Chrysaora hysoscella ( Linnaeus, 1767 )

Morandini, André C. & Marques, Antonio C., 2010, Revision of the genus Chrysaora Péron & Lesueur, 1810 (Cnidaria: Scyphozoa) 2464, Zootaxa 2464, pp. 1-97 : 26-30

Chrysaora hysoscella ( Linnaeus, 1767)

( Figures 29 –36, 77, 89)

Urtica marina Borlase 1758: 256–257 , Pl. XXV fig. 7–8, 11–12 (rejected for nomenclatural purposes).

Medusa hysoscella Linnaeus 1767: 1097 (brief description) [mouth of Tagus river (Tago Lusitaniae), Lisbon, Portugal]. Modeer 1791: 31 (brief description). Forbes 1848: 77 (mention).

Chrysaora aspilonota Péron & Lesueur 1810: 365 (description) [ France]. Blainville 1834: 299 (mention as Chrysaora apsilonota ). Goy 1995: 368 (description, the author published “ ipsis litteris ” a manuscript of Péron & Lesueur), Pl. on page 369 (medusa, lower right).

Chrysaora cyclonota Péron & Lesueur 1810: 365 (description) [English Channel]. Gosse 1853: 363–365, Pl. XXVII. Blainville 1834: 299 (mention). Goy 1995: 29 (mention), 368 (description, the author published “ ipsis litteris ” a manuscript of Péron & Lesueur), Pl. on page 369 (medusa, upper left).

Chrysaora Lesueur Péron & Lesueur 1810: 365 (description) [ France]. Blainville 1834: 299 (mention). Goy 1995: 366 (description, the author published “ ipsis litteris ” a manuscript of Péron & Lesueur), Pl. on page 367 (medusa).

Chrysaora spilhemigona Péron & Lesueur 1810: 365 (description) [ France]. Blainville 1834: 299 (mention). Goy 1995: 368 (description, the author published “ ipsis litteris ” a manuscript of Péron & Lesueur), Pl. on page 369 (medusa, lower left).

Chrysaora spilogona Péron & Lesueur 1810: 365 (description) [ France]. Blainville 1834: 299 (mention). Goy 1995: 29 (mention), 368 (description, the author published “ ipsis litteris ” a manuscript of Péron & Lesueur), Pl. on page 369 (medusa, upper right).

Chrysaora pleurophora Péron & Lesueur 1810: 365 (description) [ France]. Blainville 1834: 299 (mention). Goy 1995: 368 (description, the author published “ ipsis litteris ” a manuscript of Péron & Lesueur), Pl. on page 373 (medusa).

Chrysaora heptanema Péron & Lesueur 1810: 366 (description) [North Sea]. Blainville 1834: 299 (mention). Lesson 1843: 399 (description).

Chrysaora macrogona Péron & Lesueur 1810: 366 (description) [ France]. Blainville 1834: 299 (mention).

Chrysaora mediterranea Péron & Lesueur 1810: 366 (description) [Mediterranean Sea]. Eschscholtz 1829: 82 (brief description, probably C. hysoscella ) [Mediterranean Sea]. Blainville 1834: 299 (mention). Lesson 1843: 397–398 (description). Haeckel 1880: 511–512 (description), Pl. XXXI (medusa) [Mediterranean Sea]. Vanhöffen 1888: 14– 15 (description), 22 (brief description), 46 (distribution). Mayer 1910: 579 (synoptic table, = C. hysoscella ). Broch 1913: 19 (mention, discussion on the distinction from C. hysoscella ) [Algeciras Bay – Spain]. Stiasny 1931: 139 (mention) [Naples – Italy]. G. Chapman 1953: 155 (list), 158 (mention), 161–162 (mesoglea), fig. 3 (mesoglea) [ UK]. Vannucci 1954: 125 ( C. mediterranea = C. hysoscella ). Goy 1995: 368 (description, the author published “ ipsis litteris ” a manuscript of Péron & Lesueur). Calder 2009: 24 (junior subjective synonym).

Medusa fusca Pennant 1812: 121 (description). Forbes 1848: 77 (mention, = C. hysoscella ). Medusa tuberculata Pennant 1812: 122 (description). Forbes 1848: 77 (mention, = C. hysoscella ). Cyanea le Sueur : Lamarck 1816: 519–520 (brief description).

