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Foraging Ecology of Whale Sharks (Rhincodon Typus) Within Bahıa De Los Angeles, Baja California Norte, México

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Foraging Ecology of Whale Sharks (Rhincodon Typus) Within Bahıa De Los Angeles, Baja California Norte, México Fisheries Research 84 (2007) 47–64 Foraging ecology of whale sharks (Rhincodon typus) within Bah´ıa de Los Angeles, Baja California Norte, Mexico´ Jonathan D. Nelson a,∗, Scott A. Eckert b,1 a Department of Marine Science and Environmental Studies, University of San Diego, San Diego, CA, USA b Hubbs-SeaWorld Research Institute, San Diego, CA, USA Abstract The presence of whale sharks (Rhincodon typus)inBah´ıa de Los Angeles, Baja California Norte, Mexico, is a seasonal phenomenon, occurring during the months of June–November, with highest abundance from August to October. The foraging ecology of whale sharks in Bah´ıa de Los Angeles was studied from 28 July to 26 October 1999. During this period, 19 individual whale sharks were identified, including nine males, three females, and seven whose sex was not identified. Feeding by whale sharks was observed on 132 out of 190 sightings. Approximately 80% of the feeding events occurred in areas with ≤10 m bottom depth (mean ± S.D. = 7.0 ± 5.5 m), and were concentrated primarily in the southernmost region of the bay. The highest numbers of feeding events occurred between 12.00 and 16.00 h. Mean (±S.D.) sea surface temperature during these events was 29.7 ± 1.1 ◦C. Three feeding behaviors, “active”, “vertical”, and “passive”, and one non-feeding behavior, “cruising”, were characterized. Analysis of plankton samples collected beside feeding sharks revealed that zooplankton, primarily copepods (∼85% of total zooplankton abundance) appeared to be the primary prey source in the bay. Although whale sharks were observed feeding among large schools of baitfish (sardines, anchovies), whale sharks were never observed directly preying upon these fish. Whale sharks may target baitfish to locate zooplankton, as these fish can be indicators of plankton rich patches. Zooplankton abundance was significantly different among the three feeding behaviors, suggesting that prey abundance may influence which feeding technique was utilized. Feeding was not observed when the minimum density of zooplankton was less than ∼10.0 × 103 individuals m−3. Whale sharks may be following oceanographic cues (physical and biological) both within the bay as well as throughout the Gulf of California that are favorable for increases of prey resources. Because whale shark ecotourism is rapidly increasing in Bah´ıa de Los Angeles, it is important to identify and manage ecologically important areas utilized by whale sharks within the bay to ensure the continued integrity of the habitat that supports whale shark presence. © 2006 Elsevier B.V. All rights reserved. Keywords: Rhincodon typus; Whale shark; Distribution; Feeding; Zooplankton abundance; Conservation; Behavior 1. Introduction Whale sharks, basking sharks (Cetorhinus maximus), and megamouth sharks (Megachasma pelagios) are the only three The whale shark, Rhincodon typus, is the world’s largest fish, filter feeding sharks that occur in the world’s oceans (Compagno, yet its natural history and geographic distribution are poorly 1984). These sharks do not capture prey using their teeth, but understood. Most published literature consists of anecdotal instead filter large amounts of water over their specially adapted reports; descriptions of sightings; limited observations of gen- gills and primarily catch planktonic organisms. Ram filter feed- eral biology, foraging, and behavior; and speculative reviews of ing is a technique utilized by all three species, however, whale distribution and movement patterns (Wolfson and Notarbartolo sharks additionally can suction filter feed. This process involves di Sciara, 1981; Colman, 1997). drawing water into the mouth and forcing it out through dense filters covering the internal gills with no forward movement (Gudger, 1941a; Compagno, 1984). In addition, whale sharks ∗ Corresponding author. Current address: California Department of Fish and have been observed using a feeding technique that appears to Game, 830 S Street, Sacramento CA 95814, USA. combine ram and suction filter feeding (Clark and Nelson, 1997; E-mail addresses: [email protected] (J.D. Nelson), Heyman et al., 2001; Plues, unpublished data). The ability to [email protected] (S.A. Eckert). 1 Current address: WIDECAST, Nicholas School of the Environment Marine utilize several foraging techniques may allow whale sharks to Laboratory, Duke University, 135 Duke Marine Lab Road, Beaufort, NC, 28516, capture a wider range of prey compared to basking and meg- USA. amouth sharks (Compagno, 1984). 