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Home , Box jellyfish, Carybdeida, Chiropsalmus quadrumanus, Copula sivickisi, Tripedalia cystophora

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 129:164–172. 2016. First record of the box jellyfish (: Cubozoa: Tripedaliidae) in the

Robert M. Lasley Jr*, Cheryl Lewis Ames, Robert Erdman, Sheri Parks and Allen G. Collins (RML, SP) Fish and Wildlife Research Institute, Florida Fish and Wildlife Conservation Commission, St. Petersburg, Florida, U.S.A. (AC, CLA) Department of Zoology, National Museum of Natural History, Smithsonian Institution, Washington, DC, U.S.A. (CLA) Biological Sciences Graduate Program, University of Maryland, College Park, Maryland, U.S.A. (RE) Department of Biological Sciences, Florida Gulf Coast University, Fort Myers, Florida, U.S.A. (AGC) National Systematics Laboratory, NOAA Fisheries, National Museum of Natural History, Smithsonian Institution, Washington, DC, U.S.A.

Abstract.—Although observations of the box jellyfish Tripedalia cysto- phora have been widely documented since it was first recorded in Jamaica in 1897, to date there are no published reports of its occurrence in the Gulf of Mexico. Eighteen specimens of Tripedalia cystophora (Cubozoa: Tripedalii- dae), 12 and 6, respectively, were collected from a mangrove waterway near Bonita Springs, Florida, in 2010 and a brackish canal in Englewood, Florida, in 2015. Additional records from Tampa Bay (2007–2015) and Everglades City (2015), Florida, were documented with photography. Within the past few years, new records of T. cystophora from other localities, including Hawai‘i, northern Australia, and the east coast of Florida, have been reported. This study represents the first published report of the from the Gulf of Mexico. Several other species of cubozoans have been reported from the Gulf region, but the presence of three pedalia at each corner of the swimming bell (12 total), its small size up to 13 mm, and the presence of sexually dimorphic gonads, markedly distinguishes it from other cubozoan taxa.

Keywords: box , Tripedalia cystophora, Tripedaliidae, Cubozoa, Gulf of Mexico, distribution

