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A new species of Alertigorgia (Coelenterata: Octocorallia:
Anthothelidae) from the Indo Malayan region
L.P. van Ofwegen & P. Alderslade
Ofwegen, L.P. van & P. Alderslade. A new species of Alertigorgia (Coelenterata: Octocorallia: Anthothe-
lidae) from the Indo Malayan region.
Zool. Med. Leiden 81 (13), 8.vi.2007: 241-249, figs 1-6.— ISSN 0024-0672.
Leen P. van Ofwegen, Nationaal Natuurhistorisch Museum, P.O. Box 9517, 2300 RA Leiden, The Nether-
lands (e-mail: ofwegen@naturalis.nl).
Phil Alderslade, Museum and Art Gallery of the Northern Territory, P.O. Box 4646, Darwin, Northern
Territory 0801, Australia (e-mail: Phil.Alderslade@nt.gov.au). (Current address: P.O. Box 211, Margate,
Tasmania 7054, Australia).
Key words: Coelenterata; Anthozoa; Octocorallia; Alertigorgia; new species; Ovulidae; Oceanapia; Indo-
nesia.
A new species of Alertigorgia from Indonesian waters is described and depicted, and compared with the
hitherto only described species in the genus, A. orientalis (Ridley, 1884). The species is part of a unique
association in which a mollusc species lives on an octocoral, which on its turn lives on a sponge species.
Introduction
Kükenthal (1908) established the genus Alertigorgia for a single species described by
Ridley (1884) as Iciligorgia orientalis from Torres Strait, Australia. Since that original de-
scription the species has been found regularly in the Malay Archipelago and Australia
(for references see Verseveldt, 1940: 104), but the genus remained monotypic. However,
Colin & Arneson (1995: 89) published an underwater picture of a species named Soleno-
caulon sp., from Papua New Guinea, and a few years later, Erhardt & Baensch (1998:
271) showed a similar looking species labelled as “new genus and new species”, from
Bunaken, Sulawesi, Indonesia. The second author got involved with the identification
of these images, and was already in possession of colony fragments collected by the
Coral Reef Research Foundation and Harry Erhardt when he discussed the matter with
the first author, who also had seen images of this species, made by B.W. Hoeksema
(NNM) in Bunaken national park. During a fieldtrip in 1998, B.W. Hoeksema & L.P. van
Ofwegen photographed and collected several specimens at Bunaken and from this ma-
terial it became obvious we were dealing with an as yet undescribed species of Alertig-
orgia, which is described below. In 2001 the species was also found at Bali, Indonesia
(see material examined).
Titanideum mjöbergi Broch, 1916, should most likely also be included in Alertigorgia.
The species is described as having cylindrical branches and may represent a third spe-
cies of the genus. The original description lacks sufficient detail, and the validity of the
species will be the subject of another paper.
Abbreviations
The NNM-LIPI-WWF Expedition, 2001, was a joint project of the National Museum
of Natural History (NNM), Leiden, The Netherlands, the Research Centre for Oceanol-
ogy, Indonesian Institute of Sciences (PPO-LIPI), and the World Wildlife Fund (WWF),
242 van Ofwegen & Alderslade. A new species of Alertigorgia. Zool. Med. Leiden 81 (2007)
The material is deposited in the Zoological Museum Bogor (ZMB) of the LIPI Research
Centre for Biology (PPB-LIPI, Cibinong), and the NNM (RMNH); NTM stands for Mu-
seum and Art Gallery of the Northern Territory, Australia.
Description
Genus Alertigorgia Kükenthal, 1908
Diagnosis, amended: Anthothelidae with bushy or sparsely branched colonies.
Branches may be cylindrical, flattened with acute edges or expanded and leaf-like. Pol-
yps fully retractile into slit-like apertures, densely scattered over both faces in the spe-
cies with leaf-like branches, arranged bilaterally in two narrow grooves on edges of
conspicuously flattened branches, or singly or in small groups within short narrow
grooves in the species with cylindrical branches. The medulla has prickly needles and
thick rods with a few large tubercles; the surface layer has plump spindles with com-
plex tubercles. Polyps have a collaret and points arrangement. Sclerites are colourless.
Colonies are zooxanthellate.
Alertigorgia hoeksemai spec. nov.