Cyanea aspilonota: Lamarck 1816: 520 (brief description).

Cyanea cyclonota: Lamarck 1816: 520 (brief description). Goy 1995: 29 (mention).

Cyanea mediterranea: Lamarck 1816: 520 (brief description). Risso 1826: 302 (brief description).

Cyanea pleurophora: Lamarck 1816: 520 (brief description).

Cyanea punctulata Lamarck 1816: 520 (description).

Cyanea heptanema: Lamarck 1816: 521 (brief description).

Cyanea macrogona: Lamarck 1816: 521 (brief description).

Aurellia (?) crenata Chamisso & Eysenhardt 1821: 359 (description), Pl. XXIX figs a–c (medusa) [North Sea]. Blainville 1834: 299 (mention).

Chrysaora hysoscella: Eschscholtz 1829: 79–81 (description), Pl. VII, fig. 2 (medusa) [North Sea, West coast of Europe]. Blainville 1834: 299 (mention), Pl. XLIII. Brandt 1835: 227 (mention). Lesson 1843: 396–397 (description). L. Agassiz 1862: 165 (mention). Claus 1877: 33–42 (development), Pl. I figs 1–11, Pl. VI figs 29–30, Pl. VII figs 32–36’, Pl. IX fig. 40 2, Pl. XI fig. 50 [Adriatic Sea, Mediterranean]. Vanhöffen 1906: 47–48 (brief description), fig. 9 (medusa), fig. 10 (ephyra) [North Sea]. Vanhöffen 1908: 39 (brief description) [North Sea]. Mayer 1910: 579– 581 (synoptic table, description). Stiasny 1919: 72–73 (mention). Stiasny 1927: 73–85 (description, colour pattern), Pls I–III (medusa, aboral view). Teissier 1929: 137–177 (embryonic development, chemical composition of planulae), fig. 1 (gonad), figs 2, 7–9 (planula), figs 3–5 (segmentation), fig. 6 (gastrulation). Stiasny 1931: 139 (mention) [ Great Britain]. Kramp 1934: 211–220 (occurrence) [ Denmark]. Papenfuss 1936: 19–23 (nematocysts), figs 8, 13, 17, 22 [Kristineberg – Sweden]. Stiasny 1937: 275, 279 (mention). Ranson 1945: 316 (mention). Vannucci 1954: 125 (commented that C. hysoscella is identical to C. mediterranea , to C. blossevillei , to C. fulgida , and to C. plocamia ). Kramp 1955: 296 (brief description), 299 (nematocyst differences with D. quinquecirrha ), 305 (mention), 315 (tab. III) [ Liberia]. Vannucci 1957: 595, 596 (mention). Kramp 1961: 325–326 (synonymy). Widersten 1965: 53–55 (gonad structure, embryonic development), text-figs 8–10, Pl. II figs 3–4 (gonads). Kramp 1968: 82 (brief description) [ Liberia]. Russell 1970: 87–104 (description), text-figs 47, 54–55, Pl. III (medusa), text-fig. 48 (tentacle), textfig. 49 (oral arm), text-fig. 50 (rhopalium), text-fig. 51 (gonad), text-fig. 52, Pl. XI (colour pattern), text-fig. 53 (ephyra). Russell 1978: 3 (brief description), fig. 2 (medusa) [North Atlantic Ocean]. D.M. Chapman 1985: 618 (list), 621 (statolith), fig. 6 (spectrum of statolith crystal). Hedwig & Schäfer 1986: 114–116 (sperm structure), figs 13–19 (sperm ultrastructure) [ Helgoland – Germany]. Bingel, Avsar & Gucu 1991: 70 (occurrence) [Mersin Bay – Turkey]. Mills, Pugh, Harbison & Haddock 1996: 158 (mention). Raupp, Milde, Goerz, Plewig, Burnett & Heeger 1996: 49 (tab. 1), fig. 5 (medusa). Arai 1997: 17 (mesoglea), 32 (rhopalium), 36–37, 48 (nerves), 56 (polyp locomotion), 123– 124 (oxygen consumption), 135 (water content), 140–141, 145, 148 (gametogenesis), 150, 152 (larvae development), 163 (stolon), 164 (podocyst), 174, 177 (composition), 203–204 (feeding), 225 (classification). Heeger 1998: 99 (mention), 156 (mention, table), 252 (mention, table), 297 (brief description), 66–67 (figs 2–33, 2–34, medusa) [north Atlantic, Mediterranea]. Purcell, Malej & Benovič 1999: 248 (aggregation). Silveira & Cornelius 2000: 11 (mention, Tab. 1). Arai 2001: 76 (mention). Graham, Pagès & Hamner 2001: 204 (mention). Malej 2001: 68 (mention). Jarms 2002: 520 (brief description), figs 12.36a, c (medusa) [Mediterranean Sea]. Gershwin & Collins 2002: 128 (mention), 129 (tab. 1), 130 (mention), 143 (key), fig. 2 (phylogeny). Kokelj & Plozzer 2002: 179 (mention). Hansson, Moeslund, Kiørboe & Riisgård 2005: 121, 122 (mention, occurrence) [ Denmark]. Houghton, Doyle, Davenport & Hays 2006a: 159–170 (air monitoring technique) [ Ireland]. Houghton, Doyle, Davenport & Hays 2006b: 1239, 1241, 1242 (predation by Mola mola ) [ Ireland]. Barz & Hirche 2007: 1021–1033 (abundance, distribution) [North Sea]. Doyle, Houghton, McDevitt, Davenport & Hays 2007: 239–252 (energy density) [ Ireland]. Hays, Doyle, Houghton, Lilley, Metclafe & Righton 2008: 325–331 (diving behavior) [ Ireland]. Holst & Jarms 2007: 863– 871 (settlement preferences) [North Sea]. Houghton, Doyle, Davenport, Lilley, Wilson & Hays 2007: 1–13 (stranding specimens, seasonality) [British isles]. Arai 2009: 243 (podocysts). Colombo, Benovič, Malej, Lučić, Makovec, Onofri, Acha, Madirolas & Mianzan 2009: 108 (occurrence acoustic samples) [Mljet island, Croatia]. Purcell 2009: 25 (mention, trophic ecology).

