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First Record of the Box Jellyfish Tripedalia Cystophora

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First Record of the Box Jellyfish Tripedalia Cystophora PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 129:164–172. 2016. First record of the box jellyfish Tripedalia cystophora (Cnidaria: Cubozoa: Tripedaliidae) in the Gulf of Mexico 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 Invertebrate 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 species 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 tentacles total), its small size up to 13 mm, and the presence of sexually dimorphic gonads, markedly distinguishes it from other cubozoan taxa. Keywords: box jellyfish, 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 Copula sivickisi (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-eyes 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 opsin 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 venom 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 tentacle (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 Lemon 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 (À0.1 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 Carybdeida Gegenbaur, 1857 verified (Fig. 1B–F). After live transport to Family Tripedaliidae Conant, 1897 the Fish and Wildlife Research Institute, Genus 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).
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