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Recommended citation for this article:  Jordan, Frank and Howard L. Jelks.  2024.

Implication of diurnal activity patterns for management of invasive lionfish (Pterois spp.).  Occas. Pap. Loyola Ctr. Enviro. Communication. (6): 1-13.

IMPLICATION OF DIURNAL ACTIVITY PATTERNS FOR MANAGEMENT OF INVASIVE

LIONFISH (Pterois spp.).

by Frank Jordan1 and Howard L. Jelks2

1Department of Biological Sciences, Loyola University New Orleans, 6363 St. Charles Avenue, New Orleans, Louisiana 70118, USA

2Retired from Wetland and Aquatic Research Center, U.S. Geological Survey, 7920 NW 71st

Street, Gainesville, Florida 32653, USA

E-mail: jordan@loyno.edu (FJ), jelkshl@gmail.com (HLJ)

Lionfish foraging above Coral Gardens Reef in Turks and Caicos Islands in October 2011. Photo by Frank Jordan.

Abstract

Indo-Pacific lionfish (Pterois spp.) have invaded the Western Atlantic and are profoundly altering coral reef communities. Modeling indicates that divers must be highly efficient in order to reduce and control invasive populations of lionfish. Increasing detection of lionfish is a key strategy to increasing removal efficiency. We quantified diurnal variation in abundance while snorkeling above shallow coral reefs in Belize, Turks and Caicos Islands, and the Florida Keys to test the hypothesis that lionfish are more active and therefore likely more detectable and harvestable during crepuscular periods. Actively foraging lionfish were indeed significantly more abundant during dawn and dusk than during midday. We then used SCUBA to test the same hypothesis on deeper coral reefs in Belize and similarly found that lionfish were more active during crepuscular periods, but there was no difference in the total number of lionfish detected during midday and crepuscular periods. The disparity in abundance of lionfish detected between these surveys was because we invested greater effort while SCUBA diving into searching for actively foraging lionfish above the reef and inactive lionfish hiding in crevices and under ledges. Taken together, these snorkel and SCUBA studies suggest that conducting surveys and collections of lionfish on shallow and deeper reefs should be focused during crepuscular periods in order to increase detection, maximize harvest rates, and minimize effort and costs.

Finally, incidental behavioral observations made during this study indicate that feeding harvested lionfish to predators has negative consequences for both predators and humans.

Introduction

Invasive species often have strong negative effects on the abundance of native species and profoundly alter invaded communities (Simberloff 1997; Perrings et al. 2010). Effective control and management are therefore required to restore and conserve native communities.

Unfortunately, effective control and management require a significant investment of money and labor. One of the most significant costs, especially for mobile species, is the effort required to detect, capture, and remove invasive species. Innovations that improve monitoring and capturing of mobile species are desirable in order to reduce costs and increase the likelihood of restoring and conserving native communities.

Invasive lionfish (Pterois miles and P. volitans, after Schultz 1986) successfully invaded the Atlantic Ocean from Rhode Island southward to Venezuela and Brazil, and more recently have invaded the northern Gulf of Mexico (Aguilar-Perera and Tuz-Sulub 2010; Schofield 2010; Betancur-R et al. 2011). Lionfish are generally found in the top 50 m of the water column in their native range (Schultz 1986). Lionfish spawn year-round (Morris and Whitfield 2009), are locally abundant (Whitfield et al. 2007; Green and Côté 2009), and have strong negative effects on native communities of reef fishes (Albins and Hixon 2008; Cerino 2010; Arias-González 2011; Green et al. 2012). In addition to preying upon small coral reef fishes and invertebrates, lionfish also prey upon juvenile snappers and groupers (Morris and Akins 2009) and compete with adults of these ecologically and economically important reef predators. A more recent summary of the invasion, distribution, biology, and ecology of lionfish is provided by Del Rio et al. (2023).

Management of invasive lionfish has largely focused on removal of individual fish by SCUBA and snorkel divers using spears or hand nets during lionfish fishing tournaments (Morris

and Whitfield 2009). Indeed, a unique ecotourism industry revolving around lionfish derbies, rodeos, and safaris has developed in southern Florida and throughout much of the Caribbean. Lionfish are readily amenable to individual capture by divers because of bright warning coloration and their unique foraging behavior that involves hovering slowly above the reef with pectoral fins fully extended in order to corner or corral vulnerable prey. Lionfish are also largely indifferent to divers and other potential predators because of their arsenal of venomous spines. Given their bright coloration and slow movement, it is somewhat surprising that fish community censuses only detect about 13% of the lionfish present in a given location (Kulbicki et al. 2012). Population models indicate that highly efficient divers may be able to significantly control populations of invasive lionfish in relatively shallow habitats (Arias-González 2011, Morris et al. 2011, Frazer et al. 2012). The purpose of this study is to provide further guidance to divers and marine resource managers on how to increase detection and capture efficiency, thereby increasing the likelihood of managing populations of invasive lionfish in shallow marine habitats.

