Chrysaora chinensis Vanhöffen, 1888

Morandini, André C. & Marques, Antonio C., 2010, Revision of the genus Chrysaora Péron & Lesueur, 1810 (Cnidaria: Scyphozoa) 2464, Zootaxa 2464, pp. 1-97 : 13-16

Chrysaora chinensis Vanhöffen, 1888

( Figures 10 –12, 73, 85)

Chrysaora chinensis Vanhöffen 1888: 16 (description), 23 (description), 48 (distribution) [South China Sea, near Hong Kong – China].

Chrysaora helvola var. chinensis Mayer 1910: 580 (synoptic table), 582 (description) [non Chrysaora helvola Brandt, 1838 ].

Dactylometra quinquecirrha: Light 1914: 196 (list), 198–200 (description) [Manila Bay – Philippines]. Light 1921: 26, 28 (list), 30–31 (description) [Manila Bay – Philippines]. Stiasny 1919: 75–85 (description), text-figs a–c (medusa), Pl. II fig. 3 (medusa), Pl. II figs 4–6 (rhopalium), Pl. III fig. 7 (tentacle), Pl. III figs 8–9 (gastric filaments) [Malayan archipelago]. Stiasny 1921: 112–113 (description) [Semarang – Indonesia]. Stiasny 1929: 198 (mention) [Semarang – Indonesia] [non Chrysaora quinquecirrha ( Desor, 1848) ].

Dactylometra africana: Mayer 1915: 180 (description) [Manila Bay – Philippines, Kowloon – China]. Mayer 1917: 201– 202 (description) [Manila Bay, Luzon – Philippines, Kowloon – China]. Light 1921: 26, 28 (list); 30 (the author did not see the species, but considered the identification made by Mayer, from Manila Bay – Philippines, as doubtful) [non Chrysaora africana ( Vanhöffen, 1902) ].

Chrysaora melanaster: Mayer 1915: 179 (description), fig. 2 (medusa) [San Miguel Bay – Philippines]. Mayer 1917: 200–201 (description), fig. 8 (medusa) [San Miguel Bay, Luzon – Philippines]. Light 1921: 26 (list) [San Miguel Bay – Philippines] [non Chrysaora melanaster Brandt, 1835 ].

Chrysaora helvola: Vannucci 1954: 125 (commented that C. chinensis is identical to C. helvola ). Gao, Hong & Zhang 2002: 6 (mention), 33 (mention, tab. 7), 201–202 (brief description), fig. 118 (medusa) [ China] [non Chrysaora helvola Brandt, 1838 ].

? Dactylometra quinquecirrha: Stiasny 1940: 22 (distribution) [ Vietnam] [non Chrysaora quinquecirrha ( Desor, 1848) ].

? Chrisaora helvola: Naumov 1961: 62 (key), 64 (description), fig. 42 (medusa) [ Russia] [non Chrysaora helvola Brandt, 1838 ].

? Chrysaora quinquecirrha: Naumov 1961: 63–64 (key, description), fig. 41 (polyp), Pl. II (medusa) [ Russia]. Kramp 1968: 83 (description) [ Angola, Australia, and Indonesia] (specimens from Angola probably C. caliparea , specimens from Australia unidentifiable). Cornelius 1997: 122 (brief description), fig. (medusa) [Indian and Pacific Oceans] (part of the specimens might be identified as C. caliparea ). Heeger 1998: 156 (mention, table), 239 (mention), 295–296 (brief description), 24 (fig. 1–16, medusa), 63 (fig. 2–30d–e, tentacle), 70 (fig. 2–37, nematocyst wart), 230 (fig. 3–21c, medusa) [tropical and subtropical Pacific] [non Chrysaora quinquecirrha ( Desor, 1848) ].

? Chrysaora helvola: Heeger 1998: 156 (mention, table), 296–297 (brief description), 12 (fig. 1–1, medusa), 56 (fig. 2– 20, nematocyst warts), [tropical and subtropical Pacific] [non Chrysaora helvola Brandt, 1838 ].

Neotype specimen. Holotype is lost and a neotype specimen is designated. USNM 28680 (as Dactylometra africana , ~ 15 cm in diameter, 14.viii.1908, preserved in 4% formaldehyde solution, Kowloon – China) .