Tripedalia cystophora Conant, 1897 is a species has been the subject of several small (maximum bell height ca. 12–13 phylogenetic and developmental studies mm) box jellyfish (Cnidaria: Cubozoa) (Werner 1973, Laska-Mehnert 1985, that inhabits tropical and subtropical Bentlage et al. 2010, Gurska & Garm mangrove systems (Conant 1897, Kramp 2014). Its elaborate copulation strategies have been well documented (Werner 1971, 1961, Ekins & Gershwin 2014). Since its 1973, Stewart 1996), making it an excel- description from Jamaica in 1897, the lent cubozoan for laboratory and field * Corresponding author: Robert.Lasley@ studies examining paternity, sexual selec- myfwc.com tion, and the potential involvement of DOI: 10.2988/0006-324X-129.Q2.164 visually-guided behavior related to mat- VOLUME 129, NUMBER 2 165 ing. T. cystophora is one of three species Florida (Orellana & Collins 2011), north- in the family Tripedaliidae (Order: Car- ern Australia, (Ekins & Gershwin 2014), ybdeida), along with T. binata Moore, and Hawai‘i (Crow et al. 2015). Ekins and 1988 and (Stiasny, 1926), Gershwin (2014) reviewed the geographic all of which are sexually dimorphic and distribution of the species, listing various produce spermatophores (Lewis & Long Atlantic, Pacific, and Indian ocean local- 2005, Bentlage et al. 2010, Bentlage & ities and, like prior authors, suggested Lewis 2012, Straehler-Pohl et al. 2014). that these reports represent recent intro- While spermatophore transfer from male ductions rather than oversight of earlier to female is well documented in T. extensive surveys conducted in those cystophora and C. sivickisi,(Werner localities. Our new record of the species 1973, Lewis & Long 2005, Underwood is the first in the Gulf of Mexico Large et al. 2013), details remain unclear for T. Ecoregion, an area differing from neigh- binata.Furthermore,T. cystophora has boring regions (e.g., Caribbean) based on long been used as a model for elucidating its distinct biota, genetic breaks between it the biomechanics, physiology, and evolu- and bordering regions, extensive freshwa- tion of the complex lens- characteris- ter drainage, currents, and other abiotic tic of all box jellyfish (Nilsson et al. 2005, factors (Kumpf et al. 1999, Felder & Garm et al. 2007, Garm & Mori 2009). Camp 2009, Fautin et al. 2010). Further- The most recent study of this species used more, the northernmost records for T. a genomics approach to investigate cubo- cystophora, presented herein, lie on the zoan diversification (Liegertova´ et fringe between the Northern Gulf and al. 2015). Despite extensive research on Floridian subregions, which is the north- these various aspects of the species’ ern latitudinal limit of mangroves and a biology and the description of two types transition to oyster reef and saltmarsh of nematocysts (Orellana & Collins 2011), dominated habitats (Fautin et al. 2010). the components of the nemato- Our report therefore likely represents the cysts are yet to be described. Unlike other northernmost limit of T. cystophora in the box jellyfish that are notorious for painful Gulf of Mexico and perhaps the Atlantic and sometimes debilitating stings (Russell Ocean. et al. 1984, Brinkman & Burnell 2009, Bentlage et al. 2010), there are no documented human stings by T. cysto- Materials and Methods phora despite the extensive number of publications on this species. Furthermore, The specimens were identified as Tripe- T. cystophora has never been listed among dalia cystophora based primarily on: 1) the dangerous box jellyfish in the scientific presence of four groups of tentacular literature (e.g. Williamson et al. 1996), pedalia, one at each corner of the bell; 2) suggesting that its sting is mild to three pedalia per group; and 3) each humans. pedalium