Research Article
Research Article
A new species of Arachnanthus from the Red Sea (Cnidaria, Ceriantharia)
expand article infoSérgio N. Stampar, Suraia O. El Didi, Gustav Paulay§, Michael L. Berumen|
‡ Univ Estadual Paulista, Assis, Brazil
§ University of Florida, Gainesville, United States of America
| King Abdullah University of Science and Technology, Thuwal, Saudi Arabia
Open Access


A new species of the genus Arachnanthus (Cnidaria: Ceriantharia), Arachnanthus lilith Stampar & El Didi, sp. n., is described. This species is widely distributed in the Red Sea, and recorded from 2–30 m depths. Arachnanthus lilith Stampar & El Didi, sp. n. is the fifth species of the genus and the first recorded from the Red Sea. The number of labial tentacle pseudocycles, arrangement of mesenteries, and distribution of acontioids allow the differentiation of the new species from other species of the genus.


Anthozoa , biodiversity, coral reefs, Indo-West Pacific, marine invertebrates, taxonomy


While tube anemones are common objects for underwater photographers and are widely exhibited in aquaria, they remain undersampled in most regions of the world, and the diversity and distribution of species remains poorly documented (Stampar et al. 2016). This is especially true for species that are difficult to observe and collect, because of nocturnal habits, small body size, or deeply extended burrows. The small, nocturnal tube anemones of the family Arachnactidae are a case in point (den Hartog 1977; Stampar et al. 2012, 2015a). This family is comprised of two benthic genera, Arachnanthus Carlgren, 1912 and Isarachnanthus Carlgren, 1924 (Stampar et al. 2016), although other genera have been proposed based only on larval forms (Molodtsova 2004). However, larval genera are not currently linked to those of adults and therefore their status remains unclear (Stampar et al. 2015a). Carlgren (1912) established Arachnanthus for A. sarsi (which he described from the North Sea) together with Cerianthus oligopodus Cerfontaine, 1891 from the Mediterranean. Carlgren (1924, 1937) later described A. bockii Carlgren, 1924 from Fiji and A. australiae Carlgren, 1937 from Australia. Since these studies, the genus has received little attention, with Picton and Manuel’s (1985) study and redescription of A. sarsi being the most substantive. Here a fifth species of Arachnanthus is described, the first known from Red Sea.

Materials and methods

Specimens were collected by hand at three sites across the Red Sea, from the Gulf of Aqaba to the Farasan Islands, in Saudi Arabia (Fig. 1). Collected polyps were preserved in 10 % buffered seawater formaldehyde solution, and later transferred to 75 % ethanol. The holotype and five paratypes are deposited in the Invertebrate Collections of the Florida Museum of Natural History, University of Florida (UF Cnidaria).

Figure 1. 

Records of Arachnanthus lilith sp. n. individuals studied, collected in Saudi Arabia (dark gray). A – UF Cnidaria 9168 (Holotype), B – UF Cnidaria 9167, UF Cnidaria 9227, UF Cnidaria 9229, UF Cnidaria 9230 (Paratype) and C – UF Cnidaria 9076.

The anatomical study of polyps and cnidome were based on characters defined by previous authors (Carlgren 1912; den Hartog 1977; Stampar et al. 2012, 2015b). Six specimens were opened along the ventral side (opposite the siphonoglyph), using surgical scalpels, for anatomical study.

The classification of cnidae follows England (1991) and Stampar et al. (2015b). Thirty undischarged capsules were measured for each cnida type, sampled from each body region of two specimens (UF Cnidaria 9168 & 9229). The cnidome was studied with a Nikon Eclipse E200 microscope at 1000x magnification. Each part of the body was analyzed separately to avoid any contamination.


