Tamoya ohboya

Description. Morphology & Anatomy: Bell translucent, densely covered in conspicuously raised nematocyst warts (Fig. 1 & 3 A). Pedalia covered with nematocyst warts, both on outer keel and inner wing; pedalial wing scalpel-shaped as in Tamoya haplonema. Pedalial canal bend with prominent spike (Fig. 3 B). Tentacles with characteristic bands, red or orange to brown (Fig. 1); tentacles flared at their bases, round in cross section (pictures indicate that in life tentacles are flattened when extended). Perradial gonads as leaf-like sheets. Stomach extending to about 1 / 3 of bell height into subumbrellar cavity (Fig. 3 C); stomach attached to subumbrella with four well-developed mesenterial bands (Fig. 3 D) extending about half-way between apex and velarial turn-over. Manubrium cruciform, without nematocyst warts; lips of square shape with nematocyst warts and smooth edges (Fig. 3 C). Rhopaliar niche opening raised, frown-shaped, with single upper and lower covering scales; rhopaliar horns lacking (Fig. 3 E). Rhopalia with two median lens eyes and four lateral eyespots, two pit and two slit (Fig. 3 I & J); statolith oval (Fig. 3 J). Velarium broad; numerous canals per octant, growing close to each other with diverticula mainly at distal ends of canals (Fig. 3 F). Velarium sprinkled on exumbrellar side with many white nematocyst warts (Fig. 3 F). Frenulae four, each consisting of two sheets bracing the right-angle connection from tip of rhopalial niche to about 3 / 4 distance between velarial turnover and its margin (Fig. 3 G); perradial lappets absent. Gastric phacellae forming four bands, one in each perradius, running vertically from top of stomach down to sphincter of stomach (Fig. 3 H). Gastric cirri numerous with branches arising tree-like from each stalk (Fig. 3 K). Cnidome: The following types of nematocysts are present in T. ohboya n. sp.: club-shaped macrobasic peuryteles; lemon-shaped microbasic birhopaloids; small oval isorhizas; spherical holotrichous isorhizas; and small, oval amastigophores. Small oval isorhizas are found in all body parts except phacellae; spherical holotrichous isorhizas in all parts but apex warts. Birhopaloids are located in tentacles, manubrium and phacellae and all of them probably belong to the same type (i. e., p-birhopaloid); v-shaped notch was indiscernible in tentacles (due to image resolution) and phacellae (all nematocysts discharged) but otherwise all birhopaloids appear morphologically very similar under light microscopy. Macrobasic p-euryteles in the tentacle tip are of two types (A and B) differing in shape and size of the lancet (Fig. 4 A, C). Spherical holotrichous isorhizas in the tentacle tip belong to two distinct size classes (i. e., L and S), but do not appear to differ in morphology (Fig. 4 G – l). The size ranges and distribution of nematocyst types across different regions of T. ohboya n. sp. are presented in Table 1. Genetic Data: No variation in mitochondrial 16 S is observed between the holotype (USNM 1146080) and paratype (USNM 1124022) of T. ohboya n. sp. Mitochondrial 16 S from T. ohboya n. sp. differs from that of T. haplonema from Brazil by 2.1 – 2.5 % and T. sp. from the southeastern USA by 10.7 – 10.9 %. Mitochondrial COI from the two samples of T. ohboya n. sp. differ by 0.4 %, whereas they differ from COI from T. haplonema from Brazil by 4.4 – 4.5 %. Differential diagnosis. Tamoya ohboya n. sp. is very similar to both T. haplonema from Brazil and T. sp. from the southeastern USA in gross morphology. All three species possess a deep stomach that reaches far into the subumbrellar cavity and is connected to the subumbrella by four mesenterial filaments. All three also share the arrangement of the gastric cirri as vertical bands along the stomach walls and similar frown-shaped rhopaliar niche ostia that lack the rhopaliar horns characteristic of the genera that used to be classified within Tamoyidae (sensu Gershwin & Alderslade 2005; Carukia, Malo, Gerongia, and Morbakka). These latter taxa have recently been united in the family Carukiidae (Bentlage et al. 2010). Tamoya ohboya n. sp. can most readily be distinguished from other species of Tamoya by the distinctive red to orange banded tentacles (but see remarks below). The overall cnidome of Tamoya ohboya n. sp. is also similar to those of T. haplonema from Brazil and T. sp. from the southeastern USA. However, the distribution of nematocyst types differs among the three species and T. ohboya n. sp. possesses more nematocyst types and displays more variation across all body parts (Table 1; Figs. 4 – 6). The absence of small oval amastigophores appears to differentiate T. haplonema from Brazil from other species of Tamoya, whereas small oval isorhizas were not observed in T. sp. from the Gulf of Mexico. Tamoya ohboya n. sp. appears to be the only known species of Tamoya to possess lemon-shaped microbasic birhopaloids and small oval isorhizas in its tentacles. In contrast to T. haplonema from Brazil, no lemon-shaped microbasic birhopaloids are present in its pedalial warts. No such warts were observed in T. sp. from the Gulf of Mexico so it is unknown if this character can differentiate the two species.

