Chrysaora quinquecirrha ( Desor, 1848 )

Morandini, André C. & Marques, Antonio C., 2010, Revision of the genus Chrysaora Péron & Lesueur, 1810 (Cnidaria: Scyphozoa) 2464, Zootaxa 2464, pp. 1-97 : 48-51

Chrysaora quinquecirrha ( Desor, 1848)

( Figures 63 –67, 82, 94)

Pelagia quinquecirrha Desor 1848: 76 (original description).

Dactylometra quinquecirra: L. Agassiz 1862: 126 , 166 (mention). A. Agassiz 1865: 48–49 (description), fig. 69 (medusa) [Naushon – USA; Bermudas]. Haeckel 1880: 518 (description) [Atlantic coast of North America]. Fewkes 1881: 173 (description), Pl. VIII fig. 14 (medusa) [Narragansett Bay – USA]. Brooks 1882: 137 (mention) [Chesapeake Bay – USA]. Fewkes 1882: 293–294 (description), figs 25–27, 38–39 (rhopalium), fig. 28 (umbrella margin) [Newport – USA]. Vanhöffen 1888: 47 (distribution). A. Agassiz & Mayer 1898: 1–6 (description), Pls I–VI, VIII (medusa), Pl. VII figs 7–9, Pl. IX fig. 19 (rhopalium), Pl. VII fig. 11 (egg), Pl. VII figs 12–13 (polyp), Pl. IX figs 16– 17, Pl. X fig. 23, Pl. XI fig. 34 (gonads), Pl. IX fig. 18, Pl. X fig. 25, Pl. XI fig. 30–31 (oral arm), Pl. IX figs 20–22, Pl. X fig. 24, Pl. XI fig. 29 (nematocysts), Pl. X figs 26–27 (tertiary tentacle), Pl. X fig. 28, Pl. XI fig. 32 (tentacle), Pl. XI fig. 33 (radial septum) [New England, Tiverton – USA]. Vanhöffen 1906: 50–51 (brief description), figs 13– 14 (medusa) [Narragansett Bay – USA]. Maas 1910: 43 (mention). Fish 1926: 127–128 (occurrence), 130 (mention) [Woods Hole – USA]. Thiel 1935: 1 (index), 4–5 (stings), 13, 15, 31 (mention), fig. 4 (medusa).

Bathyluca solaris Mayer 1900: 2–3 (description), Pl. I, fig. 1 (medusa), fig. 2 (rhopalium), [Narragansett Bay – USA]. Mayer 1910: 585 (mention as a synonym of D. quinquecirrha ).

Dactylometra quinquecirrha: Mayer 1910: 585–588 (description), Pls LXII–LXIVa, figs 371–372 (medusa). Bigelow 1913: 91 (mention). Bigelow 1914: 27 (mention). Bigelow 1926: 365 (mention). Papenfuss 1936: 14–17 (nematocysts), 24, 25 (mention), figs 7, 11, 16, 20, [Chesapeake Bay – USA]. Littleford & Truitt 1937: 426–427 (variation). Littleford 1939: 368–381 (life cycle), Pl. I (embryonic development), Pl. II (planula, primary polyps), Pl. III (polyps, ephyrae) [Chesapeake Bay – USA]. Hedgepeth 1954: 277 (mention), 278 (distribution). Uchida 1955: 15 (mention). Southcott 1956: 258 (mention). Searle 1957: 74 (mention). Southcott 1963: 55 (accidents), fig. 5b (medusa). Cleland & Southcott 1965: 128 (mention), 145–148 (occurrence, stings). Thiel 1966: 20, 21 (mention). Uchida 1970: 300 (mention).

Dactylometra quinquecirrha var. chesapeakei Papenfuss 1936: 17 (brief description), 18–19 (nematocysts), 25 (mention), figs 12, 21 [Chesapeake Bay – USA].

