerrano, C.; Sarà, M. Alectona Species From North-Western Pacific (Demospongiae: Clionidae). J. Mar. Biol. Assoc. U. K. 1998, 78, 59–73. [Google Scholar] [CrossRef]
  22. Wisshak, M.; Correa, M.L.; Zibrowius, H.; Jakobsen, J.; Freiwald, A. Skeletal Reorganisation Affects Geochemical Signals, Exemplified in the Stylasterid Hydrocoral Errina dabneyi (Azores Archipelago). Mar. Ecol. Prog. Ser. 2009, 397, 197–208. [Google Scholar] [CrossRef]
  23. Costa, G.; Bavestrello, G.; Canese, S.; Canessa, M.; Mazzoli, C.; Montagna, P.; Puce, S.; Schiaparelli, S.; Bertolino, M. Sponges Associated with Stylasterid Thanatocoenosis (Cnidaria, Hydrozoa) from the Deep Ross Sea (Southern Ocean). Polar Biol. 2022, 45, 703–718. [Google Scholar] [CrossRef]
  24. Boothroyd, A.; Adams, V.; Alexander, K.; Hill, N. Priority Areas for Marine Protection in the Amundsen and Bellingshausen Seas, Antarctica. Mar. Policy 2024, 167, 106232. [Google Scholar] [CrossRef]
  25. Nissen, C.; Lovenduski, N.S.; Brooks, C.M.; Hoppema, M.; Timmermann, R.; Hauck, J. Severe 21st-Century Ocean Acidification in Antarctic Marine Protected Areas. Nat. Commun. 2024, 15, 259. [Google Scholar] [CrossRef]
  26. Núñez-Pons, L.; Mazzella, V.; Rispo, F.; Efremova, J.; Calcinai, B. DNA Barcoding Procedures for Taxonomical and Phylogenetic Studies in Marine Animals: Porifera as a Case Study. In Marine Genomics: Methods and Protocols; Verde, C., Giordano, D., Eds.; Springer: New York, NY, USA, 2022; pp. 195–223. ISBN 978-1-07-162313-8. [Google Scholar]
  27. Stosch, H.V. Pleurax, Seine Synthese Und Seine Verwendung Zur Einbettung Und Darstellung Det Zellwande von Diatomeen, Peridineen Und Anderen Algen, Sowie Für Eine Neue Methode Zur Electivfarbung von Dinoflagellaten-Penzern. Arch. Für Protistenkd. 1974, 116, 132–141. [Google Scholar]
  28. Rasband, W.S. ImageJ 2011. Available online: https://imagej.net/ij/ (accessed on 7 December 2024).
  29. Boury-Esnault, N.; Van Beveren, M. Les Démosponges Du Plateau Continental de Kerguellen-Heard. Com. Natl. Français Des Rech. Antarct. 1982, 52, 1–175. [Google Scholar]
  30. Hansen, G.A. Spongiadae. The Norwegian North-Atlantic Expedition 1876–1878. Zoology 1885, 13, 1–26. [Google Scholar]
  31. Lundbeck, W. Porifera. (Part II.) Desmacidonidae. Dan. Ingolf-Exped. 1905, 6, 1–219. [Google Scholar]
  32. Tompkins, G.; Baker, E.; Anstey, L.; Walkusz, W.; Siferd, T.; Kenchington, E. Sponges from the 2010-2014 Paamiut Multispecies Trawl Surveys, Eastern Arctic and Subarctic: Class Demospongiae, Subclass Heteroscleromorpha, Order Poecilosclerida, Family Coelosphaeridae, Genera Forcepia and Lissodendoryx; Canadian Technical Report of Fisheries and Aquatic Sciences; Dartmouth: Hanover, NH, USA, 2017; p. 129. [Google Scholar]
  33. Hentschel, E. Monaxone Kieselschwämme Und Hornschwämme Der Deutschen Südpolar-Expedition 1901–1903. Dtsch. Südpolar-Exped. 1914, 15, 35–141. [Google Scholar]
  34. Göcke, C.; Janussen, D. Demospongiae of ANT XXIV/2 (SYSTCO I) Expedition—Antarctic Eastern Weddell Sea. Zootaxa 2013, 3692, 28–101. [Google Scholar] [CrossRef]