(figs 1-6)
Material examined.— NTM C015585, holotype, BUN.07, Indonesia, N. Sulawesi, Bunaken Park, ESE of
Siladen island, 124°48’E 01°37’30”N, scuba diving, 35-40 m depth, 7.v.1998, coll. B.W. Hoeksema & L.P. van
Ofwegen; paratypes: NTM C015586, same data as holotype; NTM C014257, Indonesia, North Sulawesi
Sea, Bunaken Islands, 01°38’N 124°46’E, 12-18 m depth, 1996, coll. H. Erhardt; NTM C014258, Indonesia,
Manado, 01°45.1’N 124°58.87’E, 3 m depth , coll. Coral Reef Research Foundation, 21.v.1993; RMNH Coel.
31952, same data as holotype; RMNH Coel. 31953, NNM-LIPI-WWF Exped., BAL.09, Indonesia, Bali, Sa-
nur, Loloan Batu Agung, 08°43’31’’S 115°15’57’’E, slowly declining reef slope, sandy base, scuba-diving, 17
m depth, 3.iv.2001, coll. B.W. Hoeksema; RMNH Coel. 31954, NNM-LIPI-WWF Exped., BAL.14, Indone-
sia, Bali, NE-side Pulau Serangan, off lighthouse, 08°43’17’’S 115°15’31’’E, slowly declining reef slope,
sandy base, scuba-diving, 15-20 m depth, 5.iv.2001, coll. B.W. Hoeksema; RMNH Coel. 31955, NNM-LIPI-
WWF Exped., BAL.15, Indonesia, Bali, NE-side Pulau Serangan; 08°44’03’’S 115°15’05’’E, slowly declining
reef slope, sandy base, scuba-diving, 22 m depth, 7.iv.2001, coll. L.P. van Ofwegen & M. Slierings; MZB
Coel. 3, NNM-LIPI-WWF Exped., BAL.13, Indonesia, Bali, NE-side Pulau Serangan, Loloan Serangan,
08°43’39’’S 115°15’13’’E, slowly declining reef slope, sandy base, scuba-diving to 17 m depth, 5.iv.2001, coll.
M.I.Y.T. Hermanlimianto; MZB. Coel. 4, same data as RMNH Coel. 31955, coll. M.I.Y.T. Hermanlimianto.
Material used for comparison.— RMNH Coel. 31956, Alertigorgia orientalis, BUN.16, Indonesia, N. Su-
lawesi, S Tetapaan island, reef, 124°30’30”E 01°18’N, scuba diving -23 m. 12.v.1998, coll. B.W. Hoeksema
& L.P. van Ofwegen.
Description.— The holotype is 70 cm long and 40 cm wide (fig. 1). The lower part is
attached to a sediment dwelling species of the sponge genus Oceanapia (see base of
colony in fig 2c; Colin & Arneson, fig. 167) but it is difficult to see where exactly the
sponge ends and the octocoral begins. All branches are expanded leaf-like. The polyps
are densely scattered over both faces of the leaves. The calyces are dome-shaped and all
polyps are retracted into slit-like apertures (fig. 2f).
The polyps have a collaret of 2-3 rows of spindles and points with 4-6 spindles. All
these spindles have simple tubercles and are up to 0.25 mm long (fig. 3a-b). The tenta-
cles have small rods, up to 0.15 mm long (fig. 3c).
van Ofwegen & Alderslade. A new species of Alertigorgia. Zool. Med. Leiden 81 (2007) 243
The surface layer of the leaves has ovals, cylinders, and spindles, up to 0.30 mm
long, with densely placed complex tubercles (figs 3d, 4a); a few sclerites have some
warts fused disk-like (fig. 4a, second sclerite). Towards the interior the spindles become
less tuberculate and narrower (figs 3e, 4b).
The medulla has slender needles, up to 0.30 mm long, with simple tubercles and
spines (fig. 4c).
Colour.— Alive the colony was brown, preserved it is cream; all sclerites are colour-
less.
Etymology.— Named after Dr Bert W. Hoeksema, head of the department of Zool-
ogy/Marine Research, NNM, who collected most of the type material.
Remarks.— The species differs from Alertigorgia orientalis (fig. 2g-h) in its leaf-like
branches (fig. 2a-d; Hoeksema & Ofwegen, 2004). Furthermore, the sclerites are also
different, A. orientalis has larger polyp sclerites (fig. 5a-c); larger spindles in the surface
layer, and less ovals and cylinders (fig. 5d, 6a); and shorter rods in the medulla (fig. 6c).
