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1

Jansen van Vuuren, B., P. B. Best, J. P. Roux, and T. J. Robinson. "Phylogeographic population structure in the Heaviside's dolphin (Cephalorhynchus heavisidii): conservation implications." Animal Conservation 5, no. 4 (November 2002): 303–7. http://dx.doi.org/10.1017/s1367943002004055.

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2

Martin, Morgan J., Tess Gridley, Simon H. Elwen, and Frants H. Jensen. "Heaviside's dolphins ( Cephalorhynchus heavisidii ) relax acoustic crypsis to increase communication range." Proceedings of the Royal Society B: Biological Sciences 285, no. 1883 (July 18, 2018): 20181178. http://dx.doi.org/10.1098/rspb.2018.1178.

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The costs of predation may exert significant pressure on the mode of communication used by an animal, and many species balance the benefits of communication (e.g. mate attraction) against the potential risk of predation. Four groups of toothed whales have independently evolved narrowband high-frequency (NBHF) echolocation signals. These signals help NBHF species avoid predation through acoustic crypsis by echolocating and communicating at frequencies inaudible to predators such as mammal-eating killer whales. Heaviside's dolphins ( Cephalorhynchus heavisidii ) are thought to exclusively produce NBHF echolocation clicks with a centroid frequency around 125 kHz and little to no energy below 100 kHz. To test this, we recorded wild Heaviside's dolphins in a sheltered bay in Namibia. We demonstrate that Heaviside's dolphins produce a second type of click with lower frequency and broader bandwidth in a frequency range that is audible to killer whales. These clicks are used in burst-pulses and occasional click series but not foraging buzzes. We evaluate three different hypotheses and conclude that the most likely benefit of these clicks is to decrease transmission directivity and increase conspecific communication range. The expected increase in active space depends on background noise but ranges from 2.5 (Wenz Sea State 6) to 5 times (Wenz Sea State 1) the active space of NBHF signals. This dual click strategy therefore allows these social dolphins to maintain acoustic crypsis during navigation and foraging, and to selectively relax their crypsis to facilitate communication with conspecifics.
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3

Martin, Morgan J., Simon H. Elwen, Reshma Kassanjee, and Tess Gridley. "To buzz or burst-pulse? The functional role of Heaviside's dolphin, Cephalorhynchus heavisidii, rapidly pulsed signals." Animal Behaviour 150 (April 2019): 273–84. http://dx.doi.org/10.1016/j.anbehav.2019.01.007.

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4

Caputo, M., S. Elwen, T. Gridley, SA Kohler, JP Roux, PW Froneman, and JJ Kiszka. "Dietary plasticity of two coastal dolphin species in the Benguela upwelling ecosystem." Marine Ecology Progress Series 669 (July 8, 2021): 227–40. http://dx.doi.org/10.3354/meps13755.

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Defining the trophic relationships of marine predators and their dietary preferences is essential in understanding their role and importance in ecosystems. Here we used stable isotope analysis of skin samples (δ15N values reflecting trophic level and δ13C values reflecting foraging habitat) to investigate resource partitioning and spatial differences of the feeding ecology of dusky dolphins Lagenorhynchus obscurus and Heaviside’s dolphins Cephalorhynchus heavisidii from 2 coastal study sites separated by 400 km along the coast of central (Walvis Bay) and southern (Lüderitz) Namibia in the Benguela upwelling ecosystem. Overall, isotopic niches of both predators were significantly different, indicating partitioning of resources and foraging habitats. Despite their smaller body size, Heaviside’s dolphins fed at a significantly higher trophic level than dusky dolphins. Stable isotope mixing models revealed that both species fed on high trophic level prey (i.e. large Merluccius spp., large Sufflogobius bibarbatus, and Trachurus t. capensis) at Walvis Bay. The diet of both dolphin species included smaller pelagic fish and squid at Lüderitz. Spatial differences highlight that Heaviside’s and dusky dolphins may exhibit dietary plasticity driven by prey availability, and that they likely form distinct population segments. Important prey for both dolphin species, specifically Merluccius spp. and T. t. capensis, are the main target of trawl fisheries in the Benguela upwelling ecosystem, highlighting potential resource overlap between dolphins and fisheries.
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5

Elwen, Simon H., and Ruth H. Leeney. "Injury and Subsequent Healing of a Propeller Strike Injury to a Heaviside's Dolphin (Cephalorhynchus heavisidii)." Aquatic Mammals 36, no. 4 (December 1, 2010): 382–87. http://dx.doi.org/10.1578/am.36.4.2010.382.

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6

GOPAL, Keshni, Leszek KARCZMARSKI, and Krystal A. TOLLEY. "Patterns of geographic variation between mitochondrial and nuclear markers in Heaviside's (Benguela) dolphins ( Cephalorhynchus heavisidii )." Integrative Zoology 14, no. 5 (September 2019): 506–26. http://dx.doi.org/10.1111/1749-4877.12380.

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7

Elwen, Simon, Michael A. Meÿer, Peter B. Best, P. G. H. Kotze, Meredith Thornton, and Stephan Swanson. "RANGE AND MOVEMENTS OF FEMALE HEAVISIDE'S DOLPHINS (CEPHALORHYNCHUS HEAVISIDII), AS DETERMINED BY SATELLITE-LINKED TELEMETRY." Journal of Mammalogy 87, no. 5 (October 2006): 866–77. http://dx.doi.org/10.1644/05-mamm-a-307r2.1.

