Chiropsella bronzie, Gershwin, 2006

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 : 25-36

Chiropsella bronzie , sp. nov.

Plates 4 , 6

Chiropsalmus quadrigatus . — Southcott, 1952: 272–274. — Pope, 1953: 112 [photo], implicated in death [in part]. — Southcott, 1956: 254–280, comparison with Chironex fleckeri . — Halstead, 1959 (in part): 33, 36. — Southcott and Kingston, 1959: 443–444, in part, stings. — Barnes, 1960: 993–999, stings. — Kramp, 1961: 309 [in part]. — Southcott, 1963: 49 [in part]; general medical. — Yaldwyn, 1964: 312–313, dangerous. — Barnes, 1965: 13–22, morphological details. — Cleland and Southcott, 1965: 60, 65, 70, 73, 75, 123, 126, 129, 130, 131, 153, Fig. 7, Pl. I.1, Colour Plate II bottom right; medical aspects. — Halstead, 1965: 302 (in part); p. 329 text fig. 3; 334; pl. XL–XLII; LXXVIII, dangerous. — Barnes, 1966: 307–332, morphology and sting potential. — Barnes, 1967: 115–129, venom. — Cochrane, 1968: 16; general. — Halstead, 1971: 405 [in part]; general. — Keen, 1971: 249–254, toxin analysis. — Freeman and Turner, 1972: 31–37; comparison to Chironex venom. — Brown, 1973: Distribution. — Baxter and Marr, 1974: 223–229; venom neutralized by Chironex antivenom. — Halstead, 1980: 55, in part, fig. 26; sting. — Yamaguchi and Hartwick, 1980: 11–16, external fertilization. — Sutherland, 1981: 86, but fig. pg. 87 is actually Chironex fleckeri . — Sutherland, 1983: 374–377, but fig. 26.2 is actually Chironex fleckeri . — Kinsey, 1986: 1–76, appendix; misc. notes. — Marsh and Slack­Smith, 1986: 27–28, 34, fig. 19, [in part]; sting information, comparison with Chironex . — Fenner and Williamson, 1987: 258; nematocyst inhibition. — Williamson et al., 1987: 220, sting. — Kinsey, 1988: 1–32, 94, 106; misc. notes. — Halstead et al., 1990: 47, fig.; sting. — Ming et al., 1990: fig. 10; photo taken in Australia, probably wrongly attributed to Singapore. — Halstead, 1995: 65, in part; sting. — Williamson and Burnett, 1995: 105, sensu Barnes, 1965. —Fenner and Williamson, 1996: 661; use of antivenom. — Williamson et al., 1996b: 46, 150–154, 236, 237, 259–260, 261, 262, 277, 280–283, 292, 298, 302, 405, text figs. 6.29, 6.30, plates 1.10, 6.9.

Chiropsalmus sp. — Williamson et al., 1996a: 151, table 6.4a, b, 259–260, 267, 269, 270, 276, 277, 280. — Gordon, 1998; ecophysiology. — White et al., 1998: 113, stings. — Pereira et al., 2000; pressure immobilization bandages. —Carrette, 2002; extraction and ecology of venom. — Carrette et al., 2002: 1547–1551; relationship between nematocyst ratio and prey type. — Collins, 2002: 420, figs. 1–3; molecular phylogeny, Genbank number AF358103 .

Chiropsalmus quadrigata . —Hawdon & Winkel, 1999: 1370–1371; sting comparison to Chironex [incorrect subsequent spelling].

Chiropsalmus n. sp. A Gershwin, 2005: 125, pl. 4.9B, and throughout; taxonomy and phylogeny.

The synonymy above includes only those citations I believe refer to the taxon herein described as Chiropsella bronzie gen. et sp. nov. However, not all Australian references to C. quadrigatus can be easily understood as Chiropsella bronzie , gen. et sp. nov. In trying to tease apart the literature of decades ago, in which the identifications were not always accurate, it has become clear that Chironex fleckeri and Chiropsalmus quadrigatus were often used more or less interchangeably for Australian Chirodropida . I have excluded from Chiropsella bronzie gen. et sp. nov. those references to Chiropsalmus quadrigatus in which fatal stinging occurred, as these are most likely attributable to Chironex fleckeri (e.g. Flecker, 1952; Southcott, 1952; Pope, 1953 [in part]; Pope, 1957).