TABLE 1. Cnidome comparison of Tamoya specimens. L, W denote capsule lengths and widths, respectively, in μ.? indicates uncertainty of identification. continued next page FIGURE 4. Part one of the cnidome of Tamoya ohboya n. sp. A – I. Tentacle tip. J – P. Tentacle base. A and B. Club-shaped macrobasic p-eurytele A. C. Club-shaped macrobasic p-eurytele B. D and E. Lemon-shaped microbasic birhopaloid. F. Small oval isorhiza. G. Spherical holotrichous isorhiza L. H and I. Spherical holotrichous isorhiza S. J and K. Club-shaped macrobasic p-eurytele. L. Lemonshaped microbasic birhopaloid. M and N. Small oval isorhiza. O and P. Spherical holotrichous isorhiza. All images from paratype USNM 1124022. All scale bars represent 10 μ.

Diagnosis. Tamoya with distinctly dark brown to reddish-orange banded tentacles. Tamoya with lemon-shaped, microbasic birhopaloids and small, oval isorhizas present in cnidome of tentacles.

Remarks. One of us (WG) has been accumulating sightings of Tamoya ohboya since 2001 and has also had an opportunity to swim with the species in nature. Roughly 50 confirmed sightings, mostly by amateur naturalists, of the species between 1989 and the present have been recorded. Of the 50 tabulated sightings, 26 (including collection of the holotype and two paratypes) involve pictures or videos of the specimen that clearly show the body shape and banded tentacles. Many of these have been accumulated on the online photo sharing service Flickr (http: // www. flickr. com / photos / tesserazoa / tags / bonairebandedboxjelly /). The rest were confirmed by description and by having individuals that reported encounters examine available pictures presumed to be T. ohboya n. sp. The most distinctive feature that can be observed in photos is the prominent banding pattern of the tentacles. Occasional specimens identified as T. haplonema in Brazil also display similar tentacle bands. It is unclear if specimens of true T. haplonema sometimes have banded tentacles or whether T. haplonema and T. ohboya n. sp. have overlapping ranges in Brazil. Further remarks are limited to information garnered from the tabulated 50 sightings. Very little is known about the ecology of Tamoya ohboya n. sp., but it appears to be solitary since it has most often been encountered alone (49 of 50). The species may forage for prey during the day, given that nearly all sightings (49 of 50) have been during the day. Nevertheless, more people are in the water during this time and so it would be premature to conclude from these limited observations that T. ohboya n. sp. is a daytime predator. We also know little about the ecology of its congener, Tamoya haplonema, but the information gathered thus far suggests that it feeds during the night (Nogueira Jr. & Haddad 2008). Several Tamoya ohboya n. sp. individuals have been observed swimming back and forth between the surface and the bottom in shallow water, but the significance of this behavior remains unstudied. Specimens are reported to be fast swimmers. The vast majority of the 50 occurrences have been reported during July, August, and September (8, 14, and 9, respectively), though the species has been seen in all months of the year. Tamoya ohboya n. sp. is believed to cause serious envenomations to humans, although just three putative stings have been documented. The best-documented envenomation occurred 13 August 2006, when a 13 - to 15 - year-old girl was stung at Bachelor's Beach, Bonaire. Identity of T. ohboya n. sp. as the stinger was easily confirmed because the girl's father threw the jellyfish onto a nearby jetty after which it was photographed and collected (under the permit referred to in the introduction above; http: // blog. richterscale. org / index. php / weblog / unknown _ box _ jellyfish _ species /). This specimen was preserved and an attempt was made to bring it to the US for study, but it was lost for some time and not delivered to the Smithsonian until June, 2009 (USNM 1146084; kindly provided by Ned Deloach). The envenomation resulted in pain and agitation, and the girl received hospitalization involving compression wrap and antibiotic / analgesic cream. However, the condition of the patient worsened resulting in medical evacuation to Curaçao. Two other stings have been attributed to the Bonaire banded box jelly. One was in Bonaire and involved the daughter of an individual who is familiar with the species. This envenomation resulted in severe pain and skin damage. The victim was treated with StingThing (Remedy Marketing, Inc.) with proteolytic enzymes, and hospitalization was not needed. The third sting attributed to T. ohboya n. sp. took place on St. Lucia. The victim described the banding pattern of the tentacles and confirmed identity by viewing pictures of the species. We know little about the severity of this sting.

