Reference: Biol. Bull. 215: 57 62.

(August 2008) 2008 Marine Biological Laboratory

Preliminary Observations on the Response of Chironex eckeri (Cnidaria: Cubozoa: Chirodropida) to Different Colors of Light
LISA-ANN GERSHWIN1,* AND PETER DAWES2 School of Marine & Tropical Biology, James Cook University, Townsville, Queensland 4810, and National Marine Stinger Advisor, Surf Life Saving, Bondi Beach, New South Wales 2026, Australia; and 2 Surf Life Saving, 18 Manning Street, South Brisbane, Queensland 4101, Australia, and Royal National Lifeboat Institution, West Quay Road, Poole, Dorset BH15 1HZ, England
1

Abstract. Cubozoans are well known for their attraction to light and light-colored objects. Two highly venomous types are a public safety concern in Australian waters and elsewhere: Chironex eckeri, long considered the worlds deadliest animal and colloquially called the box jellysh; and the irukandjis, a group of at least 10 species that cause various degrees of debilitating illness. We were asked by the tourism industry whether there might be a color of light that box jellysh and irukandjis are not attracted to, such that nighttime diving activities might pose less risk of being stung. Our preliminary trials with Chironex eckeri indicated a marked positive response to lights of white, red, yellow, green, orange, and blue. All colors elicited a strong and directed attraction to light; however, medusae slowed down their pulsation rate, streamed out their tentacles, and performed a series of gure-eight patterns back and forth through the lighted area when exposed to blue light, which we interpreted as feeding behavior. This compares curiously with a report subsequent to our testing, in which the small, mangrove-inhabiting cubomedusa Tripedalia cystophora and the beach-dwelling Chiropsella bronzie demonstrate a peak sensitivity to blue-green light in the region of 500 nm, and that the former is behaviorally attracted to blue and green light, but ignores red. This leaves open the possibility that Irukandji species, which are more closely related to Tripedalia than to Chironex, may be blind to red.

Introduction Throughout the tropics and subtropics of the world, cubozoans (box jellyshes, irukandjis, and their kin) pose a signicant health threat to humans, particularly near the coasts where commercial and recreational activities take place. In tropical Australian waters, 73 deaths have been attributed to jellysh stings, and about 100 potentially life threatening stings occur each year (Gershwin and Dabinett, 2004). Research into methods to reduce the number of stings is generally focused on predicting jellysh occurrence through computer modeling (J. Seymour, pers. comm., James Cook University), or to prevention through barrier methods such as stinger-resistant swimming enclosures (Moss and Stark, 1983; Anonymous, 2004) or clothing made of Lycra fabric or other spandex fabrics (Sinclair, 2003; Gershwin and Dabinett, 2004). The tourism industry in the Great Barrier Reef region currently contributes over A$5 billion annually to Queenslands economy (Hand, 2003). Although we were unable to quantify the dollar amount attributable to nighttime activitiesfor example, swimming, boating, diving, and shingwe believe it is a respectable amount, and would likely be higher if these activities could be made safer. Currently, the number one marine aquatic health hazard in tropical Australia is dangerous marine stingers, including the large, rapidly lethal, many-tentacled Chironex-type box jellysh and the smaller, four-tentacled irukandjis, a lesser known group of 10 or more species that causes a constellation of debilitating symptoms that may be life-threatening. The attraction of cubozoans to light and light-colored objects is well documented. Matsumoto (1995) gave a good
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Received 19 May 2007; accepted 10 February 2008. * To whom correspondence should be addressed. E-mail: lisa. gershwin@stingeradvisor.com

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summary of numerous earlier observations on cubozoan light responses, and further reported that Carybdea rastonii in the eld was typically found over sandy patches and avoided dark regions such as seagrass beds. Scoresby Shepherd (pers. comm., South Australian Research and Development Institute) also reported that members of this species can often be observed hunting along the sand-seagrass border. Stewart (1996) observed that the feeding behavior of breeding individuals involved making repeated passes through light shafts among the mangroves. Ueno et al. (2000) found that Carybdea rastonii (probably C. mora) would strike grey panels more frequently than either black or white ones, and they had the best results raising medusae with black-sided aquariums. Although Chironex eckeri is well known to respond to light, few experiments have actually been performed to test the limits of this reaction. Barnes (1966) reported that in full daylight, surface and subsurface light intensities seem to have little effect, but in semi-darkness, these animals are very markedly phototaxic. The light of a match is detected at distances up to 5 ft and . . . [they] show a remarkable accuracy in turning towards the light source, even though the latter be extinguished before the turning movement is completed. Hamner et al. (1995) experimented with black and white panels and objects, and found a strong preference for light-colored panels over dark-colored panels, and avoidance of dark-colored objects but not light ones. Many fascinating discoveries have recently been made about cubozoans eye structures and optical properties (Coates, 2003; Martin, 2004; Nilsson et al., 2005; Coates et al., 2006; Garm et al., 2007), but we remain relatively nave about what they actually see and how they interpret their world. Although their attraction to light remains one of the best methods of luring cubozoans for collection, this same characteristic is, unfortunately, a hazard for night divers. The research described here was prompted by a request from the Queensland tourism industry to nd whether lights of different colors might elicit different behavioral responses from cubomedusae; specically, could a light be found that does not attract irukandjis and box jellies? In 2004, we had an opportunity to test the responses of acclimated animals to different colors of lights. A similar opportunity has not arisen since; we therefore present these data in hope of stimulating interest should an opportunity arise again for us or for another research group.