Material examined

Holotype: MTQ G55274 ( JHB J1217 ), Pretty Beach, QLD, Australia, coll. by J.H. Barnes 11 February 1964; gravid female, BH 67.77mm, DBW 88.60mm, IRW 43.11mm, TBW 1.20mm.

Paratypes: MTQ G55275 ( JHB J992.9 ), in fish trap, Buchan Point, QLD, Australia, 8 April 1962 ; BH 71.75mm, DBW 111.84mm, IRW 56.12mm. SAM H972 ( RVS A392 ), Bells Beach , Cairns, QLD, Australia, coll. G. Rowell 20 February 1959 ; 65.95mm BH, 102.45mm DBW, 52.65mm IRW, 7 tentacle stage. SAM H973 ( RVS A433 ), same data as H972 ; 57.16mm BH, 91.68mm DBW, 50.74mm IRW, 7 tentacle stage. SAM H974 ( RVS A429 ), Palm Beach , Cairns, QLD, Australia, coll. J.H. Barnes 22 February 1959 ; damaged, 44.94mm BH, 5 tentacle stage. SAM H975 ( RVS A397­8 ), Bells Beach , Cairns, QLD, Australia, coll. K. de Witte 14 January 1959 ; 2 specimens: A) 39.14mm BH, 52.60mm DBW, 5 tentacle stage; B) 46.20mm BH, 58.45mm DBW, 7 tentacle stage. SAM H976 ( RVS A371 ), Simpsons Point, QLD, Australia, coll. J.H. Barnes 22 February 1959 ; 67.76mm BH, 83.70mm DBW, 40.71mm IRW, 7 tentacle stage. SAM H977 ( RVS A422 ), Bells Beach , Cairns, QLD, Australia, coll. J.H. Barnes 9 January 1960 ; 37.96mm BH, 46.45mm DBW, 24.94mm IRW; 6 tentacle stage. SAM H978 ( RVS A420 ), same data as H977 ; 6 specimens, 20.11–23.31mm BH, 29.38–32.29mm DBW, 3–4 tentacle stage. SAM H979 ( RVS A421 ), same data as H977 ; 4 specimens, damaged, 5 tentacle stage. SAM H980 ( RVS A434 ), 33.57mm BH, 32.33mm DBW, 14.92mm IRW, 4 tentacle stage. SAM H981 ( RVS A396 ), Mossman Beach, QLD, Australia, coll. G. Rowell 22 February 1959 ; damaged, 54.59mm BH, 6 tentacle stage. SAM H983 ( RVS A393 ), Bells Beach , QLD, Australia, coll. G. Rowell 21 February 1959 ; 29.29mm BH, 28.96+mm DBW, 16.76mm IRW, 4 tentacle stage. SAM H984 ( RVS A395 ), same data as H981 ; damaged, 6 tentacle stage. SAM H985 ( RVS A397 a), same data as H975 ; 29.14mm BH, 41.35mm DBW, 22.58mm IRW, 5 tentacle stage. SAM H1057 , Cooktown , QLD, Australia, coll. J. & G. Seymour 24 December 1997 , numerous juvenile specimens of assorted sizes. SAM H1058 , 4 ­ Mile Beach , Port Douglas, QLD, Australia, coll. by J. Seymour and Gordon 28 December 1997 ; numerous juvenile specimens of assorted sizes. SAM H1048 , Palm Cove , Cairns, QLD, Australia, coll. J. & G. Seymour, 25 December 1997 ; 6 juv. specimens, 21.10–ca. 6mm BH. NTM C12749, 4 Mile Beach , Port Douglas, QLD, coll. J. Seymour 4 December 1997 ; 62.35mm BH, 92.53mm DBW, 45.47mm IRW, 7 tentacle stage. NTM C12750, Buchan Point , Cairns, QLD, Australia, coll. J. Seymour 9 May 1997 ; 81.90mm BH, 116.09mm DBW, 56.41mm IRW, 7 tentacle stage. AM G16004, same data as NTM C12749 ; 54.91mm BH, 65.65mm DBW, 33.23mm IRW, 6–7 tentacle stage. AM G16027, same data as NTM C12750 ; 64.10mm BH, 88.54mm DBW, 43.07mm IRW, 7 tentacle stage. WAM Z2926 , same data as NTM C12749 ; 60.22mm BH, 85.99mm DBW, 45.35mm IRW, 5 tentacle stage. WAM Z2927 , same data as NTM C12750 ; 56.84mm BH, 90.86mm DBW, 7 tentacle stage. USNM 100351 , same data as NTM C12750. QM G317056 , Port Douglas Marina, Port Douglas, QLD , Australia, at surface by Quicksilver mooring, coll. by B. Kilpatrick, 3 February ; 22 specimens, growth series ca. 15–50mm BH. QM G317057 , same data as G317056 ; 39 specimens, growth series ca. 12–52mm BH. QM G317069 , Ellis Beach , QLD, Australia, north of stinger enclosure, coll. by L. Gershwin and W. Porche 13 February 2000 ; 5 specimens approximately 27–50mm BH. SAM H1065 ( RVS A428 ), Palm Beach, QLD, Australia, 22 Feb 1959 ; ca 85mm. NHM 1964.5.11.1–2, Bell’s Beach , 4m South of Daintree River, coll. K. Gillett; 40mm BH, 51.81mm DBW, 25.22mm IRW, 5 tentacle stage .

