Chiropsalmus quadrumanus ( Müller 1859 )

Gershwin, Lisa-Ann, 2006, Comments on Chiropsalmus (Cnidaria: Cubozoa: Chirodropida): a preliminary revision of the Chiropsalmidae, with descriptions of two new genera and two new species, Zootaxa 1231 (1), pp. 1-42 : 6-10

Chiropsalmus quadrumanus ( Müller 1859)

Plates 1 , 5A

Tamoya quadrumana Müller, 1859: 1–12 , pl. 2, figs. 18–25, pl. 3, figs. 26–33. — Greene, 1879: 793–794, should be retained in the genus Tamoya .

Chiropsalmus quadrumanus — Agassiz, 1862: 174. — Haeckel, 1880: 447; summary. — von Lendenfeld, 1884: 248; summary. — Conant, 1898: 4. — Mayer, 1910: 515, pl. 57, fig. 3. — Krumbach, 1925: 574; Sumatra. — Ranson, 1945: 314; French Guyana. — Ranson, 1949: 123, 137; mouth of the Amazon. — Vannucci, 1954: 120–123; lengthy description, São Paulo, Brazil. — Vannucci, 1957: 594–597; summary. — Guest, 1959: 79–83; occurrence at Texas, USA, coincident with drought conditions. — Kramp, 1959: 16–17; W. Africa and Venezuela (W. African records doubtful). — Kramp, 1961: 309–310; synopsis. — Vannucci, 1966: 662; sting effects. – Phillips et al., 1969: 708, 711; feeding ecology and crab associates. — Phillips & Burke, 1970: 853–859; ecological effects of salinity, Mississippi Sound, USA. — Burke, 1975: 37; Mississippi Sound. — Burke, 1976: 20, 24–25; Mississippi Sound, USA. — Calder & Peters, 1975: 364–369, text fig. 1; cnidome and systematic comments. — Kraeuter & Setzler, 1975: 68; ecology and occurrence at Georgia, USA. — Larson, 1982: 255–257; Belize. — Werner, 1984: 133, circumtropical. —Gómez Aguirre, 1986: 227–234; ecology and occurrence in Gulf of Mexico. — Bengston et al., 1991: 1404–1406; lethal sting, near Galveston, Texas, USA. — Cairns et al., 1991: 11; name validity, Atlantic USA. — Schnadig et al., 1991: identification via nematocysts from autopsy skin scrapings. — Marques et al., 1997: 136–138; cnidome of Brazilian population. — Morandini & Marques, 1997: 188–189; discussion of sting, occurrence from Pernambuco to Santa Catarina State, Brazil. — Mianzan & Cornelius, 1999: 533, fig. 3.5; may be conspecific with C. quadrigatus . — Fernando, 2001: 140; earlier Indian reports attributed to this species were inaccurate. — Pastorino, 2001: 357; discussion. — Cairns et al., 2002: 9; name validity, USA. — Haddad et al., 2002: 1445–1450, figs. 1 (left) and 3 (left); envenomation. — Migotto et al., 2002: 22; presence in Brazil. — Morandini et al., 2005: 281–294, fig. 3. — Gershwin, 2005: 124, pl. 4.9A and throughout; taxonomy and phylogeny.

Cheiropsalamus quadrumanus Brooks, 1882: 137–138 ; Beaufort, North Carolina, USA, taken only at depth [incorrect subsequent spelling].

Chiropsalmus quadramanus Levy, 1983: 154–155 ; field guide, first aid [incorrect subsequent spelling].

Doubtful records

Chiropsalmus quadrumanus . — Stiasny, 1926: 250; occurrence in Darwin, Northern Territory, Australia. — Rao, 1931: 28–29; occurrence in Indian Ocean. — Kramp, 1959: 16–17; W. Africa and Venezuela (W. African records doubtful). — Payne, 1960: 7; Darwin, Northern Territory, Australia. — Chakrapani, 1984: 38–40; India. — Fernando, 1992: 75–76; stings in Sri Lanka; later identified as C. quadrigatus .

Material examined

Holotype: Could not be located at any major European or American institution; apparently no longer extant.

