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Assessing Seasonality and Density from Passive Acoustic Monitoring of Signals Presumed to Be from Pygmy and Dwarf Sperm Whales in the Gulf of Mexico

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Assessing Seasonality and Density from Passive Acoustic Monitoring of Signals Presumed to Be from Pygmy and Dwarf Sperm Whales in the Gulf of Mexico ORIGINAL RESEARCH published: 27 February 2019 doi: 10.3389/fmars.2019.00066 Assessing Seasonality and Density From Passive Acoustic Monitoring of Signals Presumed to be From Pygmy and Dwarf Sperm Whales in the Gulf of Mexico John A. Hildebrand 1*, Kaitlin E. Frasier 1, Simone Baumann-Pickering 1, Sean M. Wiggins 1, Karlina P. Merkens 2, Lance P. Garrison 3, Melissa S. Soldevilla 3 and Mark A. McDonald 4 1 Scripps Institution of Oceanography, University of California, San Diego, La Jolla, CA, United States, 2 National Marine Fisheries Service, Pacific Island Fisheries Science Center, Honolulu, HI, United States, 3 National Marine Fisheries Service, Southeast Fisheries Science Center, Miami, FL, United States, 4 WhaleAcoustics, Bellvue, CO, United States Pygmy sperm whales (Kogia breviceps) and dwarf sperm whales (Kogia sima) are deep diving cetaceans that commonly strand along the coast of the southeast US, but that are difficult to study visually at sea because of their elusive behavior. Conventional visual surveys are thought to significantly underestimate the presence of Kogia and they Edited by: Lars Bejder, have proven difficult to approach for tracking and tagging. An approach is presented University of Hawaii at Manoa, for density estimation of signals presumed to be from Kogia spp. based on passive United States acoustic monitoring data collected at sites in the Gulf of Mexico (GOM) from the period Reviewed by: Leigh Gabriela Torres, following the Deepwater Horizon oil spill (2010-2013). Both species of Kogia are known Oregon State University, United States to inhabit the GOM, although it is not possible to acoustically separate the two based on Robert McCauley, available knowledge of their echolocation clicks. An increasing interannual density trend is Curtin University, Australia suggested for animals near the primary zone of impact of the oil spill, and to the southeast *Correspondence: John A. Hildebrand of the spill. Densities were estimated based on both counting individual echolocation [email protected] clicks and counting the presence of groups of animals during one-min time windows. Densities derived from acoustic monitoring at three sites are all substantially higher (4–16 Specialty section: 2 This article was submitted to animals/1000 km ) than those that have been derived for Kogia from line transect visual Marine Megafauna, surveys in the same region (0.5 animals/1000 km2). The most likely explanation for the a section of the journal observed discrepancy is that the visual surveys are underestimating Kogia spp. density, Frontiers in Marine Science due to the assumption of perfect detectability on the survey trackline. We present an Received: 14 January 2018 Accepted: 06 February 2019 alternative approach for density estimation, one that derives echolocation and behavioral Published: 27 February 2019 parameters based on comparison of modeled and observed sound received levels at Citation: sites of varying depth. Hildebrand JA, Frasier KE, Baumann-Pickering S, Wiggins SM, Keywords: passive acoustic monitoring, density estimation, pygmy sprm whale, dwarf sperm whale, Gulf of Mexico Merkens KP, Garrison LP, Soldevilla MS and McDonald MA (2019) Assessing Seasonality and INTRODUCTION Density From Passive Acoustic Monitoring of Signals Presumed to be Pygmy sperm whales (Kogia breviceps) and dwarf sperm whales (Kogia sima) are deep diving From Pygmy and Dwarf Sperm Whales in the Gulf of Mexico. cetaceans that are widely distributed in tropical and temperate waters worldwide (Jefferson et al., Front. Mar. Sci. 6:66. 2015). They are typically encountered along the continental slope and in the abyssal plain (Baird doi: 10.3389/fmars.2019.00066 et al., 1996; Baird, 2005). Both species are difficult to observe, being entirely pelagic, with faint blows Frontiers in Marine Science | www.frontiersin.org 1 February 2019 | Volume 6 | Article 66 Hildebrand et al. Kogia GOM Seasonality and Density and showing only a low profile while at the water’s surface data to provide constraints on acoustic signal production and (Jefferson et al., 2015). They are not easily approached using a diving behavior. Building on these we derive population density small boat and have thus far eluded tagging attempts in the field estimates for Kogia spp. at three sites in the GOM, based on (Baird, 2015). A recent study using passive acoustic monitoring the best available information derived from analogy with beaked (Hodge et al., 2018) found that Kogia may be more common than whales. The acoustic methods we present here provide a new suggested by the visual survey record alone. tool for monitoring Kogia spp. and other cryptic populations, The pygmy sperm whale is the larger of the two species a critical aspect of making management recommendations for with a maximum length of 3.5m and weighing up to 410kg, their conservation. while the dwarf sperm whale has a maximum length of 