Revision of the genus Carybdea (Cnidaria: Cubozoa: Carybdeidae): clarifying the identity of its type species Carybdea marsupialis Author Acevedo, Melissa J. Author Straehler-Pohl, Ilka Author Morandini, André C. Author Stampar, Sergio N. Author Bentlage, Bastian Author Matsumoto, George I. Author Yanagihara, Angel Author Toshino, Sho Author Bordehore, César Author Fuentes, Verónica L. text Zootaxa 2019 2019-01-09 4543 4 515 548 journal article 27692 10.11646/zootaxa.4543.4.3 9a6e20a7-1a82-439c-bf30-07a892b110ef 1175-5326 2618013 EDFF0523-01DE-4F51-8499-ED82437EB049 Carybdea rastonii Haacke, 1886 Figs. 3 F–I Material examined: One (1) unregistered, preserved specimen, from Mirimbula ( Victoria , Australia ), collected by G. Hood, March 0 3, 2000; One (1) unregistered, preserved specimen, from Waterloo Bay ( South Australia ), collected by J. Seymour , February 1999 . Diagnosis : Gastric phacellae horizontal linear, gastric filaments with multiple roots, short stemmed; velarial canals 2 per octant, triforked; pedalial knee bend rounded, no appendage. Description: Adult medusa: Bell ( Fig. 3F ), highly transparent, colourless, slightly higher than wide, cuboid to almost cubical in shape, regularly scattered with colourless, nematocyst warts, from apex to bell margin, more dense on the bell sides than on the bell edges; nematocysts roundish to oval with different diameters, largest approx. 0.5 mm in diameter, velarium without nematocyst warts; apex plane to round convex, mesoglea thick, slight horizontal constriction near the top present. Bell height 30–35 mm high, bell width 20–30 mm (interrhopalial diameter). FIGURE 4. Schematic drawing of anatomical characters and standard measurements used (following Gershwin 2005a,b, 2006; Straehler-Pohl 2014); A) lateral view and B) subumbrella view. Gastric filaments (GF); bell (B); gonads (Go); rhopalial niche (RhN); pedalium (P); pedalial canal (PC); outer pedalial wing (oPW); inner pedalial wing (iPW); tentacle (T); bell height (BH); interrhopalial width (IRW); diagonal bell width (DBW); interpedalial diameter (IPD); interrhopalial diameter (IRD); velarial canal (VC); velarium (V); manubrium (M); stomach (Sto); mesenterium (Mes). FIGURE 5. General aspects of C. marsupialis : A) habitus live (photographer: C. Casanellas) B) preserved (Neotype: MZB 2015-1701) C) aboral view of gastric cavity D) close up of eppaulette shape phacellae E) pedalium F) pedalial knee bend G) rhopalium H) detail of the velarial canals (3 per quadrant). Photographer of different C. marsupialis individuals C-H: E. Obis. Scale bar: 1 cm. FIGURE 6. Developmental stages of C. marsupialis (preserved specimens): A) oral view, stage A, short tentacle (solid arrow) and rhopalium (dashed arrow) B) stage B starting the development of pedalia C) stage C with gastric phacellae completely developed D) stage D start the development of velarial canals E) stage E appearance of gonadal tissue F) adult stage F with ripe male gonads. Photographer: E. Obis. Scale bar: 1 mm. Pedalia ( Fig. 3H ), 4, simple, unbranched flattened, scalpel-shaped, approx. 1/3–1/2 bell height in length, situated in each interradial corner; outer wing scattered with large, white to light brown nematocyst warts or bands of nematocysts (approx. 0.5 mm in length) covering the outer keel of the pedalia; inner wing free of nematocyst warts; pedalia carrying single, white to flesh coloured tentacles in preserved specimens, in living specimens pale pink to brownish when contracted, resembling bead-chain when relaxed with white nematocyst-battery rings on a pale pink tentacle “string”. Pedalial canal slightly tapering at upper end, rounded knee bend without any hook or thorn appended, going straight through the pedalium, diamond-shaped in cross-section with sharp outer keels at proximal end up to velarium level then turning circular in cross-section towards distal end. FIGURE 7. Comparison between smaller stages of C. marsupialis (left column) and C. xaymacana (right column): A and B) general view of the small cubomedusae, 4 tentacles in C. marsupialis and 2 tentacles in C. xaymacana ; C and D) comparison of the tentacles; E and F) detail of the rhopaliar niche. Photographs taken of live specimens (with exception of E): E. Obis. Rhopalia, 4, located inside heart-shaped rhopalial niche ostium, with triangular covering scale; few small, round nematocyst warts on scale; approx. 1/6 to 1/5 of bell height up from margin; rhopalium with 6 eyes (2 median lens eyes + 2 lateral slit eyes + 2 lateral pit eyes). Velarium ( Fig. 3I ), free of nematocyst warts, containing 2 velarial canal roots per octant, canals broad in width, main canal tri-forked, square tipped, side branches sometimes bi-forked; canals flanking frenulum slightly smaller than canals flanking pedalia, all canals equally complex. Manubrium (1/2 to 3/4 of bell height in length), 4 lobes, cruciform with long, straight and blunt mouth arms,, without nematocyst warts, connected to large stomach; stomach communicates perradially with 4 gastric pockets leading into velarial canals. Gastric phacellae ( Fig. 3G ), pale pink to brownish coloured, 4, horizontal rows, in 4 stomach corners, consisting of ca. 12–15 single, short-stemmed, brush shaped filaments per quadrant. Gonads, 4 pairs narrow leaf-like- to blunt spear-head-shaped, separated by interradial septum, extending from stomach rim to bell margin, tapering towards stomach rim, tapering at rhopalia level, and broadening again towards bell margin; sexes separated but unimorph; ripe gonads yellowish to flesh coloured in preserved specimens. Remarks: Haacke (1886) gave a detailed description of a species that he had sampled in St. Vincents Gulf in Southern Australia and which he named C. rastonii . Later, the species C. arborifera , Maas 1897 and C. brevipedalia , Kishinouye 1891 were reclassified as “geographic races” ( Bigelow 1909 ) or local varieties (“lokale Varietäten”) ( Maas 1897 , 1910 ) of the same species and synonyms of C. rastonii ( Maas 1903 , 1910 ; Mayer 1906 , 1910 ; Bigelow 1938 ; Kramp 1961 ). This gave C. rastonii a wide distribution throughout the Pacific Ocean on both the eastern and western margins [ Japan , Philippines , Taiwan , Guam ( USA ), California ( USA ), Hawaii ( USA )] (e.g. Yatsu 1917 , Matsumoto 1995 , Kingsford & Mooney 2014 ). But Gershwin (2006) demonstrated that the Japanese population has a distinctive cnidome compared to C. rastonii from Australia . Moreover, Bentlage et al. (2010) showed that the populations from Hawaii and Japan were a case of several species being united under the name C. rastonii and resurrected the species C. arborifera (Hawaii) and C. brevipedalia ( Japan ) . Reported distribution of C. rastonii : Australia