  35. Van Soest, R.W.M.; Hajdu, E. Family Esperiopsidae Hentschel, 1923. In Systema Porifera: A Guide to the Classification of Sponges; Hooper, J.N.A., Van Soest, R.W.M., Willenz, P., Eds.; Springer: Boston, MA, USA, 2002; pp. 656–664. ISBN 978-1-4615-0747-5. [Google Scholar]
  36. Carter, H.J. XXIX.—Descriptions and Figures of Deep-Sea Sponges and Their Spicules from the Atlantic Ocean, Dredged up on Board H.M.S. ‘Porcupine,’ Chiefly in 1869; with Figures and Descriptions of Some Remarkable Spicules from the Agulhas Shoal and Colon, Panama. Ann. Mag. Nat. Hist. 1874, 14, 207–221. [Google Scholar] [CrossRef]
  37. de Voogd, N.; Alvarez, B.; Boury-Esnault, N.; Cárdenas, P.; Díaz, M.-C.; Dohrmann, M.; Downey, R.; Goodwin, C.; Hajdu, E.; Hooper, J.; et al. World Porifera Database. Available online: https://www.marinespecies.org/porifera (accessed on 23 January 2024).
  38. Kirkpatrick, R. Porifera (Sponges). II. Tetraxonida, Dendy. National Antarctic Expedition, 1901–1904 Natural History. Zoology 1908, 4, 1–56. [Google Scholar]
  39. Schmidt, O. Grundzüge Einer Spongien-Fauna Des Atlantischen Gebietes. Wilhelm Engelmann Leipz. 1870, iii–iv, 1–88. [Google Scholar]
  40. Topsent, E. Révision Des Mycales de l’Europe Occidentale. Ann. L’institut Océanographique 1924, 1, 77–118. [Google Scholar]
  41. Dinn, C.; Leys, S.P. Field Guide to Sponges of the Eastern Canadian Arctic; Department of Biological Sciences, University of Alberta: Edmonton, AB, USA, 2018. [Google Scholar]
  42. Koltun, V.M. Sponges of the Antarctic. 1 Tetraxonida and Cornacuspongida. In Biological Reports of the Soviet Antarctic Expedition (1955–1958); Pavlovskii, E.P., Andriyashev, A.P., Ushakov, P.V., Eds.; Akademya Nauk SSSR: Moscow, Russia, 1964. [Google Scholar]
  43. Erpenbeck, D.; Van Soest, R.W.M. Family Halichondriidae Gray, 1867. In Systema Porifera. A Guide to the Classification of Sponges; Hooper, J.N.A., Van Soest, R.W.M., Eds.; Springer: New York, NY, USA, 2002; pp. 787–816. ISBN 0-306-47260-0. [Google Scholar]
  44. Sollas, W.J. Report on the Tetractinellida Collected by H.M.S. Challenger, during the Years 1873–1876. Report on the Scientific Results of the Voyage of H.M.S. Challenger during the Years 1873–1876. Zoology 1888, 25, 1–458. [Google Scholar]
  45. Topsent, E. Spongiaires de l’Expédition Antarctique Nationale Ecossaise. Earth Environ. Sci. Trans. R. Soc. Edinb. 1913, 49, 579–643. [Google Scholar] [CrossRef]
  46. Post, A.L.; O’Brien, P.E.; Beaman, R.J.; Riddle, M.J.; Santis, L.D. Physical Controls on Deep Water Coral Communities on the George V Land Slope, East Antarctica. Antarct. Sci. 2010, 22, 371–378. [Google Scholar] [CrossRef]
  47. J. Miller, K.; Mundy, C.N.; Lindsay Chadderton, W. Ecological and Genetic Evidence of the Vulnerability of Shallow-Water Populations of the Stylasterid Hydrocoral Errina Novaezelandiae in New Zealand’s Fiords. Aquat. Conserv. Mar. Freshw. Ecosyst. 2004, 14, 75–94. [Google Scholar] [CrossRef]
  48. Waller, R.G.; Robinson, L.F. Southern Ocean Corals: Cabo de Hornos. Coral Reefs 2012, 31, 205. [Google Scholar] [CrossRef]