Fish communities on coral reefs are diverse in part because species are active at different times of the day and thereby partition available food and space (e.g., Ross 1986). Many reef predators consume relatively more prey during the twilight hours of dawn and dusk. For example, lionfish in the Indo-Pacific Ocean are crepuscular predators (Fishelson 1997). Recent studies indicate that invasive lionfish are also more active and consume more prey during crepuscular periods in the Bahamas (Green et al. 2011), although this may not be the case throughout their newly invaded range (Côté and Maljkovic 2010; Kulbicki et al. 2012). What remains unclear is whether this diurnal variation in activity and foraging behavior contributes to the overall low detectability of lionfish (Kulbicki et al. 2012). If so, then divers should focus their census and eradication efforts during dawn and dusk in order to maximize harvest and achieve desired control and management goals. This study tested the hypothesis that lionfish are more active and exposed to divers during crepuscular periods.

Materials and methods

Snorkel study

Snorkeling provides a cost-effective method to study fish abundance and behavior in shallow, clear habitats (e.g., Jordan et al. 2008). Diurnal variation in lionfish abundance was quantified while snorkeling on reefs in Belize (May 2011), Turks and Caicos Islands (October 2011), and the Florida Keys (November 2011). No attempt was made to differentiate between the two species of invasive lionfishes, although it is likely that they were all P. volitans (Hamner et al. 2007; Betancur-R et al. 2011). A mixture of 16 patch and barrier reef sites were included in the study (Table 1). Of these, ten were visited at least twice during a single day and the remaining six sites were visited once during dawn or dusk. Reefs ranged from approximately 1 to 10 m deep. During each observation period, one or two divers snorkeled along the ocean surface and located and counted actively foraging lionfish below. No attempt was made to dive down and search for lionfish hiding in crevices or under ledges. Divers snorkeled in parallel transects, used prominent reef markers, and differentiated among individual fish based on size and coloration in order to ensure counts were not duplicated. Direct measurement and evaluation of maps in Google Earth indicate that reefs ranged from 18-15,525 m2 in area (5,112 ± 1,217; mean

± 1 SE) and required 5 to 45 minutes to census.

Lionfish were counted at dawn, midday, and dusk on eight of the study reefs. Dawn was considered one hour before and two hours after sunrise, midday as two hours before and after noon, and dusk as two hours before and one hour after sunset. Lionfish were counted at midday and either dawn or dusk at another two reefs but could not be revisited for a third diurnal observation due to logistical constraints. Another six sites were visited one time at either dawn or dusk and are included herein to provide additional information on abundance of invasive lionfish.

Analysis of variance was used to test the hypothesis that abundance of lionfish varied among dawn, midday, and dusk census periods at the ten sites with repeated visitations. To control for repeated observations to the same site, each reef was used as a statistical block in this analysis. We predicted that abundance of lionfish would be lower during midday and therefore used a planned contrast to compare the abundance of invasive lionfish active during midday to the abundance of lionfish active during dawn and dusk.

SCUBA study

Our snorkeling study evaluated a cost-effective way to quickly study lionfish abundance and behavior on shallow reefs. We conducted a second study to examine abundance and activity patterns of invasive lionfish on deeper reefs (≈10-25 m) that were not amenable to snorkel surveys. Diurnal variation in lionfish activity was quantified while SCUBA diving on 12 different reef locations on the southeastern side of Glover’s Reef Atoll in Belize during May 2012. Each dive lasted 35-45 minutes depending on depth, covered similar distances (not quantified), and involved 2-4 divers moving up and down the atoll reef wall and searching for lionfish actively swimming just above the reef or hiding under crevices, ledges, and other spaces. We counted each lionfish and noted whether it was active or hiding. Working with a commercial dive operator, we were able to complete two dives before 0800 (dawn), five dives between 1100 and 1300 (midday), and five dives between 1600-1800 (dusk). Given that we were unable to quantify the distance covered, lionfish abundance data should be considered indexes of abundance (e.g., CPUE).