All had already been acclimated to a captive environment for about one week. All had been hand-fed small sh within about 30 min of the beginning of the experiment; two spat their prey before the experiment began, but all others fully completed digestion during the experiment. Acetes australis prawns, a known prey item for young C. eckeri, were available to the medusae in low density in the aquarium throughout the trials and during normal husbandry operations. Medusae were placed together in a 650-l acrylic aquarium measuring 180 60 60 cm. The aquarium was gently aerated in opposite corners throughout the experiment, but was otherwise still. The aquarium had been freshly lled with ltered natural seawater. Five colors of Philips Partylight 40-W incandescent bulbs were tested in the following order, chosen haphazardly: red, yellow, green, orange, blue. White light (Philips Classictone 40-W, clear) was used as a control, since it is often used as an effective cubozoan attractant (Arneson and Cutress, 1976; Larson, 1976; Kinsey, 1986, 1988; Hartwick, 1991) and we could therefore estimate relative response to a known stimulus. Lights were masked off so that the light beam was relatively focused; they were clamped onto a pipe 5 cm above the waters surface and aimed downward at about 45 across the width of the aquarium. Trials were run at an interval of 10 min. Three replicates of each color were tested, without breaks between samecolor trials but with breaks of 510 min between colors. Trials were conducted from late in the afternoon into the middle of the night, in order to identify confounding effects, if any, associated with behavioral periodicity. White lights (our control) were used before and after the color trials to see whether there was a difference in response associated with day-night cycles. For most trials we switched the light from one far side to the opposite far side of the aquarium; however, for the following trials we placed the light in the middle to test whether the medusae were merely swimming to an edge and staying there, or purposely remaining within the light even though they could swim past it: yellow trial 2, orange trial 3, and blue trial 2. The position of each individual was assessed at the following times: time zero, the time of the rst arrival to the light, about 5 min, about 8 min, and 10 min. Because of the small number of samples, statistics were not performed on the data; however, we believe that these preliminary results are worth presenting due to the public safety implications of light attraction in this lethal species.

Materials and Methods All experiments were carried out between the hours of 1700 and 2215 on 16 November 2004. Six juvenile Chironex eckeri ranging in bell height from 3.5 to 5 cm, plus one very young individual at 1-cm bell height, were tested. Results Our observations are summarized in Table 1. All specimens exhibited positive responses to all colors of lights,

JELLYFISH RESPONSE TO COLOR Table 1 Summary of responses of seven juvenile Chironex eckeri to different colors of light during sequential 10-min trials Number in lighted half of tank at 10 min 6 3 7 5 6 4 7 7 6 7 7 6 7 7 3 4 8 7

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Light color White 1 White 2 White 3 Red 1 Red 2 Yellow 1 Yellow 2 Yellow 3 Green 1 Green 2 Green 3 Orange 1 Orange 2 Orange 3 Blue 1 Blue 2 Blue 3 White

Time to rst arrival at light (in minutes.seconds) 5.00 3.00 4.37 3.40 2.24 6.20 4.45 4.30 2.26 5.40 1.30 6.20 1.15 2.05 2.10 2.55 2.10 6.00

Number at light at 10 min 3 2 1 1 4 1 4 4 1 2 5 4 2 2 2 1 3 3

Swimming behavior Swimming Swimming Swimming Swimming Swimming Swimming Swimming Swimming Swimming Swimming Swimming Swimming Swimming Swimming Feeding Feeding Feeding Swimming

