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Present study evaluated the impacts of adult density on key biological parameters of a tropical estuarine calanoid copepod A. tropica. Egg production, egg hatching success (EHS), adult mortality (%), nauplii cannibalism (% hour−1), population growth and intrinsic rate of population increase in response to five different adult densities viz. 125, 250, 500, 1000 and 2000 adults/L were assessed. The highest individual egg production (IEP, eggs/female/day) was recorded at 125 adults/L treatment while relative egg production (REP, eggs/L/day) was highest at 1000 adults/L. EHS (24 h and 48 h) showed significant difference (p

The last several decades of research have seen a burgeoning of data on the morphology, physiology, and evolutionary history of vertebrate auditory organs. This chapter briefly describes the status of our understanding of ear structure and function and their origins in fish, which hear using their vestibular epithelia, and land vertebrates that early evolved dedicated hearing structures. The various major lineages of land vertebrates—amphibians, lepidosaurs, archosaurs, and mammals—each have unique hearing organs. From humble beginnings as a small epithelium in their common ancestor, each lineage evolved specialized hair-cell populations and divisions of labor that led to highly sensitive and frequency-selective hearing. This chapter covers the origins, morphology, and physiological characteristics of the ears of all major groups.

Field work in Norway was funded by the Carlsberg Foundation and the National Danish Research Council to PTM. The NMFS study was funded by the U.S. Mineral Management Service. MJ is funded by the Marine Alliance for Science and Technology, Scotland, and by a Marie Curie Career Integration Grant. MW was funded by the Danish Council for Independent Research, Natural Science and NAS is currently funded by a EU Horizon 2020 MSC Fellowship. The sperm whale carries a hypertrophied nose that generates powerful clicks for long-range echolocation. However, it remains a conundrum how this bizarrely shaped apex predator catches its prey. Several hypotheses have been advanced to propose both active and passive means to acquire prey, including acoustic debilitation of prey with very powerful clicks. Here we test these hypotheses by using sound and movement recording tags in a fine-scale study of buzz sequences to relate the acoustic behaviour of sperm whales with changes in acceleration in their head region during prey capture attempts. We show that in the terminal buzz phase, sperm whales reduce inter-click intervals and estimated source levels by 1-2 orders of magnitude. As a result, received levels at the prey are more than an order of magnitude below levels required for debilitation, precluding acoustic stunning to facilitate prey capture. Rather, buzzing involves high-frequency, low amplitude clicks well suited to provide high-resolution biosonar updates during the last stages of capture. The high temporal resolution helps to guide motor patterns during occasionally prolonged chases in which prey are eventually subdued with the aid of fast jaw movements and/or buccal suction as indicated by acceleration transients (jerks) near the end of buzzes. Publisher PDF

Toothed whales use intense ultrasonic clicks to echolocate prey and it has been hypothesized that they also acoustically debilitate their prey with these intense sound pulses to facilitate capture. Cephalopods are an important food source for toothed whales, and there has probably been an evolutionary selection pressure on cephalopods to develop a mechanism for detecting and evading sound–emitting toothed whale predators. Ultrasonic detection has evolved in some insects to avoid echolocating bats, and it can be hypothesized that cephalopods might have evolved similar ultrasound detection as an anti–predation measure. We test this hypothesis in the squid Loligo pealeii in a playback experiment using intense echolocation clicks from two squid–eating toothed whale species. Twelve squid were exposed to clicks at two repetition rates (16 and 125 clicks per second) with received sound pressure levels of 199–226 dB re 1 μPa (pp) mimicking the sound exposure from an echolocating toothed whale as it approaches and captures prey. We demonstrate that intense ultrasonic clicks do not elicit any detectable anti–predator behaviour in L. pealeii and that clicks with received levels up to 226 dB re 1 μPa (pp) do not acoustically debilitate this cephalopod species.

Toothed whales and bats have independently evolved biosonar systems to navigate and locate and catch prey. Such active sensing allows them to operate in darkness, but with the potential cost of warning prey by the emission of intense ultrasonic signals. At least six orders of nocturnal insects have independently evolved ears sensitive to ultrasound and exhibit evasive maneuvers when exposed to bat calls. Among aquatic prey on the other hand, the ability to detect and avoid ultrasound emitting predators seems to be limited to only one subfamily of Clupeidae: the Alosinae (shad and menhaden). These differences are likely rooted in the different physical properties of air and water where cuticular mechanoreceptors have been adapted to serve as ultrasound sensitive ears, whereas ultrasound detection in water have called for sensory cells mechanically connected to highly specialized gas volumes that can oscillate at high frequencies. In addition, there are most likely differences in the risk of predation between insects and fish from echolocating predators. The selection pressure among insects for evolving ultrasound sensitive ears is high, because essentially all nocturnal predation on flying insects stems from echolocating bats. In the interaction between toothed whales and their prey the selection pressure seems weaker, because toothed whales are by no means the only marine predators placing a selection pressure on their prey to evolve specific means to detect and avoid them. Toothed whales can generate extremely intense sound pressure levels, and it has been suggested that they may use these to debilitate prey. Recent experiments, however, show that neither fish with swim bladders, nor squid are debilitated by such signals. This strongly suggests that the production of high amplitude ultrasonic clicks serve the function of improving the detection range of the toothed whale biosonar system rather than debilitation of prey.

