Original Article Horizontal and vertical movements of ...

ICES Journal of Marine Science (2018), 75(2), 844–857. doi:10.1093/icesjms/fsx176

Contribution to the Themed Section: ‘International Billfish Conference’

Original Article Horizontal and vertical movements of white marlin, Kajikia albida, tagged off the Yucatań Peninsula J. J. Vaudo1*, M. E. Byrne1,‡, B. M. Wetherbee1,2, G. M. Harvey1, A. Mendillo Jr 3, and M. S. Shivji1 1

The Guy Harvey Research Institute, Nova Southeastern University, 8000 North Ocean Drive, Dania Beach, FL 33004, USA Department of Biological Sciences, University of Rhode Island, 120 Flagg Road, Kingston, RI 02881, USA 3 Keen M International, Av Rueda Medina 318, Centro - Supmza. 001, Isla Mujeres, Quintana Roo 77400, Mexico 2

*Corresponding author: tel: þ1 (954) 262 3677; fax: þ1 (954) 262 4098; e-mail: [email protected] ‡ Current address: School of Natural Resources, University of Missouri, Columbia, MO, USA. Vaudo, J. J., Byrne, M. E., Wetherbee, B. M., Harvey, G. M., Mendillo, A., and Shivji, M. S. 2017. Horizontal and vertical movements of white marlin, Kajikia albida, tagged off the Yucatań Peninsula. – ICES Journal of Marine Science, 75: 844–857. Received 31 December 2016; revised 31 July 2017; accepted 15 August 2017; advance access publication 27 September 2017. The white marlin, Kajikia albida, is a highly migratory, prized sport fish of conservation concern. Improved understanding of white marlin ecology, including habitat use, will inform management measures. To improve white marlin movement knowledge in a region with limited information, we tagged 18 individuals off the eastern Yucatan Peninsula, Mexico with pop-up satellite archival transmitters. Tracks lasting 9–328 d, yielded data across 1277 d, and covered distances of 891–10 579 km. Horizontal movements varied greatly with ten individuals remaining in the Gulf of Mexico/northwestern Caribbean and eight individuals entering the western North Atlantic. Although white marlin experienced a temperature range of 10.0–33.6 C, the majority of time was spent in waters >24 C. Marlin displayed diel diving patterns with deeper dives occurring more frequently during the daytime. As water columns warmed, dive duration, maximum daily depth, and dive depth all increased. As a result, 18% of the time was spent at depths >100 m in the warmest water columns compared with 25% of the “white marlin” catch from the western North Atlantic (Beerkircher et al., 2009), highlighting management and conservation concerns for white marlin. Because white marlin mortality is largely the result of bycatch in commercial fisheries, management policies have depended largely on quotas and the practice of releasing individuals alive when captured in fisheries (Graves and Horodysky, 2015). However, management could also be improved with increased understanding of white marlin movements, particularly in