2.7 m and weight of up to 272kg (Mcalpine, 2018). They live in groups of less than 10 individuals with varying age and sex METHODS composition. Group size and their inshore-offshore presence may vary seasonally as documented in the Bahamas (Dunphy-Daly Data Collection et al., 2008). They have relatively short lives with a maximum The data presented here were collected from five deepwater known longevity of 23 years (Willis and Baird, 1998; Jefferson locations in the northern and eastern GOM (Figure 1), an area et al., 2015). Their primary prey is cephalopods (particularly which is an important habitat for a diverse and abundant group of Histioteuthidae and Cranchiidae), but stomach contents also have cetaceans (Davis et al., 2002). The circulation of the northeastern shown consumption of fish and crustaceans (West et al., 2009; GOM is dominated by the Loop Current, an area of warm water Mcalpine, 2014). Based on stomach contents and isotope analysis, that enters the GOM from the Caribbean and exits through the the two Kogia species may feed at different depths and on slightly Florida strait. The oceanographic dynamics of the GOM also different prey (Barros et al., 1998; Willis and Baird, 1998). Kogia include a large freshwater inflow from the Mississippi and other may feed both in the water column and at or near the bottom. rivers, along with their associated nutrients and sediment loads. Their sightings are most frequently reported in water depths The input of nutrients from the Mississippi River creates high between 400 - 1000m, although they are also seen in deeper phytoplankton productivity and subsequently high zooplankton waters (Baumgartner et al., 2001). productivity. Ecosystem dynamics for the deepwater GOM are Much of what we know about Kogia has been inferred from poorly understood, although it is clear that sperm whales, Kogia stranding records (Willis and Baird, 1998; Wursig et al., 2000). and other deepwater cetaceans are important upper trophic Strandings of these two species are relatively common in the level predators. southeastern United States. They were reported to be the second Acoustic data were collected in the GOM using multiple most common cetacean (after bottlenose dolphins) to strand deployments of High-frequency Acoustic Recording Packages from North Carolina to Texas between 1978 and 1987 (Odell, (HARPs) (Wiggins and Hildebrand, 2007) during and following 1991), with a total of 189 animals. In the Gulf of Mexico the 2010 Deepwater Horizon oil spill. HARPs are bottom- (GOM) there appears to be no seasonal pattern for strandings mounted acoustic recorders capable of recording continuously (Caldwell et al., 1960; Delgado-Estrella and Vasquez, 1998). The at high sample rates (up to 320 kHz) for extended periods. For relatively high rate of Kogia spp. strandings suggests they may the GOM deployments, the HARP instrumentation package was have a higher population than indicated by the relatively few located at or near the seafloor with the hydrophone sensor that are observed during visual surveys (Garrison et al., 2010). tethered to the instrument and buoyed approximately 10 m A combined (K. breviceps and K. sima) abundance of 186 (CV above the seafloor. All acoustic data were converted to sound = 1.04) animals is reported within the entire US Gulf of Mexico pressure levels based on hydrophone and electronic system (GOM) exclusive economic zone (Waring et al., 2013). calibrations. The hydrophones were composed of two stages, one On two occasions, pygmy sperm whales in captivity have for low-frequency (<2 kHz) and the other for high-frequency been shown to produce high-frequency, narrow-band clicks (>2 kHz), although we focus on only the high-frequency band with peak frequencies around 125–130 kHz (Marten, 2000; for this paper. The high-frequency stage uses a spherical omni- Ridgway and Carder, 2001; Madsen et al., 2005). Based on directional transducer (ITC-1042, www.itc-transducers.com) field recordings, dwarf sperm whales are known to produce which has an approximately flat (±2 dB) sensitivity response similarly high-frequency clicks (Merkens et al., 2018). The high- of about −200 dBrms re 1V/µPa from 1 Hz to 100 kHz. Each frequency echolocation signals of pygmy and dwarf sperm whales individual hydrophone has a frequency dependent sensitivity are similar to those of phocoenids, cephalorhynchids and two supplied by the manufacturer. The signals from the hydrophone lagenorhynchid species (Au, 1993; Bassett et al., 2009; Kyhn et al., transducer are fed into a preamplifier with approximately 50 2009, 2010); however, none of the latter species are known to dB of gain and a 10-pole low-pass filter to reduce high- occur in the GOM (Wursig et al., 2000). frequency aliasing effects above 100 kHz and digitized with 16- Passive acoustic monitoring in the northern GOM, conducted bits of resolution at 200 kHz sample rate. The response of each in response to the Deepwater Horizon oil spill (Merkens, 2013), preamplifier and filter were measured and these were combined yielded ample detections of high frequency echolocation clicks with the hydrophone sensitivity to create a system transfer that are most likely produced by one or both of the Kogia function unique to each instrument deployment.
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