Chrysaora quinquecirrha: Kramp 1961: 327–328 (synonymy). Cleland & Southcott 1965: 128 (mention), 145–148 (occurrence, stings), 165 (tab. II), fig. 13 (medusa), fig. 13 (medusa). Rice & Powell 1970: 180–186 (toxin) [Chesapeake Bay – USA]. Calder 1971: 270–274 (nematocysts of scyphistoma); figs 4–6 (nematocysts) [Virginia – USA]. Calder 1972: 40–43 (life cycle), figs 1–2 (ephyra), figs 3–6 (medusa) [Virginia – USA]. Loeb 1972: 279–288 (temperature effects on strobilation) [Chesapeake Bay – USA]. Loeb 1973: 144–146 (light influence on strobilation) [Chesapeake Bay – USA]. Loeb & Blanquet 1973: 150–157 (feeding behaviour of polyps in response to amino acid), fig. 1 (polyp) [Chesapeake Bay – USA]. Calder 1974: 326–333 (strobilation in nature) [Virginia – USA]. Loeb 1974: 423–431 (dissociation of strobilation inducing factor) [Chesapeake Bay – USA]. Blanquet & Wetzel 1975: 181–191 (ultrastructure polyp), figs 1–17 (polyp) [Maryland – USA]. Cargo 1975: 146–148 (cultivation). Kraeuter & Setzler 1975: 69 (abundance) [Georgia – USA]. Loeb & Gordon 1975: 37–41 (strobilation related to iodate compounds) [Chesapeake Bay – USA]. Larson 1976: 12 (key), 15 (brief description), figs 16a–b (ephyra), figs 24a–b (medusa) [northeast coast– USA]. Lin & Zubkoff 1976: 268–277 (isozymes) [Chesapeake Bay]. Calder 1977: 13–18 (nematocysts of ephyra); figs 4–7 (nematocysts) [Virginia – USA]. Clifford & Cargo 1978: 58–60 (feeding in laboratory) [Maryland – USA]. Cargo 1979: 279–286 (larval settlement) [Chesapeake Bay – USA]. Cargo & Rabenold 1980: 20–26 (polyp asexual reproduction), fig. 1 (polyp) [Chesapeake Bay – USA]. Williamson 1985: 37–38 (mention). Hutton, DeLisle, Roberts & Hepworth 1986: 154–155 (predation of polyps over Mysidacea), fig. 1 (polyp) [Virginia – USA]. Arai 1988: 1921 (mention). Cairns et al. 1991: 12 (list), 70 (index), fig. 2 (medusa). Purcell, Creswell, Cargo & Kennedy 1991: 103–110 (ingestion of bivalve larvae) [Choptank River, Maryland – USA]. Purcell, White & Roman 1994: 263 (mention), 264, 273, 275 (predation on copepods) [Chesapeake Bay – USA]. Thuesen & Childress 1994: 96 (mention). Burnett, Bloom, Imafuku, Houck, Vanucci, Aurelian & Morris 1996: 1378 (mention). Houck, Lipsky, Marzella & Burnett 1996: 771–778 (toxin action on rat hepatocytes) [Maryland – USA]. Olesen, Purcell & Stoecker 1996: 149–158 (feeding and growth of ephyrae) [Chesapeake Bay – USA]. Raupp, Milde, Goerz, Plewig, Burnett & Heeger 1996: 48 (mention), 49 (tab. 1), 51 (mention). Tsikhon-Lukanina, Reznichenko & Lukasheva 1996: 817 (mention). Ford, Costello, Heidelberg & Purcell 1997: 355–361 (swimming and feeding behaviour), figs 2, 4–5 (medusa) [Chesapeake Bay – USA]. Kreps, Purcell & Heidelberg 1997: 441–446 (reaction of ctenophore on medusae presence), fig. 2 (medusa) [Maryland – USA]. Wright & Purcell 1997: 332–338 (salinity effects) [Patuxent River, Maryland – USA]. Suchman & Sullivan 1998: 237–244 (predation on copepods) [Rhode Island – USA]. Purcell, Malej & Benovič 1999: 241–243 (mention), 244–248 (aggregation), 252 (biomass), 255– 258 (predation). Purcell, White, Nemazie & Wright 1999: 187–195 (environmental effects on reproduction) [Choptank River, Maryland – USA]. Silveira & Cornelius 2000: 11 (mention, Tab. 1). Arai 2001: 74, 79, 80, 81 (mention). Båmstedt, Wild & Martinussen 2001: 646, 648 (mention). Bloom, Radwan & Burnett 2001: 76–88 (comparison of toxic action with Chironex fleckeri Southcott, 1956 ) [Maryland – USA]. Buecher 2001: 63 (mention). Burnett 2001: 5 (tab. 2), 6–7 (mention). Condon, Decker & Purcell 2001: 89–94 (low oxygen effect on asexual reproduction) [Chesapeake Bay – USA]. D’Ambra, Costello & Bentivegna 2001: 226 (mention). Graham, Pagès & Hamner 2001: 201, 204 (mention). Johnson, Perry & Burke 2001: 213, 215 (mention, patterns of distribution). Kremer 2001: 88 (mention). Lalana, Ortiz & Varela 2001: 161 (mention). Lucas 2001: 234 (mention). Martinussen & Båmstedt 2001: 21, 27, 28 (mention). Matanoski, Hood & Purcell 2001: 191–199 (effect of prey on swimming and feeding efficiency) [Chesapeake Bay – USA]. Mills 2001: 59 (mention, blooms). Finenko, Anninsky, Romanova, Abolmasova & Kideys 2001: 184 (mention). Purcell & Arai 2001: 28–29, 31, 32, 34 (mention), 36 (tab. 5). Purcell, Shiganova, Decker & Houde 2001b: 150, 152, 153, 154, 155, 156, 158, 159 (mention, ctenophore predator), 153 (tab. 6). Purcell & Sturdevant 2001: 68, 76 (mention). Raskoff 2001: 127 (mention). Watanabe & Ishii 2001: 256 (mention). Abarca- Arenas & Ulanowicz 2002: 286 (aggregation on network analysis) [Chesapeake Bay – USA]. Bullard & Hay 2002: 1457, 1462, 1464 (mention) (defense against consumers) [east coast USA]. Costello & Colin 2002: 941 (feeding preferences, mention). Gershwin & Collins 2002: 128 (mention), 129 (tab. 1), 130 (mention), 131 (absence of quadralinga), 133 (mention), 137 (mention), 141 (mention), 142 (key), fig. 2 (phylogeny). Parsons & Lalli 2002: 115 (tab. 1, aggregations). Purcell 2003: 148, 149 (mention). Ishikawa, Vucenik, Shamsuddin, Niculescu & Burnett 2004: 895–899 (venom action) [Maryland – USA]. Barz & Hirche 2005: 473, 474 (mention). Dabiri, Colin, Costello & Gharib 2005: 1257 (mention). Grove & Breitburg 2005: 185–198 (effects low dissolved oxygen on growth and reproduction) [Chesapeake Bay – USA]. Kremer 2005: 619 (mention, comparison Carbon ratio). Ma & Purcell 2005a: 216, 217, 219, 220, 221 (environmental effects over predation) [Chesapeake Bay – USA]. Ma & Purcell 2005b: 231 (mention, comparison environmental effects) [Chesapeake Bay – USA]. Purcell 2005: 463, 464, 469, 472 (mention, climate effects on blooms). Purcell & Decker 2005: 376–385 (effect of climate on predation) [Chesapeake Bay, USA]. Thuesen, Rutherford, Brommer, Garrison, Gutowska & Towanda 2005: 2476, 2477, 2479, 2480 (mesoglea oxygen concentration) [Chesapeake Bay – USA]. Bayha 2006a: 82–132 (molecular systematics), figs 3– 2, 3-3 (collection map), 3-4, 3-5, 3-6 (phylogenies) [East USA]. Breitburg & Fulford 2006: 776–784 (polyp oyster dependence) [Chesapeake Bay – USA]. Hansson 2006: 124 (mention). Kimmel, Roman & Zhang 2006: 136, 137, 139 (mention, occurrence) [Chesapeake Bay – USA]. Matanoski & Hood 2006: 595–597, 600–606 (model of swimming behavior). Richmond, Marcus, Sedlacek, Miller & Oppert 2006: 193 (mention). Titelman & Hansson 2006: 302, 303 (mention, comparison of feeding rate). Decker, Brown, Hood, Purcell, Gross, Matanoski, Bannon & Setzler-Hamilton 2007: 99–112 (distribution) [Chesapeake Bay – USA]. Purcell 2007: 184, 190–192 (environment effects on polyps). Purcell, Uye & Lo 2007: 156, 159, 160–162, 166–168 (mention, blooms). Titelman, Gandon, Goarant & Nilsen 2007: 751–754 (mention, predation). Condon & Steiberg 2008: 153–168 (predation over ctenophore) [Chesapeake Bay – USA]. Costello, Colin & Dabiri 2008: 284–285 (type of swimming movement) [Chesapeake Bay – USA]. Arai 2009: 242–244 (podocysts). Bayha & Graham 2009: 221 (molecular identification of polyps). Dawson & Hamner 2009: 199 (mention). Ishii, Ohba & Kobayashi 2008: 108, 110 (low oxygen effect comparison with Aurelia ). Liu, Lo, Purcell & Chang 2009: 248 (mention). Pitt, Welsh & Condon 2009: 136, 139, 140 (comparison assimilation efficiency). Purcell 2009: 25–27, 29–30, 32–35, 38–41, 43–44, (trophic ecology methods). Purcell, Hoover & Schwarck 2009: 146–148 (effects over strobilation) [Chesapeake Bay – USA]. Calder 2009: 10 (list), 11 (key), 24–28 (description), 29, 33, 47 (mention), fig. 8 (medusa) [southeastern USA].

