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| A rare sight of the fast and shy Ganges River Dolphin Platanista g. gangetica.
(Photo by Rubaiyat Mansur, Whale and Dolphin Conservation Society)
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Abstract
Toothed whales (Cetacea, odontoceti) use biosonar to navigate their environment and to find and catch prey. All studied toothed whale species have evolved highly directional, high-amplitude ultrasonic clicks suited for long-range echolocation of prey in open water. Little is known about the biosonar signals of toothed whale species inhabiting freshwater habitats such as endangered river dolphins. To address the evolutionary pressures shaping the echolocation signal parameters of non-marine toothed whales, we investigated the biosonar source parameters of Ganges river dolphins (Platanista gangetica gangetica) and Irrawaddy dolphins (Orcaella brevirostris) within the river systems of the Sundarban mangrove forest. Both Ganges and Irrawaddy dolphins produced echolocation clicks with a high repetition rate and low source level compared to marine species. Irrawaddy dolphins, inhabiting coastal and riverine habitats, produced a mean source level of 195 dB (max 203 dB) re 1 µPapp whereas Ganges river dolphins, living exclusively upriver, produced a mean source level of 184 dB (max 191) re 1 µPapp. These source levels are 1–2 orders of magnitude lower than those of similar sized marine delphinids and may reflect an adaptation to a shallow, acoustically complex freshwater habitat with high reverberation and acoustic clutter. The centroid frequency of Ganges river dolphin clicks are an octave lower than predicted from scaling, but with an estimated beamwidth comparable to that of porpoises. The unique bony maxillary crests found in the Platanista forehead may help achieve a higher directionality than expected using clicks nearly an octave lower than similar sized odontocetes.
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An Ancient Biosonar Sheds New Light On the Evolution of Echolocation in Toothed Whales
reprinted from materials provided by Woods Hole Oceanographic Institution.
— Some 30 million years ago, Ganges river dolphins diverged from other toothed whales, making them one of the oldest species of aquatic mammals that use echolocation, or biosonar, to navigate and find food. This also makes them ideal subjects for scientists working to understand the evolution of echolocation among toothed whales.
New research, led by Frants Havmand Jensen, a Danish Council for Independent Research / Natural Sciences postdoctoral fellow at Woods Hole Oceanographic Institution, shows that freshwater dolphins produce echolocation signals at very low sound intensities compared to marine dolphins, and that Ganges river dolphins echolocate at surprisingly low sound frequencies. The study, "Clicking in shallow rivers," was published in the journal PLOS ONE.
"Ganges River dolphins are one of the most ancient evolutionary branches of toothed whales," says Jensen. "We believe our findings help explain the differences in echolocation between freshwater and marine dolphins. Our findings imply that the sound intensity and frequency of Ganges river dolphin may have been closer to the 'starting point' from which marine dolphins gradually evolved their high-frequent, powerful biosonar."
The scientists believe these differences evolved due to differences in freshwater and marine environments and the location and distribution of prey in those environments.
A complex, underwater environment
To sustain themselves, river dolphins must find their food, often small fish or crustaceans, in highly turbid water where visibility seldom exceeds a few inches. Like their marine relatives, they manage this using echolocation: They continuously emit sound pulses into the environment and listen for the faint echoes reflected off obstacles while paying special attention to the small details in the echoes that might signify a possible meal.
The environment that freshwater dolphins operate in poses very different challenges to a biosonar than the vast expanses of the sea where most dolphins later evolved. "Dolphins that range through the open ocean often feed on patchily distributed prey, such as schools of fish," Jensen says. "They have had a large advantage from evolving an intense biosonar that would help them detect prey over long distances, but we have little idea of how the complex river habitats of freshwater dolphins shape their biosonar signals."
Shy study animals with a surprisingly deep voice
To answer that question, the researchers recorded the echolocation signals of two species of toothed whales inhabiting the same mangrove forest in the southern part of Bangladesh: The Ganges river dolphin, an exclusively riverine species that is actually not part of the dolphin family but rather the Platanistidae family, and the Irrawaddy, a freshwater toothed whale from the dolphin family that lives in both coastal and riverine habitats.
Surprisingly, the echolocation signals turned out to be much less intense than those employed by marine dolphins of similar size and it seemed that the freshwater dolphins were looking for prey at much shorter distances. From this, the researchers surmise that both the dolphin species and the river dolphin were echolocating at short range due to the complex and circuitous river system that they were foraging in.
While both Irawaddy and Ganges river dolphin produced lower intensity biosonar, the Ganges river dolphin had an unexpectedly low frequency biosonar, nearly half as high as expected if this species had been a marine dolphin.
"It is very surprising to see these animals produce such low-frequent biosonar sounds. We are talking about a small toothed whale the size of a porpoise producing sounds that would be more typical for a killer whale or a large pilot whale," says Professor Peter Teglberg Madsen from Aarhus University in Denmark, an expert on toothed whale biosonar and co-author of the study.
A new perspective on the evolution of biosonar
The study suggests that echolocation in toothed whales initially evolved as a short, broadband and low-frequent click. As dolphins and other toothed whales evolved in the open ocean, the need to detect schools of fish or other prey items quickly favored a long-distance biosonar system. As animals gradually evolved to produce and to hear higher sound frequencies, the biosonar beam became more focused and the toothed whales were able to detect prey further away.
However, the Ganges river dolphin separated from other toothed whales early throughout this evolutionary process, adapting to a life in shallow, winding river systems where a high-frequency, long-distance sonar system may have been less important than other factors such as high maneuverability or the flexible neck that helps these animals capture prey at close range or hiding within mangrove roots or similar obstructions.
Improved tools for counting animals
Freshwater dolphins are among the most endangered animal species. Only around a thousand Ganges river dolphins are thought to remain, and they inhabit some of the most polluted and overfished river systems on Earth. The results of this study will help provide local collaborators with a new tool in their struggle to conserve these highly threatened freshwater cetaceans. Using acoustic monitoring devices to identify the local species may help researchers estimate how many animals remain, and to identify what areas are most important to them.
Frants H. Jensen, Alice Rocco, Rubaiyat M. Mansur, Brian D. Smith, Vincent M. Janik, Peter T. Madsen. 2013. Clicking in Shallow Rivers: Short-Range Echolocation of Irrawaddy and Ganges River Dolphins in a Shallow, Acoustically Complex Habitat. PLoS ONE. 8 (4): e59284 DOI: 10.1371/journal.pone.0059284