0165-7836/$ – see front matter © 2006 Elsevier B.V. All rights reserved. doi:10.1016/j.fishres.2006.11.013 48 J.D. Nelson, S.A. Eckert / Fisheries Research 84 (2007) 47–64 Most studies on prey composition of whale sharks have been bivalves) (Plues, unpublished data). Although whale sharks have limited to anecdotal observations and analysis of stomach con- been observed in the surface waters of Bah´ıa de Los Angeles for tents from fishery-killed or stranded animals (Gudger, 1941b; the past decade, the only published research conducted in this Springer, 1957; Silas and Rajagopalan, 1963; Karbhari and area was by Eckert and Stewart (2001) who used satellite teleme- Josekutty, 1986; Satyanarayana Rao, 1986; Silas, 1986). Prey try to study the movements, submergence behavior, and thermal items documented in the literature include planktonic species preferences of whale sharks in the Sea of Cortez. Their research such as copepods, shrimps, and crab larvae, as well as nektonic illustrated that Bah´ıa de Los Angeles is a viable area to conduct species such as sardines, anchovies, small tuna, albacore, and studies on habitat and food utilization, distribution patterns and squid (Compagno, 1984; Last and Stevens, 1994). Other studies local movement. have observed whale sharks targeting primarily planktonic prey The goals of this project were to examine the foraging ecol- items. In Western Australia along Ningaloo Reef, whale sharks ogy of whale sharks in Bah´ıa de Los Angeles. Data on whale have been observed foraging among large amounts of tropical sharks morphology and behavior were collected to characterize krill (Pseudeuphausia latifrons)(Taylor, 1994, 1996; Wilson and foraging and non-foraging strategies. Plankton samples were Newbound, 2001; Wilson et al., 2001). In Belize, off the Glad- collected during whale shark foraging events to determine prey den Spit, whale sharks have been observed foraging within the abundance and taxonomic composition and were compared to clouds of gametes during spawning events of cubara snapper samples collected at designated stations throughout the bay. In (Lutjanus cyanopterus) and dog snapper (L. jocu)(Heyman et addition, whale shark ecotourism was monitored to determine al., 2001). Clark and Nelson (1997) observed whale sharks for- impacts of human interactions on natural behaviors. aging in dense patches of copepods (Acartia clause), in Bah´ıa de la Paz in the Gulf of California, Mexico. 2. Methods Whale sharks have been observed between June and Novem- ber in Bah´ıa de Los Angeles, located in the northwestern Gulf Bah´ıa de Los Angeles (28◦51–29◦03N, 113◦26– of California (Sea of Cortez), Baja California, Mexico (Eckert 113◦36W) is located on the eastern coast of Baja Cali- and Stewart, 2001; Plues, unpublished data; Resendiz, per- fornia Norte, approximately 600 km south of the U.S.-Mexican sonal communication). Their presence appears to coincide with border (Fig. 1). The bay opens north toward the Ballenas late summer increases in zooplankton abundance. Preliminary Channel and water exchange between the bay and channel research conducted in 1998 on whale shark foraging revealed allows the mixing of cold, nutrient rich water with the warmer that the primary prey targeted in Bah´ıa de Los Angeles were waters of the bay, supporting increased primary productivity for small zooplankton, including copepods, shrimps, euphausi- the area (Bustos-Serrano et al., 1996; Delgadillo-Hinojosa et ids, mysids, crab larvae, and mollusc larvae (gastropods and al., 1997). A series of small islands creates a boundary between Fig. 1. Map of Baja California including the Gulf of California and the whale shark study site within Bah´ıa de Los Angeles. Coordinates indicate the latitude and longitude of the study site. J.D. Nelson, S.A. Eckert / Fisheries Research 84 (2007) 47–64 49 Fig. 2. Study site at Bah´ıa de Los Angeles. The dashed line represents the division created for this project, distinguishing North Bay and South Bay. The North Bay study site extended from Punta la Gringa to Punta la Herradura and was limited to the western side of the Islands including Borrego, Ventana, and Cabeza de Caballo. Sampling stations (1–16) are represented by stars and station #. Stations 01–08 are located in South Bay and 09–16 in North Bay. Numbers indicate depth contours (m). Distinctive landmarks also are indicated. the bay and the Ballenas Channel. This study covered the region period of time sufficient to permit the recording of behavioral inside these islands, an area approximately 17 km long by and morphological data. 8 km wide (Fig. 2). For logistical purposes, the study site was Observations were carried out during daylight hours divided into a northern and southern region along an east–west (06.00–19.00 h). Throughout the field season, an equal amount line from Punta Arena to Punta la Herradura, creating almost of time was allocated for observing shark activities in both North equal areas of approximately 45 km2 (Fig. 2). We denoted these and South Bay, usually on alternating days. However, if sharks regions as North and South Bay. Research was conducted from were concentrated in a particular location, that area was mon- 28 July through 26 October 1999. itored daily until sharks were no longer present, after which Throughout the text, the terms “sighting” and “observation” alternate day rotations resumed. are used. Sighting refers to detection, visually or acoustically, When a whale shark was sighted, the location was recorded of an individual whale shark. Observation refers to the sighting using a global positioning system (GPS) receiver, and the time and subsequent visual monitoring of an individual shark for a noted.
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