Chrysaora quinquecirrha: Larson 1982: 256 (list) [Carrie Bow Cay – Belize]. Duffy, Epifanio & Fuiman 1997: 123–130 (predation over fish) [Port Aransas, Texas – USA]. Graham 2001: 97–110 (numerical increases) [Gulf of Mexico]. Segura-Puertas, Suárez-Morales & Celis 2003: 9 (list) [Gulf of Mexico] [non Chrysaora quinquecirrha ( Desor, 1848) ] probably Chrysaora lactea Eschscholtz, 1829 .

Chrysaora quinequecirrha: Pitt & Kingsford 2003: 122 (mention).

Dactylometra: Patil 1951: 132 (mention).

Chrysaora sp. : Kramp 1962: 358 (probably = C. quinquecirrha ), 359 (mention), 360 (tab. I).

? Chrysaora blossevillei: Heeger 1998: 295 (synonym) [non Chrysaora blossevillei Lesson, 1830 ].

? Dactylometra africana: Heeger 1998: 295 (synonym) [non Chrysaora africana ( Vanhöffen, 1902) ].

Holotype specimen. Not available. Althought the holotype of the junior synonym Bathyluca solaris Mayer, 1900 is available (MCZ 1481), it was not seen. It is not adequate as a neotype because it is a young specimen with incomplete development.

Examined material. CASIZ 95518 (~ 7 cm in diameter, 75% ethanol, Woods Hole , Barnstable County, Massachussetts – USA) ; MZUSP 793 (~ 2 cm in diameter, 06.v.2004, from laboratory culture, 4% fomaldehyde solution in seawater) ; USNM 33457 (~ 8 cm in diameter, viii.1913, 4% fomaldehyde solution, Plum Point , Maryland – USA) , USNM 33458 (specimens ~12, 12 and 12 cm in diameter, viii.1913, ethanol, Plum Point , Maryland – USA) , USNM 53828 (~ 11.4 cm in diameter, 12.vii.1926, 4% fomaldehyde solution, Drum Point , Chesapeake Bay, Maryland – USA) , USNM 54404 (~ 15 cm in diameter, 26.vii.1916, 4% fomaldehyde solution, Cape Chas City , Chesapeake Bay – USA) , USNM 54511 (specimens ~10 and 20 cm in diameter, 18.vii.1976, 4% fomaldehyde solution, Cape Henlopen , Delaware Bay, Delaware – USA) , USNM 56702 (~ 7.6 cm in diameter, 11.ix.1977, 4% fomaldehyde solution, Cape Henlopen , Delaware Bay, Delaware – USA) , USNM 56703 (specimens ~9, 10.5, and 12 cm in diameter, 10.viii.1977, 4% fomaldehyde solution, 37º14’N 76º02.30’W, Chesapeake Bay – USA) , USNM 56758 (specimens ~2.5, 3, 3.2, 5, and 6 cm in diameter, 04.ix.1977, 4% fomaldehyde solution, Charlestown , Rhode Island – USA) .

Type locality. Nantucket Bay, Massachusetts, east coast of USA.

Distribution. Western North Atlantic, east coast of USA ( Fig. 82).

Diagnosis. Living medusae of medium size, up to 40 cm in bell diameter; marginal lappets (adults) rounded, 4–6 per octant, rhopalar lappets slightly longer than tentacular ones, without canals; tentacles 24–40 [3–5 per octant, primary tentacle central, secondary and tertiary tentacles laterally (3-2-1-2-3)]; quadralinga absent; colouration (adults) greatly variable (transparent, whitish, with reddish-brown radial pattern).

