Chrysaora fulgida ( Reynaud, 1830 )

Morandini, André C. & Marques, Antonio C., 2010, Revision of the genus Chrysaora Péron & Lesueur, 1810 (Cnidaria: Scyphozoa) 2464, Zootaxa 2464, pp. 1-97 : 19-22

Chrysaora fulgida ( Reynaud, 1830)

( Figures 19 –23, 75, 87)

Medusa (Rhyzostoma) fulgidum Reynaud 1830: 79–80 (original description), Pl. XXV (medusa) [False Bay – South Africa].

Chrysaora Reynaudii Lesson 1843: 401–402 (description) (the species described by Reynaud was transferred to the genus Chrysaora under a new name) [non Chrysaorae Reynodii Brandt, 1835].

Chrysaora Reynaudi : L. Agassiz 1862: 166 (mention).

Chrysaora fulgida: Haeckel 1880: 514 (description) [ South Africa]. von Lendenfeld 1884: 269 (description) [False Bay – South Africa]. Vanhöffen 1888: 23 (brief description), 47 (distribution). Vanhöffen 1902: 38 (mention). Vanhöffen 1908: 39 (mention). Vanhöffen 1920: 17 (mention). Mayer 1910: 579 (synoptic table). Stiasny 1934: 388–389 (description, commented that the specimens are probably C. hysoscella ) [Hoetjes Bay – South Africa]. Ranson 1945: 312, 316 (mention, types) [Cape of Good Hope – South Africa]. Vannucci 1954: 125 (commented that C. fulgida is identical to C. hysoscella ). Kramp 1955: 296 (mention), 298 (mention, considered the synonymy C. fulgida = Dactylometra africana as doubtful). Kramp 1961: 324 (synonymy). O’Sullivan 1982: 29 (mention, = C. africana ). Pagès, Gili & Bouillon 1992: 50 (mention). Gershwin & Collins 2002: 128 (mentioned as nominal species with insufficient data).

Dactylometra africana Vanhöffen 1902: 40 (original description), Pl. IV fig. 20 (medusa). Mayer 1910: 588 (description), fig. 373 (medusa, figure reproduced from Vanhöffen, 1902). Bigelow 1913: 91 (mention, similarities with C. quinquecirrha – never more than 5 tentacles per octant). Stiasny 1939: 174–182 (description; D. africana probably an older fulgida ), fig. 2 (medusa, reproduction of original figure), figs 3–7 (medusa, aboral view of octants) [ South Africa]. Ranson 1949: 142–143 (description) [ Angola, Mauritania, Guinea-Bissau, Namibia]. Kramp 1955: 297 (synonymy), 298 ( D. africana = D. quinquecirrha ; considered the synonymy C. fulgida = D. africana as doubtful), 309–310 (mention, D. africana = C. quinquecirrha ). Thiel 1966: 19–22 (description), Pl. III fig. 5 (rhopalium) [Point Noire – Congo].

Chrysaora hysoscella var. fulgida Mayer 1910: 581 (brief description). Pagès, Gili & Bouillon 1992: 50 (mention) [non Chrysaora hysoscella ( Linnaeus, 1767) ].

Dactylometra quinquecirrha: Stiasny 1931: 139 (mention) [Lagos – Nigeria] [non Chrysaora quinquecirrha ( Desor, 1848) ].

Dactylometra fulgida: Stiasny 1939: 172–173 (description), 173–184 (discussion, probably the Chrysaora stage of D. africana ), fig. 1 (medusa, reproduction of the original figure) [ South Africa].

? Chrysaora quinquecirrha: Kramp 1955: 297–300 (description, D. africana = D. quinquecirrha , comments on differences with C. lactea , comparison of nematocysts with C. hysoscella ), 305 (ephyrae), 309 (mention), 314 (tab. III), 317 (mention) [ Angola; Nigeria]. Mianzan & Cornelius 1999: 538 (description), fig. 2.17 (distribution), figs 5.14a– b (medusa) [eastern Atlantic Ocean, Africa] [non Chrysaora quinquecirrha ( Desor, 1848) ].

Chrysaora africana: Kramp 1961: 323–324 (synonymy). O’Sullivan 1982: 29 (mention). Gershwin & Collins 2002: 128 (mentioned as nominal species with insufficient data).

