Novel Deep-Water Spawning Patterns of Bonefish (Albula Vulpes), a Shallow Water Fish
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Marine Biology (2020) 167:187 https://doi.org/10.1007/s00227-020-03799-3 SHORT NOTE Novel deep‑water spawning patterns of bonefsh (Albula vulpes), a shallow water fsh Steven M. Lombardo1 · Aaron J. Adams1,2 · Andy J. Danylchuk3 · Cameron A. Luck1,4 · Matthew J. Ajemian1 Received: 28 May 2020 / Accepted: 28 October 2020 © Springer-Verlag GmbH Germany, part of Springer Nature 2020 Abstract Coastal marine fshes that form spawning aggregations most commonly exhibit a two-point movement pattern, with loca- tions separated by migration: home range to spawning aggregation site and return to home range. However, the bonefsh, Albula vulpes, partakes in a unique three-point spawning migration. Bonefsh migrate up to 80 km from shallow water home fats to form nearshore pre-spawning aggregations (PSA) before moving ofshore to spawn. Although these broad patterns have previously been documented, details of the ofshore spawning-associated diving behavior have yet to be rigor- ously examined. Using active acoustic telemetry and sonar data in 2019 in Abaco, The Bahamas, we provide a complete account of bonefsh ofshore spawning movements and novel deep diving behavior to 137.9 m. Bonefsh were continuously observed at depths ≥ 100 m for 2 h; a time period that included multiple depth changes and culminated in a spawning ascent to 67.3 m at 0.57 m s−1. These new data on bonefsh ofshore movements and deep dives, coupled with CTD data, suggest that bonefsh actively spawn at pycnoclines and thermoclines. Two previous tracking attempts (2013, 2018) at this location refect spatiotemporal plasticity in spawning, a behavior counter to other aggregation forming fshes. This is the frst detailed documentation of such deep spawning for a shallow water coastal fsh species. The ecological motivation for diving to the deepest depths remains speculative. Future work must examine the dynamic relationship between bonefsh diving behavior, spawning site selection, and oceanographic features. Introduction and Colin 1997). Unlike the somewhat plastic selection of foraging grounds, and to some extent nursery habitat Spawning aggregations are a unique ephemeral reproductive (Petitgas et al. 2012), transient spawning aggregations are strategy where fsh come together en masse under certain highly predictable in that they are temporally and spatially environmental conditions to spawn. Spawning aggregations restrictive, occurring at the same locations and times with for coastal marine fshes may either be resident, where fsh regularity (Johannes 1978; Ciannelli et al. 2015). Spawn- spawn within or near their home range, or transient, migrat- ing aggregation formation is synchronous with seasonal ing beyond the bounds of a realized home range (Domeier and physical cycles, allowing fsh to migrate and arrive at spatially and temporally distant spawning grounds under consistent conditions. Responsible Editor: S. Hamilton. Many identifed spawning aggregation sites for coastal fshes have been located at promontories and slope drop-ofs Reviewed by B. M. Binder and an undisclosed expert. (Sadovy de Mitcheson et al. 2008), positioned to take advan- * Steven M. Lombardo tage of hydrodynamic retention features (Johannes 1978). [email protected] Migrations from a home range to a spawning aggregation often repeatedly follow landmarks and bathymetric features 1 Harbor Branch Oceanographic Institute, Florida Atlantic University, Fort Pierce, FL, USA (Colin 1992; Mazeroll and Montgomery 1998; Feeley et al. 2018). For coastal aggregating fshes, the migration distance 2 Bonefsh and Tarpon Trust, Miami, FL, USA can be tens to thousands of kilometers to reach an aggrega- 3 Department of Environmental Conservation, University tion site (Bolden 2000; Feeley et al. 2018). Some grouper of Massachusetts Amherst, Amherst, MA, USA species have been documented using unconsolidated proxi- 4 Present Address: North Carolina Division of Marine mal staging areas surrounding the aggregation site, where Fisheries, Morehead City, NC, USA Vol.:(0123456789)1 3 187 Page 2 of 11 Marine Biology (2020) 167:187 they exhibit no reproductive behaviors or coloration (Rho- throughout their distributions (Adams et al. 2014; Fedler des and Sadovy 2002a; Robinson et al. 2008). More closely 2013, 2014, 2019; Filous et al. 2019b). Danylchuk et al. associated to the spawning aggregation site, the courtship (2011) used both passive and active acoustic telemetry tech- arena is where reproductive behaviors begin (Nemeth 2012). niques to characterize the timing and location of ofshore Delineation of the components of spawning habitat that sur- spawning movements in Cape Eleuthera, The Bahamas; the rounds an aggregation site can often be difcult (Feeley et al. frst study to do so. Logistical and technological limitations 2018), as such the proximal areas have often been treated of both the passive and active acoustic equipment precluded as a contiguous destination (Rhodes and Sadovy 2002b). any determination of spawning depth, but did