ited the fastest overall response; the yellow elicited the least interest; the green and orange the most variable responses. However, all colors of light elicited positive response, typically characterized by a rapid and deliberate swimming toward the light with the tentacles retracted, and once the light was reached, a continued pattern of pulsations upward, as if to come up out of the water. The blue light was remarkable in that it elicited what we believed to be feeding behaviorthat is, a slower, more rhythmic pulsation pattern, with tentacles streaming out behind, and circling through the light beam in repeated gure-eight patterns. Discussion Many types of animals and plants exhibit positive phototaxis: brine shrimp (Bradley and Forward, 1984), mosquitos (Barghini et al., 2004), chaetognaths (Goto and Yoshida, 1981), cephalopods (Moltschaniwskyj and Doherty, 1995), turtles (Salmon and Witherington, 1995), corals (Yamashiro and Nishihira, 1995), and algae (Clifton and Clifton, 1999), to name just a few. Many types of organisms respond to colored light, including butteries (Zaccardi et al., 2006), bees (Backhaus, 1992), mosquitos (Burkett and Butler, 2005), agellates (Suzaki and Williamson, 1983), and sh (Marshall et al., 2003). Other than Coatess (2005) unpublished thesis work, as far as we know, this experiment is the rst of its kind to test the behavioral responses of a species of jellysh to colors of light. It was hoped that one of the colors might be invisible to Chironex eckeri, as is the case with red light for sandies (Mellor and Hamilton, 2003), for the green alga Monostroma angicava (Togashi et al., 1999), and for the cubozoan Tripedalia cystophora (Coates et al., 2006), but this experiment clearly demonstrates that juvenile C. eckeri are attracted to the ve major colors of lightthat is, C. eckeri is not blind to any of the tested colors. The recent spectral studies of Coates and her colleagues (2006) and Garm and his colleagues (2007) on Tripedalia cystophora and Chiropsella bronzie (identied as Chiropsalmus sp.) indicated that both species displayed peak sensitivity in the blue-green region near 500 nm. Garm et al. (2007) speculated that both species were color-blind. However, in their behavioral experiments, Coates et al. (2006) found that T. cystophora was attracted to blue or green light shafts but ignored the red; they hypothesized that because the mangrove world of T. cystophora is primarily green, the blue bias in sensitivity might help improve contrast. We nd this particularly intriguing, because our experiment on Chironex eckeri indicated a very special reaction to blue light, which we interpreted as feeding behavior. Therefore, C. eckeri seemed to us to have two levels of response: rst, the positive response to all colors of light, and second, the feeding behavioral response to the blue light.

with two notable exceptions. One specimen that was badly damaged remained inactive on the bottom of the aquarium throughout most of the experiments, responding only to a single trial of the blue light. Another specimen, also fairly damaged, was highly disoriented most of the time; nonetheless, although it was rarely able to nd the light, it was frequently observed making an effort to move toward the light. Thus, although only 5 specimens participated in all trials, all 7 specimens made positive responses, and no specimens made negative responses. While we intentionally used only white light as a control, darkness also provided somewhat of a control, since we observed medusae to disperse around the tank in the intermissions between the color trials. Position of the light (in the middle half of the tank compared to at one end or the other) had no apparent effect. The fastest times to the light were independent of the light position, as were the numbers drawn to the light. Qualitatively, the smallest juvenile tended to be the fastest to respond and the fastest to arrive at the light stimulus, despite having to work harder due to its smaller size. Not all specimens came all the way to the light, or stayed in the light once arriving; however, the majority of specimens for all colors were in the lighted half of the tank at 10 min, regardless of where they began at time zero and regardless of position of the light. A graphical comparison of time to rst arrival (in seconds) is presented in Figure 1, in the order in which the trials were conducted. The red, white, and blue lights elic-

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Figure 1. Summary of time (in seconds) to rst arrival at light in different trials. Each set of trials for each color was conducted consecutively in the order presented.

It was unclear to us at the time whether the conspicuous behavior difference that we observed in reaction to blue was due to luminosity or to some spectral property of the color blue. Our red and blue lights seemed to us qualitatively to give off similar low levels of light, but the feeding behavior observed with the blue light was not exhibited with the red light, which produced an attraction similar to that of the other colors. However, in light of Coatess observations, it would seem that blue may have some quality that, to the eyes of C. eckeri, is visually different from other colors. Other unsophisticated organisms display unique positive responses to blue light, including agellates (Suzaki and Williamson, 1983) and mosquito larvae (Hribar and Hribar, 2006). Mellor and Hamilton (2003) noticed a pattern similar to that for Tripedalia with their sandies, and interpreted their attraction to blue light as an aid to navigation at dusk or in moonlight or starlight. It was previously reported that C. eckeri sleeps at night (Seymour et al., 2004) based on few observations; however, this was rapidly demonstrated to be erroneous (Currie and Jacups, 2005). Indeed, more than 30% of fatal and nonfatal box jellysh stings occur during the proposed nap time (P. Fenner, Surf Life Saving Australia, pers. comm. 2004). Furthermore, preferential response to low levels of light was reported for Carybdea rastonii (probably C. mora) from Japan (Yatsu, 1917), in observing that medusae come to the surface only in the morning and towards the evening (pg. 2). As a strange coincidence, blue lights are commonly used along foreshores throughout North Queensland; if blue light is further demonstrated to stimulate C. eckeri, and especially if a similar phenomenon is demonstrated for irukandji spe-