  49. Kaiser, S.; Brandão, S.N.; Brix, S.; Barnes, D.K.A.; Bowden, D.A.; Ingels, J.; Leese, F.; Schiaparelli, S.; Arango, C.P.; Badhe, R.; et al. Patterns, Processes and Vulnerability of Southern Ocean Benthos: A Decadal Leap in Knowledge and Understanding. Mar. Biol. 2013, 160, 2295–2317. [Google Scholar] [CrossRef]
  50. Cerrano, C.; Bertolino, M.; Valisano, L.; Bavestrello, G.; Calcinai, B. Epibiotic Demosponges on the Antarctic Scallop Adamussium Colbecki (Smith, 1902) and the Cidaroid Urchins Ctenocidaris perrieri Koehler, 1912 in the Nearshore Habitats of the Victoria Land, Ross Sea, Antarctica. Polar Biol. 2009, 32, 1067–1076. [Google Scholar] [CrossRef]
  51. Gutt, J.; Schickan, T. Epibiotic Relationships in the Antarctic Benthos. Antarct. Sci. 1998, 10, 398–405. [Google Scholar] [CrossRef]
  52. Sánchez, J.A.; Aguilar, C.; Dorado, D.; Manrique, N. Phenotypic Plasticity and Morphological Integration in a Marine Modular Invertebrate. BMC Evol. Biol. 2007, 7, 122. [Google Scholar] [CrossRef]
  53. Guardiola, M.; Frotscher, J.; Uriz, M.-J. High Genetic Diversity, Phenotypic Plasticity, and Invasive Potential of a Recently Introduced Calcareous Sponge, Fast Spreading across the Atlanto-Mediterranean Basin. Mar. Biol. 2016, 163, 123. [Google Scholar] [CrossRef]
  54. Wolfe, K.; Kenyon, T.M.; Mumby, P.J. The Biology and Ecology of Coral Rubble and Implications for the Future of Coral Reefs. Coral Reefs 2021, 40, 1769–1806. [Google Scholar] [CrossRef]
  55. Kenyon, T.M.; Doropoulos, C.; Wolfe, K.; Webb, G.E.; Dove, S.; Harris, D.; Mumby, P.J. Coral Rubble Dynamics in the Anthropocene and Implications for Reef Recovery. Limnol. Oceanogr. 2023, 68, 110–147. [Google Scholar] [CrossRef]
  56. Tunnicliffe, V. Breakage and Propagation of the Stony Coral Acropora cervicornis. Proc. Natl. Acad. Sci. USA 1981, 78, 2427–2431. [Google Scholar] [CrossRef]
  57. Pica, D.; Calcinai, B.; Anderson, M. Excavating sponges boring into the precious red coral from Cape Verde Archipelago. Biol. Mar. Mediterr. 2011, 18, 278–279. [Google Scholar]
  58. Weinstein, D.K.; Maher, R.L.; Correa, A.M.S. Bioerosion. In Mesophotic Coral Ecosystems; Loya, Y., Puglise, K.A., Bridge, T.C.L., Eds.; Springer International Publishing: Cham, Switzerland, 2019; pp. 829–847. ISBN 978-3-319-92735-0. [Google Scholar]
  59. Pulido Mantas, T.; Bavestrello, G.; Bertolino, M.; Cerrano, C.; Pica, D.; Roveta, C.; Calcinai, B. A 3D Innovative Approach Supporting the Description of Boring Sponges of the Precious Red Coral Corallium rubrum. J. Mar. Sci. Eng. 2022, 10, 868. [Google Scholar] [CrossRef]
  60. Meyer, N.; Wisshak, M.; Freiwald, A. Bioerosion Ichnodiversity in Barnacles from the Ross Sea, Antarctica. Polar Biol 2021, 44, 667–682. [Google Scholar] [CrossRef]
  61. Brandt, A.; De Broyer, C.; Gooday, A.; Hilbig, B.; Thomson, M. Introduction to ANDEEP (ANtarctic Benthic DEEP-Sea Biodiversity: Colonization History and Recent Community Patterns)—A Tribute to Howard L. Sanders. Deep Sea Res. Part II Top. Stud. Oceanogr. 2004, 51, 1457–1465. [Google Scholar] [CrossRef]
Figure 1. Map of the location of the sampling stations in the Ross Sea continental shelf.