Moreover, the surface layer has more sclerites with warts fused disk-like (fig. 6a).
Specimens of both Alertigorgia species grow almost exclusively on specimens of a
sponge species of the genus Oceanapia. On the leaf-like branches of several specimens of
A. hoeksemai an unidentified egg cowry (Gastropoda: Ovulidae) was observed, probably
feeding on the tissue (fig. 2e). This creates a unique association of an egg cowry living
on an octocoral, which on its turn lives on a sponge.
Distribution.— N. Sulawesi, Papua New Guinea, Bali.
Acknowledgements
M. Slierings (NNM) and Sue Horner (NTM) are thanked for curatorial assistance,
B.W. Hoeksema for use of his underwater photographs (fig. 2c-f).
References
Broch, H., 1916. Results of Dr. E. Mjöbergs Swedisch scientific expeditions to Australia 1910-1913. XI.
Alcyonarien.— Kungliga Svenska Vetenskapsakademiens Handlingar 52 (11): 3-48.
Colin, P.L. & C. Arneson, 1995. Tropical Pacific Invertebrates. A field guide to the marine invertebrates
occurring on tropical Pacific coral reefs, seagrass beds and mangroves.— Coral Reef Press, Beverly
Hills, California: i-vii + 1-296.
Erhardt, H. & H.A. Baensch, 1998. Meerwasser atlas band 4: Wirbellose.— Mergus, Melle, Germany: 1-
1214.
Hoeksema, B.W. & L.P. van Ofwegen, 2004. Indo-Malayan Reef Corals: a generic overview.— World
Biodiversity Database CD-ROM Series ETI, Amsterdam.
Kükenthal, W., 1908. Diagnosen neuer Gorgoniden (4. Mitteilung).— Zoologische Anzeiger 33 (1): 9-
20.
Ridley, S.O., 1884. Alcyonaria. Report on the Zoological Collections made in the Indo-Pacific Ocean dur-
ing the Voyage of H.M.S. ‘Alert’: 327-365, pls 36-38.
Verseveldt, J., 1940. Studies on Octocorallia of the families Briareidae, Paragorgiidae and Anthotheli-
dae.— Temminckia 5: i-v + 1-142, figs 1-52.
Received: 9.iv. 2007
Accepted: 17.iv.2007
Edited: C.H.J.M. Fransen
244 van Ofwegen & Alderslade. A new species of Alertigorgia. Zool. Med. Leiden 81 (2007)
van Ofwegen & Alderslade. A new species of Alertigorgia. Zool. Med. Leiden 81 (2007) 245
Fig. 2a-b, d, live colonies of Alertigorgia hoeksemai spec. nov. at Bunaken; c, at Bali; e, Ovulidae on A. hoek-
semai; f, close-up calyces; g-h, A. orientalis from Ambon.
Fig. 1. NTM C015585, holotype of Alertigorgia hoeksemai spec. nov.ᮤ
246 van Ofwegen & Alderslade. A new species of Alertigorgia. Zool. Med. Leiden 81 (2007)
Fig. 3. Sclerites of Alertigorgia hoeksemai spec. nov.; a, point spindles; b, collaret spindles; c, tentacle rod;
d, spindles and ovals of surface layer; e, spindles of subsurface layer. Scale 0.10 mm.
van Ofwegen & Alderslade. A new species of Alertigorgia. Zool. Med. Leiden 81 (2007) 247
Fig. 4. Sclerites of Alertigorgia hoeksemai spec. nov.; a, ovals of surface layer; b, spindles of subsurface
layer; c, needles from medulla. Scale 0.10 mm.
248 van Ofwegen & Alderslade. A new species of Alertigorgia. Zool. Med. Leiden 81 (2007)
Fig. 5. Sclerites of Alertigorgia orientalis (Ridley, 1884); a, point spindles; b, collaret spindles; c, tentacle
rods; d, spindles of surface layer. Scale 0.10 mm.
van Ofwegen & Alderslade. A new species of Alertigorgia. Zool. Med. Leiden 81 (2007) 249
Fig. 6. Sclerites of Alertigorgia orientalis (Ridley, 1884); a, ovals of surface layer; b, ovals from subsurface
layer; c, needles from medulla. Scale 0.10 mm.

