ALATINIDAE Gershwin, 2005 a

Bentlage, Bastian & Lewis, Cheryl, 2012, An illustrated key and synopsis of the families and genera of carybdeid box jellyfishes (Cnidaria: Cubozoa: Carybdeida), with emphasis on the “ Irukandji family ” (Carukiidae), Journal of Natural History (J. Nat. Hist.) 46 (41 - 42), pp. 2595-2620 : 2599-2602

Family ALATINIDAE Gershwin, 2005 a

Tentacles simple................................ Genus Alatina ( Figures 2E , 3B, K )

Tentacles branching distally............................ Genus Manokia (no figure) adradial Lying halfway between perradial and interradial plane area corrugata Area of muscular tissue that displays alternating ridges and grooves diverticula Sacs or pouches stemming from velarial canals exumbrella Outer surface of the bell of a medusa frenulum Perradial sheet of muscular tissue bracing the right-angle connection of velarium and subumbrella gastric phacellae Part of the digestive system of box jellyfish; usually comprised of bundles of cirri in the corners of the stomach gonad Interradial sheet of tissue containing eggs or spermatozoa interradial In box jellyfish the planes that are marked by the presence of pedalia and tentacles manubrium Cruciform or quadrangular tube of varying length or thickness projecting downward from the stomach mesentery Perradial sheet of tissue that attaches the stomach to the subumbrella nematocyst Capsule containing an oftentimes barbed tubule that delivers venom to predator or prey. Commonly referred to as

“stinging cell” nematocyst wart Dense accumulation of numerous cells containing nematocysts; visible to the naked eye as freckles or warts on the body of

the medusa pedalium Interradial muscular structures inserting at the base of each corner of the bell. Pedalia bear the tentacles of box jellyfish.

In Carybdeida each pedalium bears a single tentacle; in Chirodropida pedalia branch into numerous “fingers” distally, each

bearing a single tentacle pedalial canal Canal in the pedalium that connects the gastro-vascular system of the bell to the hollow tentacles perradial In box jellyfish the planes that are marked by the presence of rhopalia perradial lappet Muscular, triangular lappet of tissue in the perradius of the velarium rhopaliar horn Blind-ending canal that possesses an opening to the inside of the rhopaliar niche; function unknown rhopaliar niche Cavity that is open to the environment at the exumbrellar side of the bell; contains rhopalium rhopaliar niche ostium The exumbrellar opening of the rhopaliar niche; usually bearing covering scales rhopaliar window Tissue covering the rhopaliar niche on the subumbrellar side rhopalium Sensory structure bearing eyes and statocyst. In box jellyfish rhopalia possess a muscular stalk that allows for active

movement of the rhopalium stomach Sac-like enlargement of the gastro-vascular system in the upper portion of the subumbrellar cavity subumbrella Underside of the bell of a medusa subumbrellar cavity Cavity formed by the subumbrella velarium A circular flap of muscular tissue forming the opening of the subumbrellar cavity velarial canal Canal in the velarium that is connected to the gastro-vascular system

Alatina Gershwin, 2005a (refer to Gershwin 2005a; Bentlage et al. 2010; Bentlage 2010 for reviews of the genus). Type species: Alatina mordens Gershwin, 2005a , by original designation. Species: Alatina alata ( Reynaud, 1830) nomen dubium; A. grandis (Agassiz and Mayer, 1902) ; A. madraspanata (Menon, 1930) ; A. moseri (Mayer, 1906) ; A. mordens Gershwin, 2005a = A. moseri ( Bentlage et al. 2010; Bentlage, 2010); A. obeliscus ( Haeckel, 1880) nomen dubium; A. philippina Haeckel, 1880 nomen dubium; A. pyramis ( Haeckel, 1880) nomen dubium; A. rainiensis Gershwin, 2005 a ; A. turricula ( Haeckel, 1880) nomen dubium.

Manokia Southcott, 1967 (refer to Gershwin 2005a for a review of the genus). Type species: Carybdea stiasnyi Bigelow, 1938 , by original designation. Species: Manokia stiasnyi (Bigelow, 1938) .

Geographic distribution of the family

Global; tropical to temperate; neritic, and oceanic.