Figure 1. Map of the location of the sampling stations in the Ross Sea continental shelf.
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Figure 2. (A) Bar plots representing the total number of specimens for sponge species with more than 2 samples. (B) Donut charts showing the percentage of sponge species and specimens with an encrusting (Ec) or massive erect (ME) habit, or both (ME/Ec).
Figure 2. (A) Bar plots representing the total number of specimens for sponge species with more than 2 samples. (B) Donut charts showing the percentage of sponge species and specimens with an encrusting (Ec) or massive erect (ME) habit, or both (ME/Ec).
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Figure 3. Some sponge specimens with larger sizes: (A) Acanthascus (Rhabdocalyptus) australis (MNA 16005, GCR-02-223 D), (B) Iophon unicorne (MNA 16007, GRC-08-023 DC), (C) Haliclona (Gellius) rudis (MNA 16006, GRC-08-023 DE).
Figure 3. Some sponge specimens with larger sizes: (A) Acanthascus (Rhabdocalyptus) australis (MNA 16005, GCR-02-223 D), (B) Iophon unicorne (MNA 16007, GRC-08-023 DC), (C) Haliclona (Gellius) rudis (MNA 16006, GRC-08-023 DE).
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Figure 4. Bar plots showing (A) the total number of specimens in relation to the area covered on the stylasterid Inferiolabiata labiata and (B) the sponge species with the highest percentage cover on I. labiata (expressed as average ± standard deviation).
Figure 4. Bar plots showing (A) the total number of specimens in relation to the area covered on the stylasterid Inferiolabiata labiata and (B) the sponge species with the highest percentage cover on I. labiata (expressed as average ± standard deviation).
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Figure 5. (A) Lissodendoryx (Lissodendoryx) stylosa sp. nov. (holotype MNA 15959, GRC-02-223 O1); (B) L. (L.) styloderma (MNA 13340, GRC-02-223 (1) sp. 3); (C) Crella (Crella) tubifex (MNA 15968, GRC-02-223 BO1); (D) Esperiopsis flagellata sp. nov. (holotype MNA 15962, GRC-02-223 (8) sp. 6); (E) Artemisina plumosa (MNA 15989, GRC-02-223 AN2); (F) Mycale (Anomomycale) cf. titubans (MNA 15994, GRC-02-223 AH1); (G) Tetilla coronida (MNA 15999, GRC-TR17-007 CP1); (H) Poecillastra antarctica comb. nov. (MNA 13302, GRC-02-223 (26) sp. 1). White arrows indicate the position of the sponge specimens on Inferiolabiata labiata fragments.