Additional material examined: SAM H982 ( RVS A446 ) , Cairns, QLD, Australia, coll. J.H. Barnes ~ December 1960; considerably damaged .

Type locality Pretty Beach, Queensland, Australia.

Etymology

The specific name is to honour the men and women of Surf Life Saving, on this the eve of the centenary of Life Saving in Australia. The ‘Bronze Medallion’ is the hallmark of Life Saving, and is here used in the affectionate and familiar sense, ‘bronzie’ (noun in apposition).

Diagnosis

Chiropsella with 8 conspicuous solid, knob­like gastric saccules in the upper adradii of the subumbrellar cavity; with well­developed lateral gonads spanning the entire interradial septum length; with up to 9 fine, round tentacles per pedalium.

Description

Bell ( Plate 4A, B, F ) small and rounded, cuboid overall, typically less than 80mm high and wide, with a shallow subapical coronal groove defining a thick dome of apical jelly; exumbrella smooth, lacking nematocysts or warts. Interradial furrows well marked, dividing the central portion of the interradial pillars, reaching neither the subapical furrow nor the pedalia. Adradial furrows nearly vertical, accentuating the pillars as defined from the perradial region; well defined throughout the height of the bell from the subapical furrow to below the rhopalia, disappearing near the velarium.

Rhopalial niche very slightly raised from bell wall. Rhopalial niches approximately ¾ down from apex of bell, with dome­shaped opening, flat orally, creating a 90 corner with the exumbrellar surface; aboral edge of opening convex, interrupted by small median tongue­like flap hanging downward into ostium protecting rhopalium ( Plate 4C ). Rhopalial horns lacking. Rhopalia 4, deeply set within perradial niches, with 2 large median eyes with lenses and 4 smaller lateral pigment streaks, looking in toward bell cavity. Statoliths short, rounded, rod­shaped, with straight sides. Subumbrellar rhopaliar windows flat, without convexity or concavity. Rhopaliar warts not observed.