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2007 -- Ofwegen and Alderslade_Alertigorgia

  • 1. A new species of Alertigorgia (Coelenterata: Octocorallia: Anthothelidae) from the Indo Malayan region L.P. van Ofwegen & P. Alderslade Ofwegen, L.P. van & P. Alderslade. A new species of Alertigorgia (Coelenterata: Octocorallia: Anthothe- lidae) from the Indo Malayan region. Zool. Med. Leiden 81 (13), 8.vi.2007: 241-249, figs 1-6.— ISSN 0024-0672. Leen P. van Ofwegen, Nationaal Natuurhistorisch Museum, P.O. Box 9517, 2300 RA Leiden, The Nether- lands (e-mail: ofwegen@naturalis.nl). Phil Alderslade, Museum and Art Gallery of the Northern Territory, P.O. Box 4646, Darwin, Northern Territory 0801, Australia (e-mail: Phil.Alderslade@nt.gov.au). (Current address: P.O. Box 211, Margate, Tasmania 7054, Australia). Key words: Coelenterata; Anthozoa; Octocorallia; Alertigorgia; new species; Ovulidae; Oceanapia; Indo- nesia. A new species of Alertigorgia from Indonesian waters is described and depicted, and compared with the hitherto only described species in the genus, A. orientalis (Ridley, 1884). The species is part of a unique association in which a mollusc species lives on an octocoral, which on its turn lives on a sponge species. Introduction Kükenthal (1908) established the genus Alertigorgia for a single species described by Ridley (1884) as Iciligorgia orientalis from Torres Strait, Australia. Since that original de- scription the species has been found regularly in the Malay Archipelago and Australia (for references see Verseveldt, 1940: 104), but the genus remained monotypic. However, Colin & Arneson (1995: 89) published an underwater picture of a species named Soleno- caulon sp., from Papua New Guinea, and a few years later, Erhardt & Baensch (1998: 271) showed a similar looking species labelled as “new genus and new species”, from Bunaken, Sulawesi, Indonesia. The second author got involved with the identification of these images, and was already in possession of colony fragments collected by the Coral Reef Research Foundation and Harry Erhardt when he discussed the matter with the first author, who also had seen images of this species, made by B.W. Hoeksema (NNM) in Bunaken national park. During a fieldtrip in 1998, B.W. Hoeksema & L.P. van Ofwegen photographed and collected several specimens at Bunaken and from this ma- terial it became obvious we were dealing with an as yet undescribed species of Alertig- orgia, which is described below. In 2001 the species was also found at Bali, Indonesia (see material examined). Titanideum mjöbergi Broch, 1916, should most likely also be included in Alertigorgia. The species is described as having cylindrical branches and may represent a third spe- cies of the genus. The original description lacks sufficient detail, and the validity of the species will be the subject of another paper. Abbreviations The NNM-LIPI-WWF Expedition, 2001, was a joint project of the National Museum of Natural History (NNM), Leiden, The Netherlands, the Research Centre for Oceanol- ogy, Indonesian Institute of Sciences (PPO-LIPI), and the World Wildlife Fund (WWF),