Remarks

Alatinidae sensu Gershwin (2005a) contains all those Carybdeida with crescentic gastric phacellae, comprised of long cirri arranged (more or less) in a single plane (i.e. the genera Alatina and Manokia ; Figure 3K ). The rhopaliar niche ostia of Alatinidae are T-shaped (i.e. comprised of a single upper covering scale and two well-developed lower covering scales; Figure 3B ). Gershwin (2005a) also includes “3 or 4 more or less simple velarial canals per octant” in her diagnosis. This, however, provides problems for accommodating Alatina madraspanata , which clearly belongs to the family Alatinidae ( Gershwin 2005a) but possesses more than four velarial canals per octant. Therefore, velarial canals should be omitted from the diagnosis of the family.

Among the Alatinidae is A. alata , a species originally classified as Carybdea alata ( Reynaud 1830; Gershwin 2005a). Over time, numerous species were synonymized with A. alata ( Mayer 1910; Kramp 1961), but were recently considered to be separate species ( Gershwin 2005a). Molecular genetic analysis, however, suggests that at least some of the nominal Alatina species may indeed be part of a more widespread speciesgroup ( Bentlage et al. 2010). In particular, A. moseri and A. mordens are presumed to be synonymous based on molecular genetic data (the mitochondrial ribosomal 16S gene; Bentlage et al. 2010), suggesting that the characters used to distinguish the two result from intra-specific rather than inter-specific variation. The biogeography of Alatina makes it possible that additional species are part of a widespread species-group (Bentlage 2010). Therefore, the validity of many species of Alatina remains questionable and the taxonomic characters used for species delineation need to be re-evaluated in light of molecular genetic data (Bentlage 2010; Bentlage et al. 2010). In particular, a neotype for A. alata , the type species of the genus Alatina , should be designated, as this species was the earliest one described in the genus Alatina . The name A. alata is prevalent throughout the literature (usually as Carybdea alata ), but based on its original description the species is unidentifiable ( Gershwin 2005a). A redescription of specimens from the Atlantic (the type locality of A. alata ) is forthcoming, but we do not designate a neotype for A. alata here.

Figures

Figure 2. Representative species of box jellyfish from each family of Carybdeida (A–E) and one from Chirodropida (F). Note that the chirodropid possesses multiple tentacles per pedalium (F) whereas the carybdeid species possess only a single tentacle per pedalium (A–E). (A) Tamoya ohboya (Tamoyidae; image courtesy of Tim Lowry), (B) Malo kingi (Carukiidae), (C) male and female of Copula sivickisi coupling during courtship (Tripedaliidae; image courtesy of Alvaro Migotto), (D) Carybdea branchi (Carybdeidae; image courtesy of Brent Viljoen), (E) Alatina sp. (Alatinidae), (F) Chiropsalmus quadrumanus (Chiropsalmidae).sb: swimming bell, pe: pedalium, te: tentacle.

Figure 3. Key characters for identification of the Carybdeida: (A) location of taxonomic characters (Carybdea brevipedalia; image courtesy of Alvaro Migotto), (B) T-shaped rhopaliar niche ostium; note the three scales covering the niche, thus leaving a T-shaped ostium, (C) heartshaped rhopaliar niche ostium; niche ostium with only one upper covering scale, (D) key-hole shaped rhopaliar niche ostium in Copula sivickisi, (E) frown-shaped rhopaliar niche ostium in Tripedalia, (F) frown-shaped rhopaliar niche ostium in Tamoya, (G) frown-shaped rhopaliar niche ostium with rhopaliar horns in Malo, (H, I) vertical gastric phacellae in the stomach of Tamoya, (J) multiple pedalia on each corner of the swimming bell in Tripedalia, (K) crescentric gastric phacellae in Alatina (apical view). cs: covering scale, ma: manubrium, ph: gastric phacellae (comprises gastric cirri) in the stomach cavity, pc: pedalial canal, pcb: pedalial canal bend, pe: pedalium, rh: rhopalium, rn: rhopaliar niche that opens on the exumbrellar side of the bell with the rhopaliar niche ostium, rno: rhopaliar niche ostium, rho: rhopaliar horn, sto: stomach, te: tentacle, ve: velarium, vec: velarial canal.