Distribution. Confirmed sightings have been made in shallow (<10 m) near-shore waters off Bonaire (Netherlands Antilles), St. Lucia (Lesser Antilles), St. Vincent (Lesser Antilles), Cozumel (Mexico) and Utila (Honduras), suggesting that the species has a widespread distribution across the Caribbean Sea. A picture taken from Fernando de Noronha archipelago suggests that its range may extend to northern Brazil. Of the 50 tabulated sightings of T. ohboya n. sp., 45 have been on the western margin of Bonaire.

Etymology. The specific name, ohboya, has been chosen by the public through the following process. Individuals were permitted to submit potential names and reasons for them via the Internet between 14 May and 14 June 2009 (see http: // www. yearofscience 2009. org / jellyfish). Seven potential epithets were then selected from all submitted names (n = 309) by the authors of this paper. Choice among these seven names was then tabulated with an Internet poll of more than 830 votes cast from 18 – 26 June, 2009. The winning entry, ohboya, was submitted by Lisa Peck, a high school marine biology teacher who provided the following reason for submitting the name: “ I bet " Oh Boy " is the first thing said when a biologist or layman encounters the Bonaire Banded Box Jellyfish. It is not often that a new amazing species is discovered, especially one so beautiful. I am confident that Bud Gillan exclaimed " Oh Boy " when he saw tourist Vicki Carr's video of the Bonaire Banded Box Jellyfish (a never before seen species of jellyfish). Vicki probably exclaimed " Oh Boy " when Bud called her to tell her the news of her amazing discovery. I am sure that " Oh Boy " was uttered by the three people who were stung by the Bonaire Banded Box Jellyfish. In addition, I bet that " Oh Boy " was spoken by the doctors when they saw the wounds inflicted by the nematocysts of the Bonaire Banded Box Jellyfish. Later, the doctors probably uttered " Oh Boy " when they were confident that these 3 people would survive their encounter with the Bonaire Banded Box Jellyfish. And finally, when I saw the website Year of Science: Species naming contest of the Bonaire Banded Box Jellyfish I loudly exclaimed " Oh Boy " !! Therefore I am suggesting the scientific name Tamoya ohboya be given to this amazing creature. ”

Material examined. Holotype: USNM 1146080, sex undetermined, 61 mm BH (bell height; measured from velarial turn-over to top of bell), 29 mm IRW (interradial bell width), collected 30 June 2010 at Divi Flamingo, Bonaire, Netherlands (~ 12 ° 8.61 ” N, 68 ° 16.60) by W. Gillan and M. Wilhemus. Paratype: USNM 1146084, sex undetermined, 67 mm BH, 32 mm IRW, collected 15 August 2006 at Windsock, Bonaire, Netherlands (~ 12 ° 8.03 ” N, 68 ° 16.98) by J. Richter, T. Dace and P. Keyes. Paratype: USNM 1124022, mature male, 63 mm BH, 30 IRW, collected 29 July 2008 mid-afternoon, at or close to surface a few meters from shore at No Name Beach, Klein Bonaire, Netherlands (~ 12 ° 10.03 ’ N, 68 ° 18.03 ' W) by Johan van Blerk. Type locality. Divi Flamingo, Bonaire, Netherlands (Dutch Caribbean).