reveal consist- Generally, migrations to and from spawning aggregation ent ofshore movement behaviors towards the shelf edge and sites are uninterrupted (Nemeth 2012), following a home waters exceeding 42 m, followed by a return to the shallow range—spawning aggregation—home range migration pat- water fats habitat. Danylchuk et al. (2019) provided a more tern (McCann et al. 2005). Bonefsh (Albula spp.), which detailed description of pre-spawning behavior and ofshore comprises 12 circumtropical shallow water species (Pickett movements from active acoustic tracking eforts from 2013 et al. 2020), are a unique exception in that their migration is and 2014 in Andros and Abaco, The Bahamas. The 2013 interrupted spatially and temporally by a distinct pre-spawn- Abaco active tracking event yielded the frst ever recording ing behavior (Danylchuk et al. 2019). of diving movements during the ofshore spawning migra- Albula vulpes (hereafter bonefsh) are an economically tion. However, the description of fsh vertical movement and culturally important fish that supports recreational through the water column was restricted due to the depth catch-and-release and artisanal fsheries throughout the rating (50 m) of the pressure sensor within the acoustic tags. Caribbean Sea and western North Atlantic Ocean (Adams In this study, we build upon the eforts and data of Danyl- et al. 2014). The estimated annual economic impact of the chuk et al. (2019), presenting the spatiotemporal variability catch-and-release fats fshery, which includes bonefsh, of spawning movements in Abaco. Most signifcantly, we tarpon (Megalops atlanticus) and permit (Trachinotus fal- document for the frst time the complete deep diving and catus), is US$50 million in Belize (Fedler 2014), US$169 presumed spawning behavior of bonefsh via several addi- million in The Bahamas (Fedler 2019), and US$465 mil- tional active tracks. lion in the Florida Keys (Fedler 2013). Bonefsh show high site fdelity to their home range, foraging for invertebrates over a habitat mosaic of intertidal sand fats, seagrass beds, Materials and methods mangroves, and hardbottom (Boucek et al. 2019; Brown- scombe et al. 2017, 2019; Murchie et al. 2013). In The Study site Bahamas, bonefsh migrate from their shallow water home fats to nearshore deeper-water locations where they form The study took place along the southern shore of Abaco, The pre-spawning aggregations (PSA). These activities occur Bahamas. Previous work by the authors identifed a bonefsh over 4–7 days spanning the new and full moons of October PSA at Long Bay that occurs with regularity that allows through April (Danylchuk et al. 2011). The initial migration for repeatable observation. Long Bay is a south-facing bay can cover distances of up to 80 km (Boucek et al. 2019) and along a western-extending point, comprised of a mosaic of culminates with an aggregation of 2000–5000 fsh in 5–10 m marl, seagrass (Thalassia testudinum and Halodule wrightii) of water (Danylchuk et al. 2011, 2019). At sunset bonefsh and patch reef habitat. To the south of Long Bay the con- begin “porpoising,” where they gulp air at the surface (see tinental shelf narrows, and at 1.5 km ofshore the benthos Danylchuk et al. 2019), and then proceed to move ofshore transitions to a steep vertical drop-of of more than 1000 m. following the edge of the continental shelf (Danylchuk et al. The location of Long Bay provides 180-degree protection 2011, 2019). Similar pre-spawning behavior and potential from NNW to SSE winds and access to the adjacent north ofshore migrations by A. vulpes have been observed in other bay with protection from S winds. Ocean surface currents locations such as Belize (Perez et al. 2019), Cuba (Posada typically fow SSE to NNW with quasi-permanent cyclonic- et al. 2008), and Mexico (Zeng et al. 2019). Additionally, gyres forming on the west-side of Abaco and south of Grand this behavior has also been observed in the congener A. Bahama (Chérubin 2014). glossodonta, in Kiribati (Johannes and Yeeting 2000) and Tetiaroa and Anaa, French Polynesia (Filous et al. 2019a, Fish collection and tag deployment 2020). Spawning behavior of any kind has yet to be docu- mented for the remaining ten albulid species. Ofshore spawning migrations from the PSA site were Data describing the ofshore spawning movements of actively tracked with acoustic telemetry in 2013 (see bonefsh are limited, but of high importance due to the eco- Danylchuk et al. 2019), 2018, and 2019. Active track- nomic value and conservation needs of the species and genus ing methods were chosen rather than the deployment of a 1 3 Marine Biology (2020) 167:187 Page 3 of 11 187 passive acoustic telemetry array due to the logistical and Data collection and analysis technological constraints of working in ofshore waters of > 1000 m, and experience gained by previous eforts Fish were actively tracked using a Vemco VH-110 direc- detailed in Danylchuk et al. (2011, 2019). Methods for fsh tional hydrophone connected to a Vemco VR100 receiver collection were maintained throughout the study, while tag and deck box (InnovaSea Systems Inc., Massachusetts).