Pedalia 4, interradial, claw­like; with uppermost branch projecting outward in the interradial axis, all other branches typically (but not always) paired, approximately dichotomously, along main flat, tapering portion of pedalium; each branch leading to a single tentacle. Pedalial main canal with knee­shaped bend ( Plate 4D ) in the proximal third, laterally compressed throughout main canal and uppermost branch; main canal bifurcating just below first branch point, each side giving rise to compressed unilateral branches, straight­sided at tentacle insertion. Tentacles rarely reaching 9 per pedalium, hollow, fine, round in cross section ( Plate 4B, F ); straight­sided at base. Tentacular nematocysts borne on raised bands of alternating sizes, with major bands (1) separated by approximately 7 alternating thicker (2) and thinner (3) bands (i.e., 1­3­2­3­2­3­2­3­1).

Velarial canals highly diverticulated, with a single main canal­root extending from the stomach in each octant, with inter­digitating non­anastomosing branches, bearing lateral lobations, completely reticulating throughout velarium such that the number of canals cannot be easily counted. Perradial lappets broadly triangular, with branches emanating off the sides as velarial canals ( Plate 4E ). Frenulum a single, simple flexible sheet of tissue, thickened at base, reaching three­fourths the distance toward velarial margin.

Gastro­gonadal pouches 4, completely separated by 4 interradial septa. Gastric phacellae 4, strongly V­shaped with angles pointing toward corners, nearly connected at the perradii; appearing as a 4­pointed star when viewed from the aboral surface ( Plate 4F ). Cirri singly rooted, short. Manubrium long, extending to about 1/3 distance to bell margin; bearing mouth with 4 short, folded lips with simple margins. Mesenteries as very narrow flap­like structures in stomach region only; extending as wide, unraised clear strips all the way to rhopalia. Lateral gonads originating in center half of interradius, growing in both vertical directions as well as laterally; along entire length of interradial septum in mature individuals.

Gastric ‘saccules’ 8, interradial; solid spherical convexities arising from body wall, smooth, lacking digitations ( Plate 4B ); typically narrowed across the perradius and covered over by gonad in mature specimens.

Observations on living specimens

Swimming behaviour: Most often in the wild I have seen individuals of all sizes swimming horizontally at the water’s surface, against the prevailing current or just randomly in still water. However, I have also caught hundreds in brief net drags in less than a meter of clear water, and not seen them until the net was pulled in, indicating that they also like to swim at some depth below the surface. The species has also been observed to rotate ¼ turn every 4–5 pulsations ( R. Hore, pers. comm. Feb. 2000).

Feeding behaviour: In captivity, they exhibit typical cubozoan feeding behaviour, i.e., catch food, sink to the bottom, ingest food, scoot along the bottom for awhile while digesting, then come back up and look for more. They thrive and grow in captivity on frequent hand feedings of live baby fish touched to the tentacles (P. Petersen, pers. comm. March 2000), but they prefer small prawns (unpublished data).

Nematocysts and sting power

In Australia, ‘ Chiropsalmus quadrigatus ’ has often been maligned for fatalities ( Southcott 1952; Flecker 1952; Pope 1953; Pope 1957). But not all references to the Australian Chiropsalmus quadrigatus are referable to Chiropsella bronzie . In fact, there is no evidence that C. bronzie can deliver a lethal sting.

Various aspects of the nematocysts and venom of C. bronzie have been investigated ( Barnes 1967; Keen 1971; Freeman and Turner 1972; Baxter and Marr 1974; Carrette et al. 2002; Carrette 2002). The tentacular cnidome ( Plate 6 ) is comprised of cigar­shaped microbasic p ­mastigophores (38.90–45.98 x 9.39–10.26, n=6), large football­shaped isorhizas (20.77–24.07 x 11.76–13.77, n=7), small spherical p ­rhopaloids (9.20–10.09 x 7.93–8.68, n=3), small oval isorhizas with a ‘beehive’ appearance of the coiled tubule (8.61–9.70 x 6.77–7.89, n=14), and small rod­shaped, straight­sided isorhizas (10.97–13.08 x 3.64–4.64, n=16).

Distribution

So far, the species is documented from Cooktown in the North to Magnetic Island (off Townsville) to the South. It is exceedingly abundant from Port Douglas to Cairns in the summer months.