Neotype: MZUSP 493 , 22 February 2001, collected by bottom trawl over very muddy bottom, 7–10m, Costão do Carneiro , in front of Enseada beach, Ubatuba, São Paulo State, Brazil, by A.C. Marques, preserved in 4% formaldehyde solution in seawater; 8 cm BH, 7­ tentacle stage.

Other material: Unregistered, Ubatuba, trawl net, 27 March 1996, 10– 15m; 5.5cm BH, 8.5cm DBW, female. MNRJ 184 , Baía de Traição, Paraíba, Brazil, coll. 4 June 1978, P. Duarte; 6 specimens, 2.5–3cm BH. MNRJ 931 , Praia de Itaipava , Espírito Santo, 28 April 1986; 3.5cm BH, 5.5cm DBW. MZUSP 816 (1 specimen in ethanol) and 820 (2 specimens in formalin), 22 February 2001, collected by bottom trawl over very muddy bottom, 7–10m, Costão do Carneiro , in front of Enseada beach, Ubatuba, São Paulo State, Brazil, 3 other specimens examined live with neotype, 5–6cm BH, 7­tentacle stage. RMNH 17733 , 06°09.4’N, 54°02.8’W, Surinam, 10 April 1969; 3 specimens, 4–8cm BH, up to 9­tentacle stage, with exumbrellar nematocysts, opposite pedalia .

Cautiously identify as C. quadrumanus: ZMUC unregistered, Rio de Janeiro, Brazil, 1957; 78mm BH, 91mm DBW, 45mm IRW, 1mm TBW, tentacles per pedalium: 8, 8, 7, 9, alternate; lacking bell nematocysts. NHM 2000.1799, Nova Visçosa, Bahia, Brazil, coll. 3 February 1995, A.C. Morandini (lacking bell nematocysts, alternate pedalial branching), 49mm BH, 72mm DBW, 33mm IRW, 1mm TBW. NHM unregistered, Praia de Caiobá, Paraná, Brazil, 27 May 1980 (inaccurately indicated on the specimen label as 1880); 53mm BH, 72mm DBW, 9­tentacle stage (lacking bell nematocysts, alternate pedalial branching).

Type locality

Praia de Fora , Palhoça county , Santa Catarina, Brazil; originally reported as stranded at Praia de Fora, in the county of Desterro, Santa Catarina State. According to A. Morandini (pers. comm.), Desterro (now called Florianópolis) is on Santa Catarina Island, and has no beach called Praia de Fora; however, a beach by that name exists on the nearby mainland in the county of Palhoça, which was formerly probably included in the county of Desterro, and this is likely to have been the actual type locality .

Neotype locality Ubatuba , São Paulo, Brazil.

Emended diagnosis

Chiropsalmus with exumbrellar nematocyst warts; with up to ten, oppositely­arranged tentacles on each pedalium; with short, hollow, finger­like gastric saccules.

Revised description, based on neotype

Body bell­shaped ( Plate 1A ), somewhat broader than high, to about 8cm tall; with rounded, thickened apex, lacking circumaboral groove. Interradial pillars shallow. Adradial furrows shallow in upper half, defining rhopalial region but not pillars in lower half. Exumbrella liberally sprinkled with small but conspicuous raised nematocyst warts, moderately concentrated near the velarial turnover.

Pedalia 4, interradial, about as long as bell is tall, each with about 7 gelatinous ‘fingers’ and tentacles; with right and left ‘fingers’ held somewhat tightly together; lacking nematocysts. Pedalial canals flat in cross section, with angular knee­like or shallow volcano­like bend near pedalial base, lacking upward­pointing ‘thorn’; lateral canal branches emanate from both sides of undivided main canal, right and left side branches arranged opposite to one another ( Plate 1B ); truncate­bulbous at tentacle insertion. Tentacles round to somewhat flattened in cross section, about 1mm diameter at base; tentacle banding in a 1­2­3­2­1 pattern, without other ornamentation; straight­sided at insertion point.

Rhopalial niche dome­shaped, with a single narrowly convex covering scale above, with an open horizontal shelf below; immediate rhopalial niche region is prominently raised from body wall; lacking rhopalial horns. Rhopalia with 6 eyes, 2 median with lenses and 4 lateral eye spots, the upper one rounded, the lower one linear and horizontal ( Plate 1C ). Rhopalial stem warts lacking. Statolith large, globular, hanging well below the main lensed eye rather than behind it; outline shape was not studied prior to preservation.