Figure 5. (A) Lissodendoryx (Lissodendoryx) stylosa sp. nov. (holotype MNA 15959, GRC-02-223 O1); (B) L. (L.) styloderma (MNA 13340, GRC-02-223 (1) sp. 3); (C) Crella (Crella) tubifex (MNA 15968, GRC-02-223 BO1); (D) Esperiopsis flagellata sp. nov. (holotype MNA 15962, GRC-02-223 (8) sp. 6); (E) Artemisina plumosa (MNA 15989, GRC-02-223 AN2); (F) Mycale (Anomomycale) cf. titubans (MNA 15994, GRC-02-223 AH1); (G) Tetilla coronida (MNA 15999, GRC-TR17-007 CP1); (H) Poecillastra antarctica comb. nov. (MNA 13302, GRC-02-223 (26) sp. 1). White arrows indicate the position of the sponge specimens on Inferiolabiata labiata fragments.
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Figure 6. SEM pictures of Lissodendoryx (Lissodendoryx) stylosa sp. nov.: (A) style I; (B) style II; (C) arcuate isochelae; (D) sigmas.
Figure 6. SEM pictures of Lissodendoryx (Lissodendoryx) stylosa sp. nov.: (A) style I; (B) style II; (C) arcuate isochelae; (D) sigmas.
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Figure 7. SEM pictures of Lissodendoryx (Lissodendoryx) styloderma: (A) subtylostyle; (B) magnification of the head and the pointed tip of a subtylostyle; (C) tornotes; (D) magnification of a pointed end of a tornote; (E) arcuate isochelae.
Figure 7. SEM pictures of Lissodendoryx (Lissodendoryx) styloderma: (A) subtylostyle; (B) magnification of the head and the pointed tip of a subtylostyle; (C) tornotes; (D) magnification of a pointed end of a tornote; (E) arcuate isochelae.
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Figure 8. SEM pictures of Crella (Crella) tubifex: (A) acanthostyle; (B) anisostrongyle; (C) acanthostrongyle; (D) magnification of the central portion and extremities of an acanthostrongyle.
Figure 8. SEM pictures of Crella (Crella) tubifex: (A) acanthostyle; (B) anisostrongyle; (C) acanthostrongyle; (D) magnification of the central portion and extremities of an acanthostrongyle.
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Figure 9. Optical microscope pictures of Esperiopsis flagellata sp. nov.: (A) style; (B) end of a style; (C) isochelae I; (D) isochelae II; (E) C-shaped sigma; (F) flagellated sigma.
Figure 9. Optical microscope pictures of Esperiopsis flagellata sp. nov.: (A) style; (B) end of a style; (C) isochelae I; (D) isochelae II; (E) C-shaped sigma; (F) flagellated sigma.
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Figure 10. SEM pictures of Artemisina plumosa: (A) style I; (B) style II; (C) tylote; (D) magnification of the spined head of a tylote; (E) isochelae; (F) toxas.
Figure 10. SEM pictures of Artemisina plumosa: (A) style I; (B) style II; (C) tylote; (D) magnification of the spined head of a tylote; (E) isochelae; (F) toxas.
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Figure 11. SEM pictures of Mycale (Anomomycale) cf. titubans: (A) large mycalostyle; (B) thin mycalostyle; (C) anomochelae; (D) sigmas.
Figure 11. SEM pictures of Mycale (Anomomycale) cf. titubans: (A) large mycalostyle; (B) thin mycalostyle; (C) anomochelae; (D) sigmas.
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Figure 12. Hymeniacidon fragilis (Koltun, 1964) comb. nov.: (A) massively encrusting specimen MNA 13377 (GRC-02-223 (22) sp. 1); (B) laminar specimen MNA 16004 (GRC-02-223 (37) D); (C,D) choanosomal skeleton; (E) ectosomal skeleton; (F) spicules. White arrows indicate the position of the sponge attachment to the stylasterid.
Figure 12. Hymeniacidon fragilis (Koltun, 1964) comb. nov.: (A) massively encrusting specimen MNA 13377 (GRC-02-223 (22) sp. 1); (B) laminar specimen MNA 16004 (GRC-02-223 (37) D); (C,D) choanosomal skeleton; (E) ectosomal skeleton; (F) spicules. White arrows indicate the position of the sponge attachment to the stylasterid.