Additional literature The best descriptions and figures have been given by Barnes (1965).

Remarks

This description of Chiropsella bronzie will hopefully remedy some of the confusion which has long surrounded this form.

In Australian waters, fatal stingings were originally attributed to blue­bottles ( Physalia utriculus (La Martiniéré)) or Portuguese man­o­war ( Physalia physalis (Linnaeus)) ; however, it soon became clear that a cubomedusa was responsible (for historical account see Cleland and Southcott, 1965). McNeill (in Flecker, 1952) was apparently the first to attribute the stings to Chiropsalmus quadrigatus . Later, Southcott (1956) showed that the lethal medusa, which he named Chironex fleckeri , was indeed different from Chiropsalmus quadrigatus (sensu Mayer, 1910) , and for reasons that are not currently clear, the superficially similar­appearing Australian form, herein described as Chiropsella bronzie , was left with the name Chiropsalmus quadrigatus , despite the fact that it looks less like the Philippine form than does Chironex fleckeri .

To add to the confusion, certain structures have been nomenclaturally unclear. Take, for example, the structures called ‘gastric saccules’ by Kramp (1961) for other Chirodropida , ‘superior gonad’ by Southcott (1956) in describing Chironex fleckeri , and ‘perradial nuclei’ by Barnes (1965; 1966) in differentiating C. fleckeri from Chiropsella bronzie (as Chiropsalmus quadrigatus ). Part of the confusion no doubt arose because the gastric swellings in Chiropsella bronzie are solid, not hollow, as in all other species, so the term ‘saccules’ is misleading, and because they are not actually perradial (they straddle the perradii, but the structures are adradial).

The name Chiropsalmus quadrigatus was first applied to an Australian jellyfish by McNeill as cited in Flecker (1952), for a medusa responsible for the death of a boy in Darwin in 1938. The specimen in question was poorly preserved and badly mutilated, but Southcott (1956) believed it to be referable to Chironex fleckeri . When the existence of two distinct, sympatric forms became apparent, the name Chiropsalmus quadrigatus was applied to the ‘non­ Chironex ’ form. However, doubt as to the accuracy of this designation was expressed by Southcott (1956) and Barnes (1966). Indeed, Mayer (1910; 1915) redescribed Chiropsalmus quadrigatus from the Philippines as having cock’s­combshaped gastric saccules, more similar in form to those of Chironex fleckeri than to Chiropsella bronzie .

Chiropsella bronzie is easily distinguished in the field from other species in the unbranched­saccule forms (i.e., the Chiropsalmidae ) as follows: From Chiropsoides buitendijki (which has long finger­shaped gastric saccules) and C. quadrigatus , which have pedalia branching in a single plane, whereas they branch bilaterally in Chiropsella bronzie ; from C. quadrumanus , which has exumbrellar nematocysts and finger­shaped gastric saccules; and from C. zygonema , which has long, blade­like pedalia with only 2 asymmetrical fingers and tentacles.

Figures

PLATE 4. Chiropsella bronzie, gen. et sp. nov. A. Holotype, preserved. B. In life (non­type specimen); note gastric saccules (arrow). C. Rhopalial niche ostium, note ‘tag’ on upper covering scale (arrow). D. Pedalium, with knee­shaped canal bend (arrow). E. Velarium, note numerous branched canals (v) and smooth perradial lappet (l); frenulum is visible as a whitish shadow up the middle of the lappet. F. Phacellae (arrow).

PLATE 6. Chiropsella bronzie, gen. et sp. nov., tentacular nematocysts. A. Cigar­shaped microbasic p­mastigophore (a), small oval ‘beehive’ isorhizas (b), and rod­shaped isorhizas (c). B. Cigar­shaped microbasic p­mastigophore (a), small spherical p­rhopaloid (b), and small rod­shaped isorhizas (c). C. Small spherical p­rhopaloid (a), small oval ‘beehive’ isorhizas (b), and rod­shaped isorhizas (c).