cies, the choice of light color for swimming and boating areas would need to be reevaluated. A urry of media interest in mid-2005 speculated that irukandjis might be deterred by the color red (Gerard, 2005; Jeffery, 2005; Robinson, 2005). Our results, along with those of Coates and her colleagues (2006) indicate that this is unlikely to be accurate. However, Coatess results indicate the possibility that irukandjis might nonetheless be blind to red light; if this is someday demonstrated to be the case, it would be a tremendous step forward for the tropical diving industry. Another curious pattern that we observed was the overwhelmingly faster and more assertive response by the smallest juvenile than by any of the other larger study subjects. Whether this was just this individuals response or something inherent in very small individuals could not be ascertained with our sample group. Positive phototaxis in larvae and juveniles has been well documented in many other types of animals and plants, including mosquitos (Hribar and Hribar, 2006), cephalopods (Moltschaniwskyj and Doherty, 1995), gastropods (Barile et al., 1994), intertidal crustaceans (Forward and Cronin, 1979), and even bathyal larvae (Bingham and Young, 1993). We speculate that this juvenile response in C. eckeri may be due to ontogenetic prioritiesthat is, the priority of a juvenile is to survive and grow, and thus food capture is of the utmost importance. Food-linked positive phototaxis was demonstrated in Tripedalia cystophora (Stewart, 1996), by a study in which juveniles were drawn to light shafts to feed on copepods, but breeding males and gravid females did not feed. It might also be interesting to test sexually mature spawned and

JELLYFISH RESPONSE TO COLOR

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unspawned individuals, hungry individuals, and gorged individuals, to determine whether there is a reproductivelinked or hunger-linked response to light. Finally, some taxa show a marked sexual dimorphism in their response to lightfor example, glow worms (Booth et al., 2004), copepods (Chae and Nishida, 2004), sandies (Mellor and Hamilton, 2003), mosquitos (Burkett and Butler, 2005), and fairy shrimp (Luc et al., 1995). A sex-linked response was not tested in C. eckeri, but the possibility seems intriguing. To date, the only sexually dimorphic phenomena noted for cubozoans are velarial spots, which are present in females and absent in males of Carybdea sivickisi (Hartwick, 1991) and Tripedalia cystophora (Lewis and Long, 2005), and dimorphic gonads in the same two species (Werner, 1973; Hartwick, 1991); but then again, nobody has really pursued the question of sex-linked phenomena in jellysh. The behavioral morphology and swimming form that we observed in Chironex eckeri in response to blue light was unique among the responses to different colors of light, and we believe that this phenomenon bears further study. We remain most curious as to whether the blue light feeding response was due to luminosity or to the spectrality of blue. Knowing which is the case will provide useful information for improved management of marine stingers in regions where they threaten human life, and will also provide interesting insights into the limits of brainless visual response. Colors should be examined at random, rather than grouped as in our study. It was interesting to us that the rst replicate of some colors elicited a lesser response than later replicates; we imagined a priori that the opposite might be true, based on initial response to the light without a food payoff. The indication that there was a habituation response at all seems curious, and should be tested. A more rigorously quantied experiment could be conducted on a grid, to track the distance and direction traveled by each individual during the experiment. However, previous studies have indicated that cubozoans respond to substrate and background patterns (Matsumoto, 1995; Stewart, 1996; Ueno et al., 2000); therefore, one must be cautious about confounding the results with patterns that may confuse or deter the animals. The highest priority for research would be to test whether Irukandji jellyshes have a response similar to that we have demonstrated for C. eckeri, or one closer to that previously reported for Tripedalia (Coates et al., 2006). The results of such a test could have important implications for night diving and shing operations throughout the tropics and subtropics of the world where irukandjis occur. Acknowledgments We sincerely thank Tom Hatley for assistance on so many things, and the Reef HQ for allowing us to use their

tanks for this experiment; the statistical analysis benetted tremendously from insights by Ronnie Baker. We are grateful to a charter operator in the Whitsundays for originally asking us whether any color light that the jellysh would not see could be used for night diving; we have regrettably forgotten his identity, but his question sparked our interest to do this experiment. We sincerely thank two anonymous reviewers for helpful comments on the manuscript. Funding for this experiment was provided by Surf Life Saving Queensland and the Reef HQ (Great Barrier Reef Marine Park Authority).

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