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Horizontal and vertical movements of white marlin regards to interactions with fisheries and in parts of the extensive range of white marlin, such as the Caribbean Sea and Gulf of Mexico, where their movements are not well-documented (Brill and Lutcavage, 2001; Goodyear et al., 2008; Sippel et al., 2015). For example, in some cases modifying fishing gear depth distributions on the basis of vertical depth distributions of non-target species can reduce bycatch without harming catch rate of target pelagic species; eliminating shallow hooks from experimental tuna longlines significantly reduced the catch of many non-target species, including billfish (Beverly et al., 2009). Information of horizontal movements could also identify areas of concentrated use by non-target species that could be designated as fisheryclosure zones. Analysis of fisheries data suggests such zones could substantially reduce catch of non-target species with only limited reduction in target catch (Goodyear, 1999). Although white marlin have been tagged for a variety of studies, information about their movements is relatively limited. Tagand-recapture data suggest a single stock that encompasses the North and South Atlantic, with individuals making long distance, even transatlantic, and migratory movements. However, recapture rates, which are fishery dependent, are low and provide coarse movement resolution (Orbesen et al., 2008). Pop-up satellite archival transmitting tags provide a fishery independent method of examining white marlin movements, including vertical movements. Initial work using these tags focused on post-release survival and was limited to short durations (primarily 5–10 d; Horodysky and Graves, 2005; Kerstetter and Graves, 2006). More recent work has examined longer-term movements (tag deployments up to 1 year) confirming long-distance movements and migratory patterns for white marlin in the western North Atlantic, as well as primarily epipelagic habitat use (Horodysky et al., 2007; Hoolihan et al., 2015; Loose and Graves, 2016; Schlenker and Graves, 2016). Such long-term studies are critical to identifying the full breadth of behaviours and their relationships with environmental factors because they allow observations of individuals experiencing spatiotemporal variations in environmental conditions. Although several white marlin have been tracked in the eastern Caribbean (Hoolihan et al., 2015), previous satellite telemetry studies have focused primarily on fish tagged off the Mid-Atlantic Bight region of the United States, which is near the northern limit of the white marlin range. This region experiences large seasonal temperature changes and cooler waters, not characteristic of the bulk of the white marlin distribution (Longhurst, 2007). Because of temperature’s effect on physiology (Fry, 1971), it is a major driver of fish movements. Therefore, vertical movements and depth preferences of white marlin tagged off the Mid-Atlantic Bight may not be representative of individuals throughout the species range. Many pelagic species, including those sympatric with white marlin, alter their dive behaviours, expanding and contracting their vertical distributions in response to changing thermal conditions over the course of their travels (Walli et al., 2009; Weng et al., 2009; Vaudo et al., 2016). Because behavioural information can help craft management decisions, understanding the relationship of white marlin movement behaviours with temperature is vital for the conservation of this species. The waters of the western Caribbean and Gulf of Mexico, which support a thriving white marlin recreational fishery and serve as white marlin spawning grounds (Rooker et al., 2012), are warmer than and do not experience the dramatic seasonal environmental changes observed off the Mid-Atlantic Bight (Longhurst, 2007), thus


providing a useful environmental contrast with regions white marlin have been previously tracked. In addition, currents and eddies (e.g. the loop current) create thermal variability within the Gulf of Mexico and Caribbean that marlin are likely to encounter, which could elucidate the effect of temperature on white marlin vertical distribution and dive behaviour. We used pop-up satellite archival transmitting tags to investigate the long-term continuous movements of white marlin tagged off the Yucatań Peninsula. Our goals were to examine the horizontal and vertical movements of white marlin in an area that does not experience the dramatic seasonal environmental changes found in other parts of the white marlin’s range. In particular, we describe white marlin habitat use with regards to depth and temperature, and examine the effect of the thermal environment on white marlin vertical movements in this region. We also compare white marlin vertical habitat use here with their movements in other regions, expecting the warmer waters of the Gulf of Mexico/Caribbean to provide preferred habitat to deeper depths, and therefore an expanded depth distribution compared with cooler regions of their distribution.