  • 2. 242 van Ofwegen & Alderslade. A new species of Alertigorgia. Zool. Med. Leiden 81 (2007) The material is deposited in the Zoological Museum Bogor (ZMB) of the LIPI Research Centre for Biology (PPB-LIPI, Cibinong), and the NNM (RMNH); NTM stands for Mu- seum and Art Gallery of the Northern Territory, Australia. Description Genus Alertigorgia Kükenthal, 1908 Diagnosis, amended: Anthothelidae with bushy or sparsely branched colonies. Branches may be cylindrical, flattened with acute edges or expanded and leaf-like. Pol- yps fully retractile into slit-like apertures, densely scattered over both faces in the spe- cies with leaf-like branches, arranged bilaterally in two narrow grooves on edges of conspicuously flattened branches, or singly or in small groups within short narrow grooves in the species with cylindrical branches. The medulla has prickly needles and thick rods with a few large tubercles; the surface layer has plump spindles with com- plex tubercles. Polyps have a collaret and points arrangement. Sclerites are colourless. Colonies are zooxanthellate. Alertigorgia hoeksemai spec. nov. (figs 1-6) Material examined.— NTM C015585, holotype, BUN.07, Indonesia, N. Sulawesi, Bunaken Park, ESE of Siladen island, 124°48’E 01°37’30”N, scuba diving, 35-40 m depth, 7.v.1998, coll. B.W. Hoeksema & L.P. van Ofwegen; paratypes: NTM C015586, same data as holotype; NTM C014257, Indonesia, North Sulawesi Sea, Bunaken Islands, 01°38’N 124°46’E, 12-18 m depth, 1996, coll. H. Erhardt; NTM C014258, Indonesia, Manado, 01°45.1’N 124°58.87’E, 3 m depth , coll. Coral Reef Research Foundation, 21.v.1993; RMNH Coel. 31952, same data as holotype; RMNH Coel. 31953, NNM-LIPI-WWF Exped., BAL.09, Indonesia, Bali, Sa- nur, Loloan Batu Agung, 08°43’31’’S 115°15’57’’E, slowly declining reef slope, sandy base, scuba-diving, 17 m depth, 3.iv.2001, coll. B.W. Hoeksema; RMNH Coel. 31954, NNM-LIPI-WWF Exped., BAL.14, Indone- sia, Bali, NE-side Pulau Serangan, off lighthouse, 08°43’17’’S 115°15’31’’E, slowly declining reef slope, sandy base, scuba-diving, 15-20 m depth, 5.iv.2001, coll. B.W. Hoeksema; RMNH Coel. 31955, NNM-LIPI- WWF Exped., BAL.15, Indonesia, Bali, NE-side Pulau Serangan; 08°44’03’’S 115°15’05’’E, slowly declining reef slope, sandy base, scuba-diving, 22 m depth, 7.iv.2001, coll. L.P. van Ofwegen & M. Slierings; MZB Coel. 3, NNM-LIPI-WWF Exped., BAL.13, Indonesia, Bali, NE-side Pulau Serangan, Loloan Serangan, 08°43’39’’S 115°15’13’’E, slowly declining reef slope, sandy base, scuba-diving to 17 m depth, 5.iv.2001, coll. M.I.Y.T. Hermanlimianto; MZB. Coel. 4, same data as RMNH Coel. 31955, coll. M.I.Y.T. Hermanlimianto. Material used for comparison.— RMNH Coel. 31956, Alertigorgia orientalis, BUN.16, Indonesia, N. Su- lawesi, S Tetapaan island, reef, 124°30’30”E 01°18’N, scuba diving -23 m. 12.v.1998, coll. B.W. Hoeksema & L.P. van Ofwegen. Description.— The holotype is 70 cm long and 40 cm wide (fig. 1). The lower part is attached to a sediment dwelling species of the sponge genus Oceanapia (see base of colony in fig 2c; Colin & Arneson, fig. 167) but it is difficult to see where exactly the sponge ends and the octocoral begins. All branches are expanded leaf-like. The polyps are densely scattered over both faces of the leaves. The calyces are dome-shaped and all polyps are retracted into slit-like apertures (fig. 2f). The polyps have a collaret of 2-3 rows of spindles and points with 4-6 spindles. All these spindles have simple tubercles and are up to 0.25 mm long (fig. 3a-b). The tenta- cles have small rods, up to 0.15 mm long (fig. 3c).