Examined material. Neotype; first author private collection (as Chrysaora helvola , ~ 5 cm in diameter, xi.1927, Java Sea, part of NNM 5241); NHM 1931.8.11.3-4 (as Dactylometra quinquecirrha , specimens ~10 and 12 cm in diameter, 1918, Cheribon – Indonesia), NHM 1932.2.6.16 (as Dactylometra quinquecirrha , ~ 5 cm in diameter, 1922, Tand Jong Priok – Indonesia); NNM 5241 (as Chrysaora helvola , ~ 6 cm in diameter, xi.1927, Java Sea), NNM 5243 (as Chrysaora helvola , specimens ~7 and 9 cm in diameter, ix.1927, west of Sumatra); USNM 28679 (as Dactylometra africana , ~5.5, 6, 7.5, 8, and 9 cm in diameter, 14.vi.1909, Manila Bay, Luzon island – Philippines), USNM 28712 (as Chrysaora melanaster , ~ 15 cm in diameter, 14.vi.1909, San Miguel Bay, Luzon – Philippines).

Type locality. South China Sea (near Hong Kong).

Distribution. Western Pacific Ocean ( China, Indonesia, Philippines, Sumatra) ( Fig. 73).

Diagnosis. Medusa size medium, typical dimensions ~ 10 cm in diameter; marginal lappets slightly elongate, up to 6 per octant, without canals; tentacles 24–40 (3–5 per octant, 3-2-1-2-3), clefts with similar depth; quadralinga absent; colouration (adults) exumbrella either transparent or background pinkish with radiating reddish stripes, tentacles and lappets reddish-brown.

Neotype specimen description. USNM 28680 (as Dactylometra africana ), damaged specimen. Umbrella flatter than a hemisphere, diameter 15 cm. Exumbrellar surface finely granulated. Mesoglea flexible, about 0.5 cm thick on central portion, thinning towards margin. Marginal lappets 48, round, sharper at tip, 6 per octant (2 rhopalar and 4 tentacular); rhopalar lappets slightly longer and broader than tentacular ones. Rhopalia 8, without ocelli, in clefts; exumbrellar sensory pit deep, blind-ended. Tentacle clefts similar in depth. Tentacles 24 (3 per octant), as long as umbrellar diameter; arranged as 2-1-2 (primary tentacle central, and 2 secondary tentacles laterally). Subumbrellar and tentacular musculature not distinguishable. Mouth-disc circular, with four evident corners, grooved. Pillars evident, 1 cm wide, delimited by corners of insertion of manubrium. Subgenital ostia oval, ~ 5 cm in diameter. Oral arms ca. 25 long (28 cm according to Mayer, 1915: 180), V-shaped in cross section, edges delicate and slightly convoluted. Central stomach circular, marginal region limited by insertion of radial septa. Stomach pouches 16, width uniform at basal part; tentacular pouches enlarged distally. Radial septa thin, with rounded base; straight up to ¼ of margin, then making an “S” (first thinning tentacular pouch, then enlarging it); ending near tentacular base at rhopalar lappet. Gastric filaments in four interradial fields. Quadralinga absent. Gonads rimming gastric filaments, in semicircle, greatly folded; female.