Holotype specimen. MNHN 2318 as Medusa (Rhyzostoma) fulgida (~ 8.5 cm in diameter, ethanol, Cape of Good Hope – South Africa).

Examined material. Holotype; NHM 1937.7.19.343 (~ 18 cm in diameter, 05.x.1926, Saldanha Bay – South Africa), NHM 1937.7.19.344 (~ 23 cm in diameter, 05.x.1926, Saldanha Bay – South Africa), NHM 1987.10.8.1 (as Chrysaora cf. africana , ~ 4 cm in diameter, ii.1987, St. Paul River, Monrovia – Liberia); NNM 5361 (~ 17 cm in diameter, viii.1938, Walvis Bay – Namibia); USNM 53871 (as C. quinquecirrha , specimens ~3, 3, 5, 6, 7, 7, and 7 cm in diameter, 18.xii.1959, 4% formaldehyde solution, Ghana border, Gulf of Guinea – Togo); ZMA 2019 (as Dactylometra fulgida , ~ 17 cm in diameter, 1938, Walvis Bay – Namibia); ZMB CNI 5949 (types of Dactylometra africana , specimens ~7.5, 7.5, 9.5, and 10 cm in diameter, 10.x.1898, Fish Bay – Liberia), ZMB CNI 14803 (specimens ~ 5–7 cm in diameter, 31.x.1898, Algoa Bay – South Africa); ZMH C7451 (as Dactylometra africana , specimens ~14, 15, and 17 cm in diameter, no date, Point Noire – Congo), ZMH C7452 (as Dactylometra africana , specimens ~10, 11, 12, and 16 cm in diameter, 29.viii.1963, Point Noire – Congo); ZMUC not-numbered (specimens ~10 and 12 cm in diameter, x.1937, False Bay – South Africa), ZMUC not-numbered (as Dactylometra quinquecirrha , ~ 7 cm in diameter, 20.xii.1951, Lobito Harbour – Angola).

Type locality. Cape of Good Hope – South Africa.

Distribution. Coast of west Africa ( Fig. 75).

Diagnosis. Medusa of medium size (10–20 cm in diameter); marginal lappets rounded (in adults), up to 8 per octant, without canals; tentacles up to 56 (7 per octant, 4-3-2-1-2-3-4), but usually 24 (3 per octant, 2-1-2) or 40 tentacles (5 per octant, 3-2-1-2-3); quadralinga absent; colouration (adults) yellowish-brown, brownish, with radial pattern of darker triangles.

Holotype specimen description. Umbrella less than a hemisphere, ~ 8.5 cm in diameter ( Figs 19 –20). Exumbrellar surface finely granulated. Mesoglea flexible, thicker centrally. Marginal lappets 4 per octant (2 rhopalar, 2 tentacular), rounded; without canals of gastrovascular system; rhopalar lappets not overlapping (“open rhopalia” condition). Rhopalia 8, without ocelli, in deep clefts; exumbrellar sensory pit deep. Tentacular clefts vary in depth. Tentacles 24 (3 per octant), as long as bell diameter, arranged as 2nd-1st-2nd (primary tentacle central, secondary tentacles laterally). Subumbrellar musculature not distinguishable. Brachial disc circular, with 4 evident corners. Pillars evident, 2 cm wide, delimited by insertion corners of manubrium. Subgenital ostia rounded, 0.7 cm in diameter. Oral arms ca. bell diameter long, V-shaped. Central stomach circular, marginal region limited by insertion of radial septa. Stomach pouches 16, width uniform centrally; tentacular pouches enlarged distally. Radial septa narrow, at proximal end wider, pear-shaped; outer 1/3 bending towards rhopalia (~45º); ending near tentacular base at rhopalar lappet. Gastric filaments not observed. Quadralinga absent. Gonads semicircular ring, greatly folded; sex not determined.