Chrysaora lutea: Blainville 1834: 299 (mention, described as Orythia lutea Quoy & Gaimard, 1827: 175–176 , Pl. 4B, fig. 1, from Gibraltar strait) [not a Chrysaora , a rhizostome medusa]. Russell 1970: 88 (mention).

Medusa stella Dalyell 1847: 106 (mention), Pl. XV–XVII [ Scotland].

Cyanea chrysaora Milne Edwards & Haime, 1839 /1849, Pl. 47 [not seen].

Chrysaora hyoscella: Wright 1861: 357–359 , Pl. XVIII (gonads).

Chrysaora isosceles: Haeckel 1880: 513 (description, changing spelling) [Atlantic coasts of Europe: France, British Isles, Germany]. Vanhöffen 1888: 22 (brief description), 47 (distribution). Delap 1901: 25–28 (life cycle), Pl. I fig. 1 (polyp), Pl. I figs 2–3 (strobilation), Pl. I fig. 4 (ephyrae), Pl. II fig. 5 (ephyrae), Pl. II figs 6–7 (medusa) [Valencia Harbour – Ireland]. Delap & Delap 1906: 13 (mention) [Valencia Harbour – Ireland]. Stephens 1912: 58 6 (mention). Lebour 1923: 71 (list), 73–75 (food items), fig. 1 (medusa) [ UK]. Lambert 1935: 71, 74, 76–80, 82–83, 85 (cultivation problems), unnumbered figure (ephyra).

Taeniolhydra roscoffensis Hérouard, 1908 .

Chrysaora hysocella: Teissier 1926: 266–273 (water content).

Chrysaora isoceles : Holland, Davenport & East 1990: 762 (turtle food), 766–767 (tab. 4, fatty acid composition).

Crysaora hisoscella: Carli 1991: 77 (mention) [Ligurian Sea].

Crysaora hysoscella: Carli 1991: 77 (mention) [Ligurian Sea].

Chrysaora hisoscella: Scalera-Liaci 1991: 164 (occurrence) [Adriatic Sea].