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

Description of other specimens and additional data. Medusa: Umbrella diameter up to 40 cm (observed 2–20 cm), almost hemispherical ( Fig. 63), flattened in younger specimens. Exumbrellar surface finely granulated; transparent-whitish ( Fig. 63); whitish with triangular spots radially distributed. Mesoglea flexible, relatively thick on central portion, thinner at edge. Marginal lappets rounded, 6 per octant, without canals; rhopalar lappets slightly larger than velar, rhopalar lappets overlapping (“closed rhopalia” condition). Rhopalia 8, without ocelli, in deep clefts; deep exumbrellar sensory pit. Tentacle clefts vary in depth. Tentacles up to 40 in adults (5 per octant); arranged as primary tentacle central, secondary and tertiary tentacles laterally (3-2-1-2-3) (Fig. 65), tertiary tentacles near rhopalia; 3–4 times longer than umbrellar diameter; in some specimens slightly reddish; some specimens with additional tentacles, reaching a maximum number of 56 (7 per octant; these quaternary tentacles developing between the secondary and tertiary tentacles, i.e. 3-4-2-1-2- 4-3). Radial and ring musculature not distinguishable. Brachial disc circular. Pillars evident. Quadralinga absent. Subgenital ostia rounded, ca. 1/4–1/6 of umbrellar diameter. Oral arms up to 6 times longer than umbrellar diameter (Fig. 64), V-shaped, delicate and convoluted frilled edges, distal portion slightly spiraled. Central stomach circular, marginal region limited by insertion of radial septa. Stomach pouches 16, width uniform centrally; tentacular pouches enlarged in the 1/5 distal. Septa narrow, wider at proximal end (pearshaped); outer 1/3 bending towards rhopalia (~45º) ending straight into the margin near tentacle at rhopalar lappet (Fig. 65). Gastric filaments in four interradial fields. Gonads outlining gastric filaments, semicircular ring, greatly folded; colour whitish to pale pink. Planula. planulae elongated, pear-shaped. Scyphistoma (Fig. 66). Conical to goblet-shaped, up to 4 mm high; oral disc 1.5 mm wide; tentacles typically 16 (13–20), length up to 5 times polyp height. Mouth cruciform, with prominent lips elevated from the oral disk. Gastric septa 4. Whitish to cream. Podocysts. Trapezoid, diameter 0.35–0.45 mm; yellowish brown. Strobila. Polydisc (3–15 ephyrae), pinkish, strobilation lasting about 10 days. Ephyra (Fig. 67): typically with 8 arms (lobes); marginal lappets 16, pointed; rhopalia 8; mouth cruciform; diameter 2.5 mm just after liberation; pinkish; with one nematocyst cluster on each side of rhopalium. Cnidome ( Fig. 94): Specimen USNM 54404, medusa tentacles with holotrichous O-isorhizas [n=10; 20.5–24.5 x 17.6–21.5 µm (mean = 21.66 x 19.21 µm)]; holotrichous aisorhizas [n=10; 5.8–6.8 x 2.9–3.9 µm (mean = 6.56 x 3.33 µm)]; holotrichous A-isorhizas [n=10; 24.5–27.4 x 17.6–19.6 µm (mean = 25.68 x 18.23 µm)]; heterotrichous microbasic rhopaloids [n=10; 11.7–14.7 x 6.8–9.8 µm (mean = 12.94 x 8.33 µm)]; Specimen USNM 54511, medusa tentacles with holotrichous O-isorhizas [n=10; 16.6–19.6 x 14.7–16.6 µm (mean = 18.13 x 15.68 µm)]; holotrichous a-isorhizas [n=10; 5.8–6.8 x 2.9– 3.9 µm (mean = 6.56 x 3.82 µm)]; holotrichous A-isorhizas [n=10; 14.7–18.6 x 5.8–9.8 µm (mean = 16.86 x 8.82 µm)]; heterotrichous microbasic rhopaloids [n=10; 10.7–12.7 x 5.8–7.8 µm (mean = 11.86 x 6.76 µm)].

Systematic remarks. Chrysaora quinquecirrha is the only species of the genus occurring along the east coast of the USA ( Larson 1976). It can easily be differentiated from other northern species of Chrysaora by the presence of up to 5 tentacles per octant, with the following formula 3-2-1-2-3 ( Calder 1972). In a recent molecular study, Bayha (2006a, b) stated that Chrysaora quinquecirrha of the US east coast comprises two distinct species: C. quinquecirrha (from coastal waters) and Chrysaora sp. #1 (from estuarine, Gulf of Mexico, and Caribbean waters). There are no valid published records (see synonymy) of the species south of the state of Georgia ( USA) and, therefore, the distribution of the species is discontinuous with the other Atlantic species of the genus, C. lactea (see discussion of C. lactea above). The two species have been confused at times in the past, both having a similar number of tentacles (up to 40). However, the pattern of tentacular development in the two species is clearly different. Based on specimens observed here, we disagree with the data presented by Bayha (2006a, b) – our morphological studies have not found any distinguishing character that suggest the separation of C. quinquecirrha into two species, although further molecular data may provide new insights.

Biological data. The life cycle was described by Littleford (1939) and Calder (1972). The species has been studied extensively in terms of the effects of environmental factors on strobilation (e.g. Loeb & Gordon 1975), predation effects (e.g. Matanoski et al. 2001), and stinging ability and action of venom in comparison to other species (cubozoans) (e.g. Burnett et al. 1996; Bloom et al. 2001).

Etymology. quinquecirrha : due to the presence of five tentacles per octant in the species, derived from the Latin quinque (= 5) plus cirrus (hair, tail) ( Brown 1956).