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Figure 13. SEM pictures of Tetilla coronida: (A) oxeas; (B) extremity of a protriaene; (C) extremities of anatrienes; (D) extremities of anamonaenes; (E) sigmaspires.
Figure 13. SEM pictures of Tetilla coronida: (A) oxeas; (B) extremity of a protriaene; (C) extremities of anatrienes; (D) extremities of anamonaenes; (E) sigmaspires.
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Figure 14. Optical microscope pictures of Poecillastra antarctica: (A) oxeas; (B) calthrops and orthotrianes; (C) plesiasters; (D) amphiasters; (E) spirasters.
Figure 14. Optical microscope pictures of Poecillastra antarctica: (A) oxeas; (B) calthrops and orthotrianes; (C) plesiasters; (D) amphiasters; (E) spirasters.
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Table 1. List of the sampling stations with the corresponding codes, latitude and longitude.
Table 1. List of the sampling stations with the corresponding codes, latitude and longitude.
StationStation CodeLatitudeLongitude
Cape Hallett CanyonCHC−71.98111172.19383
−71.98752172.1767
Iselin BankIB−72.26866−176.60469
−72.26288−176.59273
Hallett RidgeHR−72.3839176.1017
−72.38978176.10349
Unknown station 1US1−71.64022172.15508
Unknown station 2US2−72.49782174.99664
Unknown station 3US3−72.49855174.99802
Unknown station 4US4−72.49892175.00003
Unknown station 5US5−72.4995174.99637
Unknown station 6US6−72.49971174.96788
Unknown station 7US7−72.494803174.94212
Unknown station 8US8−72.4956174.94267
Unknown station 9US9−72.487563174.94933
Table 2. List of the sponge species associated with Inferiolabiata labiata, with growth habits, sponge size, locality and number of specimens. Ec: encrusting; ME: massive erect. IB: Iselin Bank; CHC: Cape Hallett Canyon; HR: Hallett Ridge; US: unknown station. ° new species; * first record for Antarctica; ^ new record for the Ross Sea.
Table 2. List of the sponge species associated with Inferiolabiata labiata, with growth habits, sponge size, locality and number of specimens. Ec: encrusting; ME: massive erect. IB: Iselin Bank; CHC: Cape Hallett Canyon; HR: Hallett Ridge; US: unknown station. ° new species; * first record for Antarctica; ^ new record for the Ross Sea.
ClassSponge SpeciesGrowth
Habit
Sponge Size (Min–Max)LocalityNumber of Specimens
Demospongiae Sollas, 1885Biemna chilensis Thiele, 1905ME0.027–1.407 cm2IB2
Haliclona cf. (Flagellia) flagellifera
(Ridley & Dendy, 1886)
ME0.482 cm2IB1
Haliclona (Gellius) rudis (Topsent, 1901)Ec0.447–1.079 cm2IB3
CHC
Haliclona cf. virens (Topsent, 1908)Ec0.085–0.596 cm2IB5
Haliclona sp. 1 Ec0.350–0.665 cm2IB3
HR
Haliclona sp. 2 Ec0.115 cm2IB1
Iophon abnormalis Ridley & Dendy, 1886 ^Ec0.061–0.234 cm2IB2
Iophon radiatum Topsent, 1901Ec0.006–1.692 cm2IB47
CHC
HR
Iophon unicorne Topsent, 1907Ec0.017–3.611 cm2IB2
CHC
Asbestopluma (Asbestopluma) sinuosa