Analyses of variance were used to test the hypotheses that abundance of lionfish and percentage of active lionfish varied between dawn, midday, and dusk census periods. We predicted that both abundance and activity of lionfish would be lower during midday and therefore used planned contrasts to compare abundance and percentage of lionfish active during midday to the abundance and percentage of lionfish active during dawn and dusk.

Results

Snorkel study

We observed from 0 to 13 (2.7±1.0, mean ± SE) lionfish on reefs in Belize, Turks and Caicos Islands, and the Florida Keys during 2011 (Table 1). There was no correlation between the area of reef surveyed and the number of lionfish observed (r=-0.04, p=0.7275). Abundance of lionfish varied significantly during the day (F2, 21=3.332, p=0.0275; Figure 1), with significantly more lionfish active during dawn and dusk than during midday (planned contrast F=9.55,

p=0.0094). More extensive observations on a single patch reef in Belize indicate that diurnal lionfish activity was minimal outside of crepuscular periods (Figure 2).

SCUBA study

We observed from 1 to 23 (11.8±6.6) lionfish on deeper reefs at Glover’s Reef Atoll in Belize. There was no difference in the abundance of lionfish detected during midday than during dawn and dusk (F2,11=0.080, p=0.9238). The percentage of active lionfish varied significantly during the day (F2,11=15.687, p=0.0012), with significantly (planned contrast F=31.138, p=0.0003) more lionfish active during dawn (88%) and dusk (67%) than during midday (7%).

Discussion

Quantifying variation in daily activity patterns of lionfish is essential to accurately estimating and modeling the foraging pressure this invasive species exerts on native fish communities. Native lionfish in the Indo-Pacific (Fishelson 1997) and invasive lionfish on reefs in the Bahamas (Green et al. 2011) and in the Florida Keys, Turks and Caicos Islands, and Belize (this study) actively forage during crepuscular periods and then spend much of the remainder of daylight hours inactive in crevices, under ledges, and in other hiding spaces. Both our snorkel surveys of shallow reefs and SCUBA surveys of deeper reefs demonstrated significantly increased foraging activity during crepuscular census periods. The current study complements Green et al. (2011) by documenting similar patterns of variation in daily activity levels of lionfish from disparate geographic locations and by highlighting how this information can be used to enhance management of lionfish. Invasive lionfish in some Atlantic locations may be more active throughout the day than native lionfish in the Indo-Pacific (Côté and Maljkovic ́ 2010; Kulbicki et al. 2012). Differences in environmental conditions (e.g., amount of cloud cover) or behavioral adjustments of predators and prey (Bosiger et al. 2012) may help to account for observed differences between locations.

There is an emerging consensus that amelioration of negative effects of invasive lionfish will require efficient, direct, and regular removal of lionfish (Morris and Whitfield 2009; Albins and Hixon 2011; Morris et al. 2011; but see Chew 2011). The most commonly used method for control of lionfish populations is to spear or net adults while SCUBA diving. Our results indicate that SCUBA divers should be able to detect and capture invasive lionfish throughout the day as long as divers thoroughly search in crevices, under ledges, and in other hiding spaces. We were able to capture most of the active (100%) and inactive (82%) lionfish that we pursued while SCUBA diving on the wall at Glover’s Reef Atoll. It is likely that SCUBA divers who are not thoroughly searching for lionfish in daytime hiding spaces would have significantly lower detection and capture rates, which may help explain very low detection rates (13%) in published studies of lionfish abundance (Kulbicki et al. 2012). There is likely a tradeoff between the amount of time needed to detect hiding lionfish and the amount of reef that can be searched by SCUBA divers. Focusing search efforts during crepuscular periods when lionfish are most active and exposed should minimize this tradeoff.

Recreational SCUBA divers participating in derbies, rodeos, and safaris account for much of the current effort to manage invasive lionfish. For example, recreational divers removed 1,518 lionfish in the Florida Keys during three rodeos in 2011 prior to our surveys there (NOAA 2011).

This novel form of ecotourism is a boon to local economies and provides educational opportunities in which divers learn about invasive species, predator-prey interactions, competition, and other ecological principles while volunteering to cull populations of lionfish. There is little chance of accidental bycatch because of the unique appearance and behavior of foraging lionfish. Commercial dive operations typically schedule dives during daytime hours when lionfish are least active (i.e., after dawn and before dusk) and revisit many of the same locations. Based on our data and Green et al. (2001), these dive schedules should be shifted to crepuscular periods when the primary goal is to harvest lionfish. Also, a greater range of dive sites should be visited given the extensive geographic range of invasive lionfish and strong site fidelity of adults (Jud and Layman 2012). Increased costs of traveling to new and more distant sites could be offset because more efficient detection and capture of lionfish during crepuscular periods will reduce the amount of fuel and other supplies needed per lionfish harvested.