? Chrysaora hysoscella: Pagès, Gili & Bouillon 1992: 48–52 (description), figs 59–61 (medusa) [Bengala current]. Arai 1997: 198 (specimens aggregation). Mianzan & Cornelius 1999: 537–538 (description), fig. 2.14 (distribution), figs 5.12a–b (medusa) [South Atlantic]. Brierley, Axelsen, Buecher, Sparks, Boyer & Gibbons 2001: 55–65 (acoustic observations) [ Namibia]. Buecher 2001: 63 (mention). Buecher, Sparks, Brierley, Boyer & Gibbons 2001: 1075– 1079 (biometry) [ Namibia]. Mills 2001: 57 (mention, blooms) [Southeastern Africa]. Sparks, Buecher, Brierley, Axelsen, Boyer & Gibbons 2001: 275–284 (abundance) [southeastern Africa]. Colombo, Mianzan & Madirolas 2003: 650, 655 (mention). Brierley, Axelsen, Boyer, Lynam, Didcock, Boyer, Sparks, Purcell & Gibbons 2004: 384– 392 (acoustic tests for abundance) [ Walvis Bay – Namibia]. Brierley, Boyer, Axelsen, Lynam, Sparks, Boyer & Gibbons 2005: 105–111 (acoustic estimation) [Benguela current]. Flynn & Gibbons 2007: 303–307 (diet, feeding) [ Walvis Bay – Namibia]. Purcell, Uye & Lo 2007: 155, 157, 166 (mention, blooms). Bayha & Graham 2009: 221 (molecular identification of polyps). Palomares & Pauly 2009: 13, tab.1 (stock), 15, tab. 2 (growth parameters) [ Namibia]. Purcell 2009: 25 (mention, trophic ecology).

Holotype specimen. Not available.

Examined material. MZUSP 760 (specimens ~7 and 11 cm in diameter, 01.viii.2002, 4% formaldehyde solution in seawater, Helgoland – Germany); NHM 1903.1.28.1 (as Chrysaora mediterranea , ~ 9 cm in diameter, v.1899, Portugal), NHM 1950.3.1.16 (as Chrysaora isosceles , specimens ~1, 1.2, and 2.5 cm in diameter, 1900, Valencia – Ireland,); NNM 24089 (~ 13 cm in diameter, 3.ix.1999, formaldehyde solution, The Netherlands); PMJ Coel 891 (specimens ~4, 4.5, 5, 6, and 6.5 cm in diameter, ethanol), PMJ Coel 927 (as Chrysaora mediterranea , ~ 7 cm in diameter, 1873, ethanol, Smyrna – Turkey), PMJ Coel 928 (~ 12 cm in diameter, ethanol), PMJ Coel 937 (as Chrysaora mediterranea , specimens ~12 and 13.5 cm in diameter, 1873, ethanol, Istanbul – Turkey); ZMA 2003 (as Chrysaora mediterranea , ~ 9 cm in diameter, France); ZMB CNI 381 (as Chrysaora mediterranea , ~ 4.5 cm in diameter, Trieste – Italy); ZMUC not-numbered (as Chrysaora mediterranea , ~ 10 cm in diameter, 1879, formaldehyde solution, Istanbul – Turkey) ZMUC notnumbered (specimens ~3, 4, 4.5, 4.5, and 6 cm in diameter, 13.xi.1950, Galathea Expedition, formaldehyde solution).

Type locality. Mouth of Tagus river, Lisbon – Portugal.

Distribution. Eastern North Atlantic: North Sea, English Channel, Mediterranean Sea ( Fig. 77).

Diagnosis. Living medusae of medium size, 15–25 cm in bell diameter; marginal lappets rounded (adults), 4 per octant, rhopalar lappets slightly wider, without canals; tentacles 24 (3 per octant, 2-1-2); quadralinga absent; hermaphrodites, protandric, sperm sacs on gastrovascular cavity; colouration (adults) either transparent-white with colourless or brown lappets, dark (usually brown) radiating stripes on transparent-whitish background, light stripes on dark background.

Holotype specimen description. No holotype located, and no neotype designated.