Costa & Bertolino, 2022
ME0.01 cm2IB1
Inflatella belli (Kirkpatrick, 1907)ME0.034 cm2IB1
Lissodendoryx (Lissodendoryx) stylosa sp. nov. °Ec0.027–0.132 cm2IB3
Lissodendoryx (Lissodendoryx) styloderma
Hentschel, 1914 ^
Ec0.045–0.564 cm2IB2
CHC
Lissondendoryx (Ectyodoryx) nobilis
(Ridley & Dendy, 1886)
Ec0.105 cm2IB1
Lissondendoryx sp.Ec1.483 cm2IB1
Crella (Crella) tubifex (Hentschel, 1914) ^Ec0.084–0.103 cm2IB2
Amphilectus rugosus (Thiele, 1905) ^Ec0.037–0.178 cm2IB3
Esperiopsis flagellata sp. nov. °Ec0.037–0.464 cm2IB4
Isodictya cf. spinigera (Kirkpatrick, 1907)Ec0.003 cm2IB2
Clathria (Clathria) pauper Brøndsted, 1927Ec0.078 cm2IB1
Clathria (Microciona) sp. 1Ec0.193 cm2HR1
Clathria (Clathria) toxipraedita Topsent, 1913Ec0.086–0.125 cm2IB2
HR
Artemisina plumosa Hentschel, 1914 ^ME0.031–0.188 cm2IB5
HR
Mycale (Anomomycale) cf. titubans (Schmidt, 1870)Ec0.07–1.615 cm2IB5
CHC
Myxilla (Myxilla) elongata Topsent, 1916Ec0.005–0.440 cm2IB3
CHC
Myxilla (Myxilla) mollis Ridley & Dendy, 1886Ec0.179–0.757 cm2IB7
CHC
HR
Tedania (Tedaniopsis) oxeata Topsent, 1916Ec1.463 cm2IB1
Tedania (Tedaniopsis) tantula (Kirkpatrick, 1907)Ec0.004–0.192 cm2IB2
Polymastia invaginata Kirkpatrick, 1907Ec0.039–0.279 cm2IB6
Hymeniacidon fragilis (Koltun, 1964) ^ comb. nov.ME/Ec0.01–1.794 cm2IB20
CHC
Pseudosuberites cf. sulcatus (Thiele, 1905)Ec0.072 cm2IB1
Halichondria (Halichondria) sp.Ec0.212 cm2HR1
Halichondria (Halichondria) prostrata Thiele, 1905ME0.400 cm2CHC1
Poecillastra antarctica Koltun, 1964 ^ comb. nov.ME0.015–0.315 cm2IB3
Tetilla coronida Sollas, 1888 *ME0.300 cm2US1
Hexactinellida Schmidt, 1870Clathrochone cf. clathroclada (Lévi & Lévi, 1982)ME0.008–3.563 cm2IB21
HR
Acanthascus (Rhabdocalyptus) australis
(Topsent, 1901)
ME0.109–0.234 cm2IB2
Tot species: 37 Tot specimens: 169
Table 3. Localities of sampling, spicular characteristics and authors of the records of Lissodendoryx (Lissodendoryx) complicata (Hansen, 1885) in the Arctic, North Atlantic and Antarctica.
Table 3. Localities of sampling, spicular characteristics and authors of the records of Lissodendoryx (Lissodendoryx) complicata (Hansen, 1885) in the Arctic, North Atlantic and Antarctica.
LocalityStylesDermal SpiculesIsochelaeSigma ISigma IIAuthor
North Atlantic420–680 × 16–25strongyles with transition to tylotes: 220–400 × 3.5–740–58 × 5–1242–55 × 217–23 × 1[31]
Eastern Arctic and Subarctic448.4–690.3 × 9.3–25.5tylotes: 208.6–359.9 × 3.7–8.527.8–68.9 × 5–13.231.4–60.3 × 1.5–3.915–23.2 × 1.9–3[32]
Kerguelen Archipelago557–787 × 19–26styles: 262–314 × 6.445–58 × 10–1351–64 × 219–26 × 1[29]
Antarctic (Ross Sea)580–880 × 20–30styles II: 290–425 × 10–1525–42.5 × 1525–40 × 5Present work
Table 4. Localities of sampling, spicules characteristics and authors of the records of Mycale (Anomomycale) titubans in the North Atlantic and Antarctic areas.