Velarium broad, about 1/3 to 1/2 bell radius. Velarial canals very numerous, with lateral lobations, dendritically branching from a single root in each octant, with more than 10 canals extending past velarial turnover; with numerous scattered nematocyst warts on bases of velarial canals and on perradial lappets ( Plate 1D ). Perradial lappets broadly triangular, with canals branching off sides. Frenulum a well developed, single­sheet extending a little over halfway to bell margin.

Stomach small and shallow. Gastric cirri short, unbranched, singly­rooted, bundled into discontinous, horseshoe­shaped phacellae. Gastric saccules 8, solid, distinct, fingershaped, smooth and unbranched, pendant, about 1/3 to 1/2 as long as the subumbrellar cavity ( Plate 1A, D ). Gonads leaf­like, attached along entire height of interradial septa, extending out onto gastric saccules. Manubrium very long, nearly reaching bell margin. Mouth comprising 4 broadly pointy triangular lips, with smooth edges. Perradial mesenteries completely lacking.

Color in life transparent and colourless, with conspicuous red nematocyst warts on bell and velarium; tentacles yellowish­whitish.

Variation

Specimens in the NHM collection from northern Brazil lack any trace of exumbrellar nematocysts. Müller’s three original specimens were about 100mm tall by 120mm wide, and had 7 or 8 tentacles per pedalium (as indicated in the illustrations).

Nematocysts

The cnidome of the southeastern and northeastern Brazilian populations of C. quadrumanus was studied by Marques et al. (1997), and of the South Carolina population by Calder & Peters (1975). The cnidome comprised ellipsoid and ovoid isorhizas, clubshaped microbasic mastigophores, and three sizes of oval microbasic p ­mastigophores in both populations, but the dimensions and distribution of the various nematocyst types were different. Several of these types are shown in Plate 5A from the neotype specimen.

Ecology

Kraeuter and Setzler (1975) stated that C. quadrumanus was the second most abundant species in their plankton study at Hudson Creek and Duplin River, Georgia; they further reported that a July bloom was common and that the species overwinters in the estuaries.

Distribution

Southeast Brazil to North Carolina, USA. Reports from Sumatra, Australia, and West Africa seem dubious pending examination of material. Unfortunately, the Darwin, NT, specimen ( AM #4188) identified by Stiasny (1926) cannot presently be located.

Remarks

It may well be that the various populations of what have long been thought to be a single species of Chiropsalmus quadrumanus , actually represent more than one species. Specimens from northern Brazil lacking exumbrellar nematocysts were collected from bottom trawls, and thus may have lost their warts during collection (A. Morandini, pers. comm.); however, upon examination, these specimens did not appear to have warts even in un­abraded places such as the recessed areas adjacent to the pedalia or rhopalial niches, or on the velarium. Thus, I am extremely dubious that the northern Brazilian form lacking the exumbrellar nematocysts is the same species, and if it is not, one would have to wonder about the true relationship of the forms to the north and south of it. According to the nematocyst studies of Calder and Peters (1975) and Marques et al. (1997), the nematocyst ratios and sizes are somewhat different between the north and south. It would be helpful to do a combined morphological and molecular comparison of the American and southern Brazilian forms, plus the northern Brazilian form that lacks nematocysts on the body.

Figures

PLATE 1. Chiropsalmus quadrumanus. A. Neotype, in life; note unbranched, pendant saccules (arrow). B. Pedalium; note that the central canal is undivided along the branching portion (arrow). C. Rhopalium (removed from animal). D. Subumbrellar view.

PLATE 5. Primary tentacular nematocysts. A. Chiropsalmus quadrumanus, neotype, club­form microbasic p­mastigophore (a), large oval microbasic p­mastigophore (b), spherical microbasic p­ mastigophore (c), and straight­sided isorhizas (d). B. Chiropsoides buitendijki, syntype, elongate club­form microbasic p­mastigophore above, large oval microbasic p­mastigophore below. C. Chiropsoides buitendijki, non­type specimen from Sri Lanka (QM 317021), microbasic p­ mastigophore; note ‘hockey­stick’ shape.