with one (Conant 1897). The species has a widespread, pantrop- In September 2010, numerous T. cys- ical to subtropical distribution. Within 40 tophora medusae were noticed swarming years of the original description from near red mangroves (Rhizophora mangle) Jamaica (Conant 1897), T. cystophora in a waterway adjacent to Estero Bay near was reported from the eastern Pacific Bonita Springs, Florida. Twelve exemplars and Coral Triangle (Stiasny 1926, Thiel (bell height 5.5–13 mm), including males 1936). In the past 4 years, the reported and females, were collected by a resident range of the species has expanded to and delivered to the Florida Gulf Coast include records from the east coast of University’s Vester Field Station in Bonita 166 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Springs, where they were held in a small a Nikon D7100 DSLR with 60 mm AF aquarium for observation. The samples Micro-Nikkor f2.8D and the Nikon R1C1 were separated into three lots: 2 lots were strobe package. Specimens were photo- prepared in 5% formalin/seawater solution graphed from above in a shallow aquarium and 1 lot in 95% ethanol for future with a black velvet background. Measure- molecular studies. Material was deposited ments of bell height and bell width are into the National Museum of Natural provided below under material examined History, Smithsonian Institution (USNM). in the following format (bell height 3 bell The following year, a large number of width). medusae were seen swarming at the Vester Additional specimens were observed but Field Station in August 2011 (Lasley et al. not collected at Smacks Bayou (2007) and 2016). between Cockroach Bay and Anna Maria On July 15, 2015, six T. cystophora Island (2011), Tampa Bay, Florida (Sheri medusae (bell height 4.8–9.5 mm) were Parks pers. obs.); and Everglades City, collected in a brackish canal parallel to Florida (2015; including identifiable fe- East 3rd Street, Englewood, Florida (Fig. males) (Anders Garm pers. comm.). Spec- 1). The canal feeds into Rock Creek ca. 1.5 imens from Smacks Bayou, St. Petersburg km from the creek’s mouth at Bay. and Everglades City, Florida were photo- Specimens were caught near Rhizophora graphed, the images providing reliable mangle roots roughly one hour before identification of the specimens as T. sunset (20h25; with low tide (0.1 ft) at cystophora (Lasley et al. 2016). 20h47). One exemplar (Fig. 1A) was determined to be female based on the presence of large eggs and embryos circu- Systematics lating in the gastrovascular cavity. The other specimens appeared to be post- Phylum Cnidaria Verrill, 1865 spawned adults given their mature size Class Cubozoa Werner, 1973 but lack of gonads, so the sex could not be Order Gegenbaur, 1857 verified (Fig. 1B–F). After live transport to Family Tripedaliidae Conant, 1897 the Fish and Wildlife Research Institute, Tripedalia Conant, 1897 St. Petersburg, Florida, each specimen was Tripedalia cystophora Conant, 1897 photographed (Figs. 1, 2), measured, preserved in 10% formalin/seawater solu- Material examined.—12 medusae: 1 tion, and deposited into the Institute’s gravid female (5.5 mm 3 7.7 mm), marine collection (FSBCI). Prior to pres- gastrovascular cavity filled with planulae, ervation in formalin, one tentacle from suggesting recent (within days) fertiliza- each specimen was removed and stored in tion, 1 unsexed (9.5 mm 3 13 mm) in 95% EtOH for future molecular study. formalin (USNM 1204759); 3 unsexed in Preserved specimens from Englewood were 95% ethanol (USNM 1204760); 4 mature examined using a Zeiss Discovery V8 females (8.33 3 12 mm – 10 3 13 mm), 2 SMZ-168 stereoscope with manual 8x mature males (9 3 12.6 mm), 1 unsexed (10 zoom. Live color photos were taken using mm x 14 mm) in formalin (USNM