Table 4. Localities of sampling, spicules characteristics and authors of the records of Mycale (Anomomycale) titubans in the North Atlantic and Antarctic areas.
LocalityMycalostyle IMycalostyle IIAnomochelaeSigmaAuthor
Florida303–250[39]
West coast of Brittany875 × 27430 × 821–3550–130 × 2–5[40]
Denmark590–900 × 15–19320–400 × 5–824–5250–140 × 1.8–5.7[31]
Labrador Sea484–646 × 16–24350–595 × 7.5–1626–3055–97[41]
Kerguelen Archipelago704–960 × 22.4–25.6432.4–580.4 × 6.4–8.370.4–83.2 × 3.2–6.4Sigma I: 93.4–119 × 1.9–5.7[29]
Sigma II: 190.7–262.4 × 5.7–9.6
Antarctica (Ross Sea)600–940 × 20–30380–480 × 1027.5–5065–142 × 2.5Present work
Table 5. Comparison of shape, skeleton organization and spicule dimensions of Plicatellopsis fragilis Koltun, 1964 and Hymeniacidon fragilis (Koltun, 1964) comb. nov.
Table 5. Comparison of shape, skeleton organization and spicule dimensions of Plicatellopsis fragilis Koltun, 1964 and Hymeniacidon fragilis (Koltun, 1964) comb. nov.
SpeciesShapeEctosomal SkeletonChoanosomal SkeletonStyle I (μm)Style II (μm)Author
Plicatellopsis fragilis Koltun, 1964lamellartangential auxiliary styles irregular reticulum of poorly developed longitudinal fiber bundles and isolated spicules550–800 × 20–30250–380 × 6–11[42]
Hymeniacidon fragilis (Koltun, 1964) comb. nov.massively encrusting, lamellartangential auxiliary stylesascending bundles are connected by transversal bundles and free spicules, creating an irregular network; smaller styles are scattered between fibers640–754 × 40–50193–377 × 10Present work
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Calcinai, B.; Marrocco, T.; Roveta, C.; Puce, S.; Montagna, P.; Mazzoli, C.; Canese, S.; Vultaggio, C.; Bertolino, M. Porifera Associated with Deep-Water Stylasterids (Cnidaria, Hydrozoa): New Species and Records from the Ross Sea (Antarctica). J. Mar. Sci. Eng. 2024, 12, 2317. https://doi.org/10.3390/jmse12122317

AMA Style

Calcinai B, Marrocco T, Roveta C, Puce S, Montagna P, Mazzoli C, Canese S, Vultaggio C, Bertolino M. Porifera Associated with Deep-Water Stylasterids (Cnidaria, Hydrozoa): New Species and Records from the Ross Sea (Antarctica). Journal of Marine Science and Engineering. 2024; 12(12):2317. https://doi.org/10.3390/jmse12122317

Chicago/Turabian Style

Calcinai, Barbara, Teo Marrocco, Camilla Roveta, Stefania Puce, Paolo Montagna, Claudio Mazzoli, Simonepietro Canese, Carlo Vultaggio, and Marco Bertolino. 2024. "Porifera Associated with Deep-Water Stylasterids (Cnidaria, Hydrozoa): New Species and Records from the Ross Sea (Antarctica)" Journal of Marine Science and Engineering 12, no. 12: 2317. https://doi.org/10.3390/jmse12122317

APA Style

Calcinai, B., Marrocco, T., Roveta, C., Puce, S., Montagna, P., Mazzoli, C., Canese, S., Vultaggio, C., & Bertolino, M. (2024). Porifera Associated with Deep-Water Stylasterids (Cnidaria, Hydrozoa): New Species and Records from the Ross Sea (Antarctica). Journal of Marine Science and Engineering, 12(12), 2317. https://doi.org/10.3390/jmse12122317

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