Material and methods Marlin tagging Twenty-one white marlin were captured via rod and reel using circle hooks in the vicinity of Isla Mujeres, Mexico, off the eastern Yucatań Peninsula, between May 2014 and June 2016. This location is a popular sport fishing destination, particularly for billfish, including white marlin. Each white marlin was brought aboard the vessel for processing and a saltwater hose was inserted into the mouth to ventilate the gills. For each fish, lower jaw fork length was measured and a pop-up satellite archival transmitting tag (MiniPAT, Wildlife Computers, Redmond, WA, USA) was anchored into the dorsal musculature of the marlin using an umbrella dart (Domeier et al., 2005). Handling time for each white marlin was 24 C, deeper dives into cooler waters were frequent. This expansion of the vertical distribution beyond the preferred temperature range is likely the result of prey distribution. For example, young Pacific bluefin tuna, Thunnus orientalis, made more dives below the thermocline when food was scarce and ceased this behaviour when food became abundant in surface waters (Kitagawa et al., 2007). Like other billfish, white marlin are visual predators and show clear diel diving patterns (e.g. Sippel et al., 2007; Hoolihan et al., 2015; Lam et al., 2016; Carlisle et al., 2017). These deeper dives primarily occurred during the daytime and were probably related to foraging. In addition to being able to search for prey at greater depths during the daytime because of greater illumination, by diving down into cooler waters to forage, white marlin may have a physiological advantage over prey. In addition to their large size providing them with thermal inertia and, therefore, warmer bodies than their prey, white marlin have modified eye muscle associated with heat production (Block, 1986). The increased temperature of the retina allows for better detection of rapid motion, providing a visual advantage over prey species (Fritsches et al. 2005). Although white marlin may benefit from foraging dives into cooler waters, they may be less suited for this behaviour. White marlin make fewer excursions into cooler waters than sympatric billfishes (blue marlin and sailfish; Hoolihan et al. 2015). It is possible that these dive behaviours could also be a means of regulating body temperature to maximize metabolic scope (Claireaux et al. 2000). Despite white marlin having a sympatric distribution with important fisheries targets, and like these species, are large visual predators that exhibit diel dive behaviours, there are differences in their vertical movements that may be used to reduce bycatch of white marlin in longline fisheries. Many of the vessels in the western North Atlantic longline fishery target swordfish, Xiphias gladius, and bigeye tuna, Thunnus obesus, which have deeper depth distributions than white marlin. Bigeye tuna are found much deeper than white marlin (mean daytime depth: 200 m; Lam et al., 2014), and even the mean nighttime depth of bigeye tuna (45 m; Lam et al., 2014) is deeper than that of white marlin, which rarely dove to 50þ m during the nighttime. Similarly, in thermal conditions similar to those observed in this study, swordfish spend most of the daytime at depths that exceed the maximum depth we observed for white marlin, and although there was overlap in nighttime distributions, swordfish were typically deeper than white marlin (Dewar et al., 2011). Such differences in vertical behaviour between fisheries targets and white

856 marlin suggest that setting minimum hook depths for longlines in these fisheries could reduce white marlin bycatch. Experimental longlines for the Hawaii-based tuna fishery that eliminated hooks set shallower than 100 m reduced the catch of non-target species including three species of billfish without affecting target catch (Beverly et al., 2009). Minimum hook depths, however, may not work to minimize white marlin bycatch in all longline fisheries. Within the Gulf of Mexico, yellowfin tuna, T. albacares, are a major target of longliners, but tend to have a relatively shallow depth distribution, and therefore, have a great deal of vertical overlap with white marlin (Weng et al., 2009). In these fisheries, reducing white marlin catch will likely require exploiting spatiotemporal differences in the horizontal distribution of the two species (Goodyear, 1999). Overall, white marlin tagged off the Yucatań Peninsula displayed widely varying horizontal movements and used deeper waters than previously observed, but exploited similar temperatures to white marlin tagged in other parts of their range. This discrepancy suggests the thermal structure of the water column plays a large role in the vertical distribution of this species. Although, variations in the thermal characteristics of the water column may make predicting the vertical distribution of white marlin difficult without detailed temperature data, differences in vertical distribution of white marlin and some important fisheries targets are substantial suggesting eliminating shallow water hooks from longlines may be a means of reducing white marlin bycatch in some parts of its range.

Supplementary data Supplementary material is available at the ICESJMS online version of the manuscript.

Acknowledgements We thank Andreas Marcher, Rafael de la Parra, and the excellent crew of the Keen M for assistance with the field work. This study was supported by the Offield Family Foundation, Guy Harvey Ocean Foundation, and Guy Harvey Research Institute, Nova Southeastern University.

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Handling editor: Jonathan Grabowski