Description of other specimens and additional data. Medusa: largest medusa 23 cm in diameter; umbrella almost hemispherical (Fig. 21), flatter in younger specimens. Exumbrella finely granulated; colouration yellowish-brown, with brown lappets, sometimes with red-brown radial stripes (Fig. 23). Mesoglea flexible, thicker centrally. Marginal lappets nearly rounded, 4–8 per octant; rhopalar lappets not overlapping (“open rhopalia” condition). Rhopalia 8, without ocelli, in deep clefts; deep exumbrellar sensory pit. Tentacle clefts vary in depth. Tentacles up to 56 (3–7 per octant); arranged as 4th-3rd-2nd-1st-2nd-3rd-4th (primary tentacle central; secondary, tertiary and quaternary tentacles laterally). Musculature not distinguishable. Brachial disc circular, with four evident corners. Pillars evident, delimited by insertion corners of manubrium. Quadralinga absent. Subgenital ostia rounded, ca. 1/3–1/5 of umbrellar diameter. Oral arms 1–4 times longer than umbrellar diameter, V-shaped, delicate and convoluted edges. Central stomach circular, marginal region limited by insertion of radial septa. Stomach pouches 16, uniform in width centrally; tentacular pouches slightly enlarged distally. Septa narrow, widening gradually (pear-shaped outline) at proximal end; outer 1/3 to 1/4 bending towards rhopalia (~45º) (Fig. 22). Gastric filaments in four interradial fields. Gonads outlining gastric filaments, semicircular ring, greatly folded; non-hermaphroditic. No data on planula, scyphistoma, or ephyrae. Cnidome ( Fig. 87): Specimen NNM 5361, medusa tentacles with large holotrichous O-isorhiza [n=10; 9.8–11.7 x 8.8–9.8 µm (mean = 10.78 x 9.21 µm)]; small holotrichous a-isorhizas [n=10; 5.8–6.8 x 2.9–3.9 µm (mean = 6.17 x 3.52 µm)]; holotrichous A-isorhizas [n=10; 15.6–18.6 x 8.8–11.7 µm (mean = 17.15 x 9.90 µm)]; heterotrichous microbasic rhopaloids [n=10; 10.7–13.7 x 5.8–6.8 µm (mean = 11.76 x 6.17 µm)]; Specimen ZMH C7451, medusa tentacles with medium holotrichous O-isorhizas [n=3; 10.7–12.7 x 10.7 µm (mean = 11.76 x 10.7 µm)]; large holotrichous O-isorhizas [n=10; 16.6–20.5 x 14.7–16.6 µm (mean = 19.21 x 15.78 µm)]; small holotrichous a-isorhizas [n=10; 5.8–6.8 x 2.9–3.9 µm (mean = 6.27 x 3.13 µm)]; holotrichous A-isorhizas [n=10; 15.6–19.6 x 9.8–10.7 µm (mean = 17.84 x 10.29 µm)]; heterotrichous microbasic rhopaloids [n=10; 10.7–13.7 x 6.8–7.8 µm (mean = 12.15 x 6.96 µm)].

Systematic remarks. We agree with Stiasny (1939) in considering Dactylometra africana as just an older stage of Chrysaora fulgida with more tentacles. Chrysaora fulgida is morphologically very similar to C. hysoscella . The two species may be distinguished by the colour pattern ( C. fulgida is usually darker), the additional tentacles appearing in the larger (ca. 20 cm) specimens of C. fulgida , and the relatively separate geographic distributions. In addition, none of the examined specimens of C. fulgida had sperm sacs on the gastrovascular cavity, as seen in specimens of C. hysoscella . We prefer to keep the species as valid until detailed data on comparative development are gathered for both species, as well as sample specimens from intermediate areas in relation to the present known distributions (e.g. Morocco coast); additionally genetic data could give some hints on the identity and distribution boundaries of the species. Chrysaora fulgida might present the same tentacular arrangement as C. quinquecirrha , with five tentacles per octant and the same formula. However, these species are on different sides of the Atlantic Ocean, with C. fulgida restricted to the African coast ( Kramp 1955; 1961) and C. quinquecirrha restricted to the east coast of the USA. Additional morphological and molecular comparisons of west African Chrysaora medusae are being conducted (S. Neethling & M.J. Gibbons pers. comm. 2009).

Biological data. No available data.

Etymology. fulgida : probably refers to colouration, derived from the Greek fulgidus (= shining) ( Brown 1956).