Class Anthozoa Ehrenberg, 1834

Subclass Ceriantharia Perrier, 1883 (sensuStampar et al. 2014)

Suborder Penicillaria den Hartog, 1977

Family Arachnactidae Carlgren, 1912

Arachnanthus Carlgren, 1912


Arachnactidae with sterile protomesenteries; metamesenteries in duplets (M and B), long (‘M’) metamesenteries with gonads and a double mesenteric filament, short (B) betamesenteries sterile, with single, convoluted mesenteric filament; very long stomodeum; lacking a directive labial tentacle; cnidome with p-mastigophores and b-mastigophores (after Carlgren 1912, 1924, 1937 and den Hartog 1977).

Type species

Arachnanthus oligopodus (Cerfontaine, 1891)

Valid species

Arachnanthus australiae Carlgren, 1937

Arachnanthus bockii Carlgren, 1924

Arachnanthus oligopodus (Cerfontaine, 1891)

Arachnanthus sarsi Carlgren, 1912

Arachnanthus lilith sp. n.


North Sea, Mediterranean Sea, Red Sea, East Australia, and Melanesia.

Arachnanthus lilith Stampar & El Didi, sp. n.

Figs 1, 2, 3, 4, Tables 1, 2

Material examined (six specimens)

Holotype: UF Cnidaria 9168, adult individual (35 mm long), Saudi Arabia, island near Jaz’air Sila, (27.651°N, 35.2832°E) (Fig. 1A), 10–30 m depth, fore reef, under rocks, G. Paulay, Seabird McKeon, Daisuke Uyeno coll. (27/ix/2013). Paratypes: UF Cnidaria 9167, adult (31 mm long), same data as holotype. UF Cnidaria 9227, adult (35 mm long), UF Cnidaria 9229, adult (42 mm long), UF Cnidaria 9230, adult (26 mm long) all three from Saudi Arabia, Gulf of Aqaba, Joey’s Shipwreck Bay, (28.1846°N, 34.6381°E) (Fig. 1B), 7–13 m depth, in sand and seagrass bed, collected at night, G. Paulay, Daisuke Uyeno, Casey Zakroff coll. (01/x/2013). UF Cnidaria 9076 (Fig. 2D), adult, Saudi Arabia, Farasan Banks, Atlantis Shoal (18.1917°N, 41.1138°E) (Fig. 1–C), 9–11 m depth, sandy shoal with patch reefs, in sand, collected at night, Arthur Anker, Patrick Norby, Gustav Paulay coll. (07/iii/2013).

Figure 2. 

Arachnanthus lilith sp. n. A (Paratype UF Cnidaria 9227) (not to scale) B (Paratype UF Cnidaria 9168) (not to scale) C–D Live specimens in nature (not included as paratypes – ICZN 72.4.6) (not to scale) E Dissected specimen with detail of acontioids (arrows) (scale bar 2 mm) UF Cnidaria 9168 (Holotype) F Detail of oral disc UF Cnidaria 9229 (Paratype) with detail on tentacular pores with green fluorescent protein (GFP) (arrows) (not to scale).


Small ceriantharian, up to at least 42 mm long, 4–6 mm wide. With 19–24 translucent marginal tentacles (3–5 mm long in preserved specimens), each with 2–4 brown bands (Fig. 1); tentacle arrangement (1)…; at least 5 pores per tentacle, pores marked by concentration of green fluorescent protein (GFP) (Fig. 1–F); unpaired marginal tentacle present. With 11–15 pale labial tentacles (up to 2 mm long in preserved specimens), tentacle arrangement (0)…; unpaired labial tentacle absent. Long actinopharynx extending over 1/3 of total body length, hyposulcus 3–4 mm long, hemisulci distinct; siphonoglyph wide, connected to eight mesenteries; directive mesenteries a little shorter than hyposulcus. Three pairs of protomesenteries (P), P2 and P4 long and P3 short, metamesenteries (M), long, fertile with double mesenteric filament; betamesenteries (B) short, sterile with single mesenteric filament (double in a short part immediately below actinophrarynx) and rather convoluted; acontioids only in mesenteries M3 and M4; see Fig. 2 for schematic arrangement of mesenteries. Cnidome (Fig. 3) of spirocysts, atrichs, microbasic b-mastigophores (three types), microbasic p-mastigophores (two types), and ptychocysts; distributed as shown in Table 1.