Description of other specimens and additional data. Medusa: Umbrella diameter up to 16 cm (range 5– 15 cm), flatter than a hemisphere. Exumbrellar surface finely granulated. Colouration according to Mayer (1915): tentacles and lappets reddish-brown, exumbrella background pinkish with radiating reddish stripes, larger specimens darker; according to Light (1921): entirely transparent. Mesoglea flexible, relatively thick on central portion, thinner at edge. Marginal lappets rounded, sharper at the tip, 4–6 per octant (Figs 11–12); rhopalar lappets slightly larger than velar, rhopalar lappets sometimes overlapping (“closed rhopalia” condition). Rhopalia 8, without ocelli, in deep clefts; deep exumbrellar sensory pit. Tentacle clefts similar in depth. Tentacles 24–40 (3–5 per octant, 3-2-1-2-3) (Fig. 11–12); primary tentacle central, secondary and tertiary tentacles lateral. Musculature not distinguishable. Brachial disc circular, with four evident corners. Pillars evident, width ~ 1 / 8 of umbrellar diameter. Quadralinga absent. Subgenital ostia rounded, ca. 1 / 5 of umbrellar diameter. Oral arms 1–3 times longer than umbrellar diameter ( Fig. 10), V-shaped, delicate and convoluted edges, terminal portion slightly spiraled. Central stomach circular, marginal region limited by insertion of radial septa. Stomach pouches 16, width uniform centrally; tentacular pouches enlarged distally, but narrow before enlarge. Septa narrow, widening gradually at proximal end (rounded base); straight up to ca. 1 / 2, then bent towards each other, then bent towards rhopalia (~45º) (Figs 11–12). Gastric filaments in four interradial fields. Gonads outlining gastric filaments, semicircular ring, greatly folded. Cnidome ( Fig. 85): Specimen USNM 28680 (neotype), medusa tentacles with holotrichous O-isorhizas [n=10; 13.7–19.6 x 12.7–17.6 µm (mean = 16.07 x 14.6 µm)]; holotrichous a-isorhizas [n=10; 5.8–6.8 x 1.9–2.9 µm (mean = 6.37 x 2.84 µm)]; holotrichous Aisorhizas [n=10; 12.7–14.7 x 6.8–8.8 µm (mean = 14.01 x 7.94 µm)]; heterotrichous microbasic rhopaloids [n=10; 8.8–11.7 x 5.8–6.8 µm (mean = 10.48 x 6.76 µm)]; Specimen on the first author collection, medusa tentacles with holotrichous O-isorhizas [n=10; 17.6–21.5 x 13.7–20.5 µm (mean = 20.48 x 17.93 µm)]; holotrichous a-isorhizas [n=10; 4.9–5.8 x 2.9–3.9 µm (mean = 5.78 x 3.33 µm)]; holotrichous A-isorhizas [n=10; 13.7–16.6 x 7.8–9.8 µm (mean = 14.9 x 8.92 µm)]; heterotrichous microbasic rhopaloids [n=10; 9.8– 11.7 x 4.9–6.8 µm (mean = 10.29 x 5.88 µm)];Specimen USNM 28712, medusa tentacles with holotrichous O-isorhizas [n=10; 13.7–29.4 x 12.7–24.5 µm (mean = 20.97 x 18.03 µm)]; holotrichous a-isorhizas [n=10; 4.9–5.8 x 2.9–3.9 µm (mean = 5.68 x 3.33 µm)]; holotrichous A-isorhizas [n=10; 14.7–19.6 x 6.8–9.8 µm (mean = 16.66 x 9.11 µm)]; heterotrichous microbasic rhopaloids [n=10; 8.8–11.7 x 4.9–5.8 µm (mean = 10.39 x 5.78 µm)].

Systematic remarks. Chrysaora chinensis was described by Vanhöffen (1888) based on a single specimen collected near Hong Kong. The author gave only data on size (7 cm), colouration (faint star figure on exumbrella) and oral arms (very long, up to 25 cm). We could not locate the type specimen and no other author ever reported finding the species. Mentions of the species in the literature amount to repetition of Vanhöffen’s specimen data and record. We believe that all Chrysaora specimens occurring in China, Indonesia, and Philippines should be assigned to C. chinensis . The specimens of that area have been referred to as Dactylometra africana , C. helvola , C. melanaster , and D. quinquecirrha , probably due to their colouration pattern and presence (in some specimens) of 40 tentacles. The shape of the radial septa conforms to the group of species from the Pacific Ocean. The specimens cannot be identified as D. africana (valid name C. fulgida ) or as D. quinquecirrha (valid name C. quinquecirrha ) because of the differences in shape of the radial septa. They also cannot be referred to C. helvola (valid name C. fuscescens ) or C. melanaster due to the difference in tentacle and lappet number. Both of the latter species also attain a larger size than in observed specimens of C. chinensis . The species that most resembles C. chinensis is C. pacifica ; both have comparable sizes and colour patterns, and the number of tentacles and lappets are also similar. The only known distinction between C. chinensis and C. pacifica is the size of the holotrichous O-isorhiza nematocysts, which are two times smaller in C. pacifica (see Tab. II). Based on this cnidome difference we resurrect the species C. chinensis .

Biological data. Considered the most dangerous species of Manila Bay ( Philippines) (according to Light 1921: 31).

Etymology. chinensis : derived from the area where the species was discovered (South China Sea, near Hong Kong, northwestern Pacific Ocean).