  • 3. van Ofwegen & Alderslade. A new species of Alertigorgia. Zool. Med. Leiden 81 (2007) 243 The surface layer of the leaves has ovals, cylinders, and spindles, up to 0.30 mm long, with densely placed complex tubercles (figs 3d, 4a); a few sclerites have some warts fused disk-like (fig. 4a, second sclerite). Towards the interior the spindles become less tuberculate and narrower (figs 3e, 4b). The medulla has slender needles, up to 0.30 mm long, with simple tubercles and spines (fig. 4c). Colour.— Alive the colony was brown, preserved it is cream; all sclerites are colour- less. Etymology.— Named after Dr Bert W. Hoeksema, head of the department of Zool- ogy/Marine Research, NNM, who collected most of the type material. Remarks.— The species differs from Alertigorgia orientalis (fig. 2g-h) in its leaf-like branches (fig. 2a-d; Hoeksema & Ofwegen, 2004). Furthermore, the sclerites are also different, A. orientalis has larger polyp sclerites (fig. 5a-c); larger spindles in the surface layer, and less ovals and cylinders (fig. 5d, 6a); and shorter rods in the medulla (fig. 6c). Moreover, the surface layer has more sclerites with warts fused disk-like (fig. 6a). Specimens of both Alertigorgia species grow almost exclusively on specimens of a sponge species of the genus Oceanapia. On the leaf-like branches of several specimens of A. hoeksemai an unidentified egg cowry (Gastropoda: Ovulidae) was observed, probably feeding on the tissue (fig. 2e). This creates a unique association of an egg cowry living on an octocoral, which on its turn lives on a sponge. Distribution.— N. Sulawesi, Papua New Guinea, Bali. Acknowledgements M. Slierings (NNM) and Sue Horner (NTM) are thanked for curatorial assistance, B.W. Hoeksema for use of his underwater photographs (fig. 2c-f). References Broch, H., 1916. Results of Dr. E. Mjöbergs Swedisch scientific expeditions to Australia 1910-1913. XI. Alcyonarien.— Kungliga Svenska Vetenskapsakademiens Handlingar 52 (11): 3-48. Colin, P.L. & C. Arneson, 1995. Tropical Pacific Invertebrates. A field guide to the marine invertebrates occurring on tropical Pacific coral reefs, seagrass beds and mangroves.— Coral Reef Press, Beverly Hills, California: i-vii + 1-296. Erhardt, H. & H.A. Baensch, 1998. Meerwasser atlas band 4: Wirbellose.— Mergus, Melle, Germany: 1- 1214. Hoeksema, B.W. & L.P. van Ofwegen, 2004. Indo-Malayan Reef Corals: a generic overview.— World Biodiversity Database CD-ROM Series ETI, Amsterdam. Kükenthal, W., 1908. Diagnosen neuer Gorgoniden (4. Mitteilung).— Zoologische Anzeiger 33 (1): 9- 20. Ridley, S.O., 1884. Alcyonaria. Report on the Zoological Collections made in the Indo-Pacific Ocean dur- ing the Voyage of H.M.S. ‘Alert’: 327-365, pls 36-38. Verseveldt, J., 1940. Studies on Octocorallia of the families Briareidae, Paragorgiidae and Anthotheli- dae.— Temminckia 5: i-v + 1-142, figs 1-52. Received: 9.iv. 2007 Accepted: 17.iv.2007 Edited: C.H.J.M. Fransen
  • 4. 244 van Ofwegen & Alderslade. A new species of Alertigorgia. Zool. Med. Leiden 81 (2007)
  • 5. van Ofwegen & Alderslade. A new species of Alertigorgia. Zool. Med. Leiden 81 (2007) 245 Fig. 2a-b, d, live colonies of Alertigorgia hoeksemai spec. nov. at Bunaken; c, at Bali; e, Ovulidae on A. hoek- semai; f, close-up calyces; g-h, A. orientalis from Ambon. Fig. 1. NTM C015585, holotype of Alertigorgia hoeksemai spec. nov.ᮤ
  • 6. 246 van Ofwegen & Alderslade. A new species of Alertigorgia. Zool. Med. Leiden 81 (2007) Fig. 3. Sclerites of Alertigorgia hoeksemai spec. nov.; a, point spindles; b, collaret spindles; c, tentacle rod; d, spindles and ovals of surface layer; e, spindles of subsurface layer. Scale 0.10 mm.
  • 7. van Ofwegen & Alderslade. A new species of Alertigorgia. Zool. Med. Leiden 81 (2007) 247 Fig. 4. Sclerites of Alertigorgia hoeksemai spec. nov.; a, ovals of surface layer; b, spindles of subsurface layer; c, needles from medulla. Scale 0.10 mm.
  • 8. 248 van Ofwegen & Alderslade. A new species of Alertigorgia. Zool. Med. Leiden 81 (2007) Fig. 5. Sclerites of Alertigorgia orientalis (Ridley, 1884); a, point spindles; b, collaret spindles; c, tentacle rods; d, spindles of surface layer. Scale 0.10 mm.
  • 9. van Ofwegen & Alderslade. A new species of Alertigorgia. Zool. Med. Leiden 81 (2007) 249 Fig. 6. Sclerites of Alertigorgia orientalis (Ridley, 1884); a, ovals of surface layer; b, ovals from subsurface layer; c, needles from medulla. Scale 0.10 mm.