Harvesting of invasive lionfish by recreational divers should be encouraged to support natural resource management needs whenever possible. For example, recreational divers could be enlisted to continually remove lionfish from reefs in marine reserves or other areas with special conservation or management status (Albins and Hixon 2011). Commercial fishing of lionfish is emerging as another tool used to control lionfish abundance. Collaboration between marine natural resource managers and recreational divers provides an excellent opportunity for service learning while also helping to restore and conserve altered reef ecosystems and enhance local economies through increased ecotourism. With that said, we would like to report herein a potential risk associated with unregulated hunting of lionfish by recreational divers. A group of ten divers, including a few actively spearfishing lionfish, were intensely harassed by a 2-m long great barracuda (Sphyraena barracuda) at Glover’s Reef Atoll. This harassment included aggressive circling, repeated charges, nips at fins and creel bags, and unresponsiveness to attempts to scare the barracuda away with pole spears (personal observation). This behavior persisted for about ten minutes until we decided to abandon the site. Discussion with our dive master indicated that this fish had engaged in the same kind of behavior with a group of divers who were not spearfishing. We later learned that a different dive operator had been regularly spearing and feeding lionfish to this barracuda, either as a form of wildlife feeding tourism or in the hope of increasing predation rates on lionfish. Marine wildlife feeding is a growing ecotourist activity that clearly has serious potential risks for both wildlife and tourists (e.g., Orams 2002).

Capture of individual fish while SCUBA diving is currently the most effective method to control populations of invasive lionfish in deeper habitats. Lionfish also use shallow reef habitats that are less likely to attract or require the attention of SCUBA divers. One of our goals was to determine if snorkeling could provide a relatively time and cost-effective method to detect and capture lionfish in these shallow habitats. Our snorkel data indicate that detection and capture efficiency will be greatly increased if efforts are focused during the crepuscular periods of dawn and dusk. We captured 100% of the lionfish on unnamed patch reef 1 (see Table 1) in Belize between 06:30 and 07:30 using mask, snorkel, and pole spear. Note that we would have detected and therefore harvested zero lionfish if collections were made around midday. Active searching of crevices and other daytime hiding spaces would likely yield more lionfish than just snorkeling along the surface but it would also require greater time and effort. Our data indicate that snorkeling is a cost-effective alternative for harvesting lionfish from relatively shallow marine habitats, especially when efforts are concentrated during crepuscular time periods. Encouraging

removal of lionfish from shallow marine habitats by snorkelers increases the overall eradication effort, which is essential given the extensive geographic range of invasive lionfish (Arias- González 2011, Morris et al. 2011, Frazer et al. 2012).

Finally, monitoring abundance is central to long-term control and management of invasive lionfish in the Atlantic Ocean (Morris and Whitfield 2009). Results of this study indicate that monitoring should be performed during crepuscular periods in order to maximize detection and increase accuracy of historically poor estimates of lionfish abundance (Kulbicki et al. 2012).

Acknowledgements

We thank Danielle Pauli, Warren Cabral, and Bryan Edwards for logistical assistance at Glover’s Reef Atoll and Simone Pitre for allowing FJ to survey lionfish during their honeymoon in Turks and Caicos Islands. We also thank Susan Dye for assisting with surveys in the Florida Keys. Lisa Jelks helped edit the final manuscript. Research in Belize was conducted under the authority of Marine Scientific Research Permit 000019-12 issued to FJ by the Belize Fisheries Department and IACUC Permit 2012-10 issued to FJ by Loyola University New Orleans.

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Table 1. Number of invasive lionfish observed on reefs in Belize (BEL), Turks and Caicos Islands (TCI), and Florida Keys (FLK). Asterisks denote sites where lionfish were censused multiple times during a single day and for these sites the number of lionfish listed is the maximum observed.

Figure 1. Mean (± 1 SE) number of lionfish observed during dawn, midday, and dusk census periods on ten reefs in the Caribbean.

Figure 2. Daily abundance of invasive lionfish on a single patch reef at Glover’s Reef Atoll, Belize on May 25, 2011.

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