Description of other specimens and additional data. Medusa: Umbrella diameter up to 30 cm, almost hemispherical, flattened in younger specimens. Exumbrellar surface finely granulated; transparent-whitish with or without brown triangles radially distributed ( Figs 29 –31). Mesoglea flexible, relatively thick on central portion, thinner on umbrellar edges. Marginal lappets rounded, 4 per octant; rhopalar lappets slightly wider than velar, rhopalar lappets not overlapping (“open rhopalia” condition). Rhopalia 8, without ocelli, in deep clefts; deep exumbrellar sensory pit. Tentacle clefts vary slightly in depth. Tentacles 24 (3 per octant); arranged as 2nd-1st-2nd (primary tentacle central, secondary tentacles laterally) (Fig. 32). Musculature not discernible. Brachial disc circular, with four evident corners, grooved in larger specimens. Pillars weakly evident, delimited by insertion corners of manubrium. Quadralinga absent. Subgenital ostia rounded, ca. 1/8 of umbrellar diameter. Oral arms 1–2 times longer than umbrellar diameter, V-shaped, delicate and convoluted edges, distal portion sometimes slightly spiraled. Central stomach circular, marginal region limited by insertion of radial septa. Stomach pouches 16, width uniform centrally; tentacular pouches enlarged distally. Septa narrow, widening gradually (pear-shaped or triangular outline) at proximal end; outer 1/3 to 1/4 bending towards rhopalia (~45º) (Fig. 32). Gastric filaments in four interradial fields. Gonads outlining gastric filaments, semicircular ring, greatly folded; hermaphrodite, protandric, sperm sacs on gastrovascular cavity (Fig. 33). Planula. planulae elongated, pear-shaped; whitish. Scyphistoma (Figs 34–35). Conical to goblet-shaped, up to 2.5 mm high; oral disc 1.75 mm wide; typically with 16 tentacles (14–16), length up to 5 times polyp height. Mouth cruciform, with prominent lips elevated from oral disk. Gastric septa 4, whitish to cream. Podocysts. Trapezoid, diameter 0.25–0.44 mm; yellowish brown. Strobila. Polydisc (2–6 ephyrae), pale orange, strobilation lasts about 10 days. Ephyra (Fig. 36): typically with 8 arms (lobes); marginal lappets 16, pointed; rhopalia 8, with white-yellowish concretions; cruciform mouth; diameter 1.5–2 mm just after liberation; pale orange; with one nematocyst cluster on each side of rhopalium. Cnidome ( Fig. 89): Specimen MZUSP 760, medusa tentacles with holotrichous O-isorhizas [n=10; 11.7–12.7 x 9.8–11.7 µm (mean = 12.05 x 10.39 µm)]; holotrichous a-isorhizas [n=10; 4.9–6.8 x 2.9–3.9 µm (mean = 6.07 x 3.72 µm)]; holotrichous A-isorhizas [n=10; 11.7–14.7 x 6.8–8.8 µm (mean = 13.03 x 7.45 µm)]; heterotrichous microbasic rhopaloids [n=10; 8.8– 10.7 x 4.9–6.8 µm (mean = 9.8 x 5.97 µm)]. We consider atrichous haploneme nematocysts reported by other authors ( Papenfuss 1936; Russell 1970) as O-isorhizas and a-isorhizas.

Systematic remarks. Chrysaora hysoscella is morphologically similar to C. fulgida ; their main differences are in the colour pattern, in the additional number of tentacles in C. fulgida ; and the hermaphroditic condition of C. hysoscella (see discussion of C. fulgida above). The species presents a wide variation of colour patterns (see Stiasny 1927, Pls I–III; Russell 1970: 95–96, Text-fig. 52, Pl. XI), which led to it being referred to by several different names (see synonymy; Péron & Lesueur 1810; Haeckel 1880; Goy 1995). Agassiz (1862: 165) had already pointed out that specimens from the Mediterranean, the North Sea, and Atlantic coasts of Europe should be compared. The distinction between C. hysoscella and C. plocamia is in the discussion of the later.

Biological data. Life cycle described by Delap (1901), more recent compilation of data by Russell (1970).

Etymology. hysoscella : due to characteristic exumbrella colouration pattern, with some triangular markings, resembling an isosceles triangle ( Brown 1956).