’ Fig. 1. Tripedalia cystophora. Lateral view of live specimens. A, ripe female (7.8 mm 3 9.2 mm) (FSBCI 129558, 1); B, sex undetermined (9.5 mm 3 11.4 mm) (FSBCI 129558, 2); C, sex undetermined (8.1 mm 3 9.5 mm) (FSBCI 129558, 3); D, sex undetermined (4.8 mm 3 6.5 mm) (FSBCI 129558, 4); E, sex undetermined (7.6 mm 3 9.4 mm) (FSBCI 129558, 5); F, sex undetermined (5.2 mm 3 7.9 mm) (FSBCI 129558, 6). VOLUME 129, NUMBER 2 167 168 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Tripedalia cystophora. A–C, ventral, lateral, and dorsal views, respectively of live specimen, sex undetermined (9.5 mm 3 11.4 mm) (FSBCI 129558, 2).

1205451) – collected from red mangroves Conclusions near Bonita Springs, Florida, Gulf of Mexico (2682205300 N8185000800 W), USA, Of the seven box jellyfish species report- coll. C. Ritter, 7 September 2010, daylight ed from the Gulf of Mexico, including those in this study, three have been well hours. documented as long-term inhabitants: 1) 6 medusae: 1 female, 5 sex undetermined alata (Reynaud, 1830) (¼ (4.8 3 6.5 mm – 9.5 3 11.4 mm) (FSBCI alata) (Larson et al. 1991, Lewis et al. 129558, 1–6) – collected from red man- 2013); 2) sp. (T. cf. haplonema) groves, in canal parallel to East 3rd Street, Mu¨ller, 1859 (Phillips & Burke 1970, Englewood, Florida, Gulf of Mexico Collins et al. 2011); and 3) (26.941419, 82.335328), USA, coll. M. quadrumanus Mu¨ller, 1859 (Mu¨ller 1859, Vorkapic, R. Lasley, 15 July 2015. Guest 1959, Phillips & Burke 1970, Go´mez Range.—Tripedalia cystophora occurs in Aguirre 1986). In addition to these well- tropical and subtropical mangrove habi- documented taxa, Alatina grandis (Agassiz tats between 328N and 288S latitude. In the & Mayer, 1902) was reported (as Carybdea Atlantic, the species has been reported var. alata) a single time in the Gulf of from Jamaica, Puerto Rico, the Bahamas, Mexico (Graham 1998). Furthermore, and the east coast of Brazil (Morandini & Hedgepeth’s (1954) species checklist of Marques 1997); the east coast of Florida the Gulf of Mexico includes Carybdea (Orellana & Collins 2011); and the Gulf of aurifera Mayer, 1900, and newer checklists Mexico (this report). Pacific Ocean speci- (Segura-Puertas et al. 2003, Segura-Puer- mens have been reported from Japan tas et al. 2009) include Carybdea marsu- (Uchida 1970); Costa Rica (Rodr´ıguez- pialis (Linnaeus, 1758). However, the Sa´ enz & Segura-Puertas 2009); northern former corresponds to collections by Australia, Thailand, and Indonesia (Ekins Mayer (1900) from Tortugas and the latter & Gershwin 2014); and Hawai‘i (Crow et to reports by Sua´ rez-Morales et al. (1999) al. 2015). Indian Ocean specimens have from the western Caribbean Sea. been reported from Aldabra Atoll, Sey- All species reported from the Gulf of chelles (Ekins & Gershwin 2014). Mexico, including T. cystophora, belong to VOLUME 129, NUMBER 2 169 the order Carybdeida except Chiropsalmus mangrove systems into higher latitudes quadrumanus. Tripedalia cystophora is eas- (Saintilan et al. 2014) and the possible ily distinguished from the other species of increase in frequency of jellyfish blooms Carybdeida by the presence of three pedal- (Condon et al. 2013), the range expansion ia/tentacles at each interradial corner of of T. cystophora and its possible link to the bell (versus one in other Gulf species of climate change and human introduction is Carybdeida) (Fig. 2) (Conant 1897, Mayer worthy of investigation. Molodtsova 1910). (Chiro- (2009) reviewed the long history of medu- dropida: Chiropsalmidae), superficially re- sozoan studies in the Gulf. Despite being a sembles T. cystophora in having multiple small species that can easily blend in with tentacles at each corner, but C. quadruma- mangrove infrastructure and murky water nus possesses only a single pedalium per (Stewart 1996, Garm et al. 2012), the fact corner that is divided into 5–9 branches, that T. cystophora was previously never each giving rise to a single tentacle (Mayer reported in the Gulf of Mexico, despite 1910, Calder 2009). decades of extensive medusozoan sam- Conant’s 1897 original description of T. pling, casts doubt on their prior occur- cystophora does not allude to any type rence in the region. Correspondingly, specimens. Furthermore, the legendary F. recent new records of the species in S. Conant described T. cystophora as a Australia (2014) and (2015) are species new to science while he was assumed to have resulted from human Director at the Johns Hopkins Marine introductions (Ekins & Gershwin 2014, Laboratory at Kingston Harbor, Jamaica Crow et al. 2015), and the likelihood of (Conant 1897), but there is no indication anthropogenic influence and cryptogenesis that Conant deposited any cubozoan (introduction from an obscure locality) specimens into the affiliated Institute of was addressed in a report on the recent Jamaica Museum (IJM) (Muka 2014) appearance of Copula sivickisi, another prior to his untimely death in 1897. A member of the Tripedaliidae, in the Johns Hopkins circulatory published in western Atlantic (Bennett et al. 2011), memory of Conant and his work (Brooks which had been considered strictly an 1900) states that Conant ‘‘brought home Indo-Pacific species (Lewis et al. 2008). specimens’’, but it does not say where they The recently published T. cystophora were deposited (Brooks 1900:2). No inver- genome, along with future large-scale tebrate zoology collection currently exists morphological and molecular studies, at Johns Hopkins University, bringing the could serve as a tool to elucidate the existence of the specimens further into mechanism of dispersal resulting in the question. Nonetheless, given the complete- apparent recent global range expansion of ness of Conant’s original description and the species. accompanying illustration of the speci- mens, there is no doubt about the species’ Acknowledgments identity as T. cystophora, negating the need to erect a neotype in accordance with Thanks to Michael and Joy Vorkapic ICZN Article 75.2 (ICZN). Finally, it has for collecting the Englewood specimens been suggested that the species has been and notifying the Fish and Wildlife extirpated from its type locality of King- Research Institute. We also acknowledge ston, Jamaica (Ekins & Gershwin 2014). Chris Ritter for collecting the Bonita This new record of T. cystophora Springs samples and notifying the Vester extends the species’ known range to the Field Station. Thanks also to Carol Davis Gulf of Mexico as far north as Tampa for notifying the first author after receiv- Bay. Together with the movement of ing the report. Thank you to Geoff Keel, 170 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

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