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8

Elwen, Simon H., Desray Reeb, Meredith Thornton, and Peter B. Best. "A population estimate of Heaviside's dolphins,Cephalorhynchus heavisidii, at the southern end of their range." Marine Mammal Science 25, no. 1 (January 2009): 107–24. http://dx.doi.org/10.1111/j.1748-7692.2008.00246.x.

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9

Elwen, Simon H., Peter B. Best, Desray Reeb, and Meredith Thornton. "Diurnal Movements and Behaviour of Heaviside's Dolphins,Cephalorhynchus heavisidii, with some Comparative Data for Dusky Dolphins,Lagenothynchus obscutus." South African Journal of Wildlife Research 39, no. 2 (October 2009): 143–54. http://dx.doi.org/10.3957/056.039.0204.

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10

Elwen, S. H., M. Thornton, D. Reeb, and P. B. Best. "Near-Shore Distribution of Heaviside's (Cephalorhynchus heavisidii) and Dusky Dolphins (Lagenorhynchus obscurus) at the Southern Limit of their Range in South Africa." African Zoology 45, no. 1 (April 2010): 78–91. http://dx.doi.org/10.3377/004.045.0103.

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11

Morisaka, Tadamichi, Leszek Karczmarski, Tomonari Akamatsu, Mai Sakai, Steve Dawson, and Meredith Thornton. "Echolocation signals of Heaviside’s dolphins (Cephalorhynchus heavisidii)." Journal of the Acoustical Society of America 129, no. 1 (January 2011): 449–57. http://dx.doi.org/10.1121/1.3519401.

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12

Leeney, Ruth. "Using Static Acoustic Monitoring to Describe Echolocation Behaviour of Heaviside’s Dolphins (Cephalorhynchus heavisidii) in Namibia." Aquatic Mammals 37, no. 2 (June 1, 2011): 151–60. http://dx.doi.org/10.1578/am.37.2.2011.151.

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13

Gridley, T., MJ Martin, J. Slater, J.-P. Roux, RJ Swift, and SH Elwen. "Towed passive acoustic monitoring complements visual survey methods for Heaviside’s dolphins Cephalorhynchus heavisidii in the Namibian Islands Marine Protected Area." African Journal of Marine Science 42, no. 4 (October 1, 2020): 495–506. http://dx.doi.org/10.2989/1814232x.2020.1848925.

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14

Elwen, S. H., M. Thornton, D. Reeb, and P. B. Best. "Near-shore distribution of Heaviside’s (Cephalorhynchus heavisidii) and dusky dolphins (Lagenorhynchus obscurus) at the southern limit of their range in South Africa." African Zoology 45, no. 1 (April 2010): 78–91. http://dx.doi.org/10.1080/15627020.2010.11657256.

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15

Henry, John. "A note on concentrations of metals in cetaceans from southern Africa." J. Cetacean Res. Manage., January 1, 1999, 177–94. http://dx.doi.org/10.47536/jcrm.v1i1.260.

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Concentrations of zinc, copper, cadmium, mercury and lead were measured by atomic absorption spectrophotometry in samples of the brain, kidney , liver and muscle tissue from 178 individuals of 323 different cetacean species (4 right whales - Eubalaena australis, 2 pygmy right whales - Caperea marginara, 3 minke whales - Balaenoptera acutorostrata, 3 Bryde's whales - B. edeni, 1 humpback whale - Megaptera novaeangliae, 1 sperm whale - Physeter macrocephalus, 11 pygmy sperm whale - Kogia breviceps, 6 dwarf sperm whales - K. simus, 1 southern bottlenose whale - Hyperoodon planifrons, 1 Cuvier's beaked whale - Ziphius cavirostris, 9 Blainville's beaked whales - Mesoplodon densirostris, 5 strap-tooth whales - M. layardii, 2 True's beaked whales - M. mirus, 3 long-finned pilot whales - Globicephala me/as, 30 Risso's dolphins - Grampus griseus, 12 bottlenose dolphins - Tursiops truncatus, 5 striped dolphins - Stenella coeruleoalba, 1 pantropical spotted dolphin - S. attenuata, 1 hump-backed dolphin - Sousa chinensis, 21 dusky dolphins - Lagenorhynchus obscurus, 1 hourglass dolphin - L. cruciger, 12 Heaviside's dolphins - Cephalorhynchus heavisidii and 43 common dolphins - Delphinus delphis). All but the hourglass dolphin were strandings or animals taken incidental to fishing operations or under scientific permit in coastal waters of South Africa or Namibia. Highest concentrations of Zn, Cu and Hg were generally found in the liver and of Cd in the kidney. Comparisons of animals pre-and post puberty indicated accumulation of hepatic mercury in the pygmy sperm whale, Risso's dolphin, dusky dolphin and common dolphin. Loss of a metal (zinc) after puberty was only shown in the common dolphin. No individual analyses exceeded proposed (human) tolerance limits for hepatic mercury and hepatic or renal cadmium
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16

Martin, Morgan Jennifer, Tess Gridley, Jean-Paul Roux, and Simon Harvey Elwen. "First Abundance Estimates of Heaviside’s (Cephalorhynchus heavisidii) and Dusky (Lagenorhynchus obscurus) Dolphins Off Namibia Using a Novel Visual and Acoustic Line Transect Survey." Frontiers in Marine Science 7 (September 18, 2020). http://dx.doi.org/10.3389/fmars.2020.555659.

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