Table 1.

Cnidome of Arachnanthus lilith sp. n. based on two specimens (UF 9229; 9168). Mean and range given for each cnida.

Length (in µm) Width (in µm)
Column Pytchocysts 40.69 (37.7–44.2) 9.18 (9.1–10.4)
Atrichs 48.57 (41.6–53.3) 8.01 (6.5–10.4)
b-mastigophores I 32.32 (31.2–33.8) 4.11 (3.9–5.2)
p-mastigophores I 86.45 (83.2–89.7) 21.49 (19.5–23.4)
Marginal tentacles p-mastigophores I 84.15 (78.0–91.0) 19.84 (18.2–20.8)
p-mastigophores II 33.75 (31.2–37.7) 6.84 (6.5–7.8)
b-mastigophores I 32.63 (31.2–33.8) 3.9 (3.8–4.0)
b-mastigophores II 21.06 (19.5–27.3) 4.11 (3.9–5.2)
Atrichs 34.92 (31.2–39.0) 6.58 (5.2–7.8)
Labial tentacles p-mastigophores I 64.87 (61.1–67.6) 13.08 (11.7–14.3)
b-mastigophores II 25.3 (20.8–28.6) 5.07 (3.9–7.8)
Atrichs 25.69 (24.7–28.6) 6.02 (5.2–6.5)
Stomodeum p-mastigophores I 49.44 (45.5–54.6) 9.83 (7.8–11.7)
b-mastigophores II 24.05 (20.8–26.0) 5.76 (5.2–6.5)
Atrichs 33.28 (31.2–35.1) 6.54 (5.2–7.8)
Betamesenteries p-mastigophores I 83.8 (80.6–89.7) 23.14 (19.5–24.7)
p-mastigophores II 54.9 (52.0–58.5) 15.34 (13.0–16.9)
b-mastigophores II 19.24 (15.6–23.4) 4.03 (3.9–5.2)
Metamesenteries b-mastigophores II 25.04 (23.4–26.0) 5.76 (5.2–6.5)
b-mastigophores III 17.76 (16.9–18.2) 4.2 (3.9–5.2)
Figure 3. 

Graphical representation of the arrangement of mesenteries of Arachnanthus lilith sp. n. Abbreviations: M.C. multiplication chamber, D directives, T.P. terminal pore, S siphonoglyph, B betamesenteries (convoluted mesentery), M metamesenteries (double filament), P protomesenteries, AC acontioids

Figure 4. 

Cnidome of Arachnanthus lilith sp. n. A microbasic p-mastigophores I B microbasic p-mastigophores II C Atrich D Ptychocyst E microbasic b-mastigophores I F microbasic b-mastigophores II G microbasic b-mastigophores III.


Presently known only from the Saudi Arabian Red Sea, from the Gulf of Aqaba to the Farasan Islands in the southern Red Sea. The species was found extended only at night.


The specific name lilith refers to the mythological figure of a female night demon in the vicinity of the Red Sea to ancient Mesopotamia (Saudi Arabia to Iraq).

Live color

Column pinkish tan at basal half or along most of its length, becoming clear toward base of tentacles. Marginal tentacles whitish/transparent, with brown and light green bands; extent of banding variable, with a basal brown band commonly developed. Labial tentacles clear to brown, with whitish base and tips. Oral disk with green and white colors.

Description of holotype

(UF Cnidaria 9168). Small polyp, 35 mm long, 4 mm in diameter just below the marginal tentacles, 3 mm diameter near aboral end. With 19 marginal tentacles arranged in two pseudocycles, each 4 mm long and 0.5 mm in diameter near base, tentacle arrangement (1)…. With 12 labial tentacles, each ~1 mm long, brown with a white apical tip, directive labial tentacle absent, tentacle arrangement (0)…. Oral disc 0.7 mm wide, actinopharynx 17 mm long, light beige to light brown, siphonoglyph wide and elongate with eight mesenteries attached, hyposulcus 9 mm long. Directive mesenteries shorter than actinopharynx. Protomesenteries as in diagnosis, M-mesenteries (M), long, fertile with a double mesenteric filament; B-mesenteries (B) short, sterile with single mesenteric filament (double in a short part immediately below actinopharynx) and rather convoluted; acontioids only in mesenteries M3 and M4.

Comparison with other members of the genus

Although Fautin et al. (2007) suggested that morphology alone is insufficient to distinguish species of this genus, internal anatomical characters do actually separate all known species (Table 2). While there are cases of cryptic species among tube-dwelling anemones (Stampar et al. 2012), none are yet documented for Arachnanthus.

Table 2.

Comparison of anatomical features of species of Arachnanthus (after Carlgren 1912b; Carlgren 1924; Carlgren 1937; Picton and Manuel 1985; this study).

A. australiae A. bockii A. oligopodus A. sarsi A. lilith sp. n.
Marginal tentacles Up to 40 Up to 30 ~20 Up to 35 Up to 24
Arrangement of labial tentacles (0) (0) (0) (0) (0)
Length of actinopharynx ~2/3 of gastric cavity ~1/2 of gastric cavity ~1/2 of gastric cavity ~1/2 of gastric cavity >1/2 of gastric cavity
Hyposulcus ~1/2 size of stomodeum ~1/2 size of stomodeum ~2X size of stomodeum < size of stomodeum = size of stomodeum
Oral disc diameter ~0.7 cm ~1 cm 0.5 cm
Maximum n° of mesentery attached to siphonoglyph 12 12 4 6 8
Directive mesenteries = length of Actinopharynx < length of Actinopharynx > length of Actinopharynx < length of Actinopharynx < length of Actinopharynx
P(C)2 Short, 1/2 of gastric cavity Very short, 1/4 of gastric cavity Short, 1/2 of gastric cavity Long, 3/4 of gastric cavity Long, 6/7 of gastric cavity, almost to aboral pole
P(C)3 Very short, <1/4 of gastric cavity Very short, <1/4 of gastric cavity Short, ~1/2 of gastric cavity Short, ~1/3 of gastric cavity Short, 1/3 of gastric cavity
M1 Almost to aboral pore Almost to aboral pore To aboral pore Almost to aboral pore To aboral pore
M3 4/5 of gastric cavity Almost to aboral pore 1/5 of gastric cavity Almost to aboral pore 3/4 of gastric cavity
Cnido-glandular tract of fertile mesenteries Present (short?) Present (short?) Present Present Present
Cnido-glandular tract of B Present (short?) Present (short?) Present (short?) Present (short) Present (short)
Acontioids Only in M1, M2 and M3 Only in M1, M2 and M3 Only in M1 Only in M1, M2 and M3 Only in M3 and M4
Distribution Northern Australia Fiji Mediterranean Sea North Sea Red Sea

Arachnanthus lilith has labial tentacles in three pseudocycles, unlike A. australiae, A. oligopodus, and A. sarsi, which all have them in one pseudocycle, while in A. bockii labial tentacles are not clearly organized and may be considered to fall into one or two pseudocycles. The actinopharynx is 2/3 as long as the gastric cavity in A. australiae, less than ½ as long in the other three described species, and a little over ½ as long in A. lilith. The maximum number of the mesenteries attached to the siphonoglyph is especially useful for distinguishing species: A. australiae and A. bockii have 12 each, A. lilith has eight, A. sarsi six, while A. oligopodus has four. The organization of mesenteries, particularly the mesentery P2 and M3, also provides useful characters to separate species (Table 2). Finally, the distribution of acontioids is also quite different in some species, especially in A. lilith where acontioids are present only on mesenteries M3 and M4. These mesenterial characters serve well to differentiate species of Arachnanthus, although how they vary over the ontogeny of each species remains to be studied.

Finally, the present study demonstrates the importance of more detailed investigations using non-standard collecting techniques. Small ceriantharians are rarely collected as they are frequently nocturnal and can be difficult to extract from the sediment as they retract quickly and rapidly. There are few described species of Ceriantharia with small body sizes; however, this may be the result of sampling limitations.


SED is thankful to the CNPq for a fellowship. This work was partly supported by São Paulo Research Foundation FAPESP 2015/24408-4, 2016/50389-0, 2017/50028-0, CNPq 404121/2016-0 and CAPES/CNPQ – PROTAX II 88882.156878/2016-01 to SNS, as well as the King Abdullah University of Science and Technology (award number CRG-01-BER-2012-002 to MLB). We are also grateful to Drs. Bert W. Hoeksema and Kensuke Yanagi for their helpful comments and suggestions.


  • Carlgren O (1912) Ceriantharia. Danish Ingolf-Expeditie 5(3): 1–78.
  • Carlgren O (1924) Papers from Dr. Th. Mortensen´s Pacific Expedition 1914–16 XVI. Ceriantharia. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening 75: 169–195.
  • Carlgren O (1937) Ceriantharia and Zoantharia. Scientific Reports of the Great Barrier Reef Expedition 1928–29, 5: 177–207.
  • Cerfontaine P (1891) Notes préliminaires sur l’organisation et le développement de différentes formes d’Anthozoaires. IV. Sur un nouveau cerianthe du golfe de Naples, Cerianthus oligopodus (n. sp.). Bulletin de l’Académie Royale des Sciences, des Lettres et des Beaux-Arts de Belgique 21: 32–39.
  • England KW (1991) Nematocysts of sea anemones (Actiniaria, Ceriantharia and Corallimorpharia: Cnidaria): nomenclature. Hydrobiologia 216/217: 691–697.
  • den Hartog JC (1977) Descriptions of two new Ceriantharia from the Caribbean Region, Pachycerianthus curacaoensis n. sp. and Arachnanthus nocturnus n.sp., with a discussion of the cnidome and of the classification of the Ceriantharia. Zoologische Mededelingen 51: 211–248.
  • Molodtsova TN (2004) On the taxonomy and presumable evolutionary pathways of planktonic larvae of Ceriantharia (Anthozoa, Cnidaria). In: Fautin DG, Westfall JA, Cartwrigh P, Daly M, Wyttenbach CR (Eds) Coelenterate Biology 2003.Springer, Dordrecht, 261–266.
  • Stampar SN, Maronna MM, Vermeij MJ, Silveira FL, Morandini AC (2012) Evolutionary diversification of banded tube-dwelling anemones (Cnidaria; Ceriantharia; Isarachnanthus) in the Atlantic Ocean. PLoS ONE 7(7): e41091.
  • Stampar SN, Maronna MM, Kitahara MV, Reimer JD, Morandini AC (2014) Fast-evolving mitochondrial DNA in Ceriantharia: A reflection of Hexacorallia paraphyly? PLoS ONE 9(1): e86612.
  • Stampar SN, Morandini AC, Branco LC, Silveira FL, Migotto AE (2015a) Drifting in the oceans: Isarachnanthus nocturnus (Cnidaria, Ceriantharia, Arachnactidae), an anthozoan with an extended planktonic stage. Marine Biology 162: 2161–2169.
  • Stampar SN, Beneti JS, Acuña FH, Morandini AC (2015b) Ultrastructure and tube formation in Ceriantharia (Cnidaria, Anthozoa). Zoologischer Anzeiger 254: 67–71.
  • Stampar SN, Maronna MM, Kitahara MV, Reimer JD, Morandini AC (2016) Ceriantharia in current systematics: Life cycles, morphology and genetics. In: Goffredo S, Dubinsky Z (Eds) The Cnidaria, Past, Present and Future: The world of Medusa and her Sisters.Springer International Publishing, Cham, 61–72.
login to comment