The frequency range of hearing in fishes and frogs historically has been thought to be confined to relatively low frequencies in comparison to that of mammals. However, within the last 20 years, the audiograms of several fish and frog species have been shown to encompass ultrasonic (US) frequencies. Moreover, these animals have been shown to respond behaviorally to US playbacks. Although the evolution of US detection in these species is still an ongoing topic of study, both fishes and frogs have faced the challenge of producing very high-frequency responses from systems that evolved with low-frequency sensitivity. A short history of the behavioral responses and the electrophysiological mechanisms (when known) underlying the production and reception of US in fishes and frogs is presented, with a focus on the unique experimental approaches that have yielded this surprising upward extension of the hearing ranges of several specialized fishes and frogs.

SUMMARYToothed whales emit high-powered ultrasonic clicks to echolocate a wide range of prey. It may be hypothesized that some of their prey species have evolved capabilities to detect and respond to such ultrasonic pulses in a way that reduces predation, akin to the situation for many nocturnal insects and echolocating bats. Using high-speed film recordings and controlled exposures, we obtained behavioural evidence that simulated toothed whale biosonar clicks elicit highly directional anti-predator responses in an ultrasound-sensitive allis shad (Alosa alosa). Ten shad were exposed to 192 dB re. 1 μPa (pp) clicks centred at 40 kHz at repetition rates of 1, 20, 50 and 250 clicks s–1 with summed energy flux density levels of 148, 161, 165 and 172 dB re. 1 μPa2 s. The exposures mimicked the acoustic exposure from a delphinid toothed whale in different phases of prey search and capture. The response times of allis shad were faster for higher repetition rates of clicks with the same sound pressure level. None of the fish responded to a single click, but had median response times of 182, 93 and 57 ms when exposed to click rates of 20, 50 and 250 clicks s–1, respectively. This suggests that the ultrasound detector of allis shad is an energy detector and that shad respond faster when exposed to a nearby fast-clicking toothed whale than to a slow-clicking toothed whale far away. The findings are thus consistent with the hypothesis that shad ultrasound detection is used for reducing predation from echolocating toothed whales.

Lungfishes are the closest living relatives of the tetrapods, and the ear of recent lungfishes resembles the tetrapod ear more than the ear of ray-finned fishes and is therefore of interest for understanding the evolution of hearing in the early tetrapods. The water-to-land transition resulted in major changes in the tetrapod ear associated with the detection of air-borne sound pressure, as evidenced by the late and independent origins of tympanic ears in all of the major tetrapod groups. To investigate lungfish pressure and vibration detection, we measured the sensitivity and frequency responses of five West African lungfish ( Protopterus annectens ) using brainstem potentials evoked by calibrated sound and vibration stimuli in air and water. We find that the lungfish ear has good low-frequency vibration sensitivity, like recent amphibians, but poor sensitivity to air-borne sound. The skull shows measurable vibrations above 100 Hz when stimulated by air-borne sound, but the ear is apparently insensitive at these frequencies, suggesting that the lungfish ear is neither adapted nor pre-adapted for aerial hearing. Thus, if the lungfish ear is a model of the ear of early tetrapods, their auditory sensitivity was limited to very low frequencies on land, mostly mediated by substrate-borne vibrations.

Clupeiform fish species, including the Gulf menhaden (Brevoortia patronus) that belong to the subfamily Alosinae, can detect ultrasound. Clupeiform fishes are unique in that they have specialized gas-filled bullae in the head associated with the ear via the bulla membrane and with the lateral line via the lateral recess membrane. It has been hypothesized that the utricle of the inner ear is responsible for ultrasound detection through a specialized connection to the gas-filled bullae complex. Here, we show that the lateral line and its connection to the gas-filled bullae complex via the lateral recess are involved in ultrasound detection in Gulf menhaden. Removal of a small portion of the lateral line overlying the lateral recess membrane eliminates the ability of Gulf menhaden to detect ultrasound. We further show that the gas-filled bullae vibrates in response to ultrasound, that the gas-filled bullae are necessary for detecting ultrasound, and that the bullae connections to the lateral line via the lateral recess membrane play an important role in ultrasound detection. These results add a new dimension to the role of the lateral line and bullae as part of the ultrasonic detection system in Gulf menhaden

Most fish cannot hear frequencies above 3 kHz, but a few species belonging to the subfamily Alosinae (family Clupeidae) can detect intense ultrasound. The response of adult specimens of the European allis shad (Alosa alosa) to sinusoidal ultrasonic pulses at 70 and 120 kHz is tested. The fish showed an intensity-graded response to the ultrasonic pulses with a response threshold between 161 and 167 dB re 1 µPa (pp) for both frequencies. These response thresholds are similar to thresholds derived from juvenile American shad (Alosa sapidissima) in previous studies, supporting the suggestion that these members of Alosinae have evolved a dedicated ultrasound detector adapted to detect and respond to approaching echolocating toothed whales.

Squid play an important role in bio- mass turnover in marine ecosystems and constitute a food source for ~90% of all echolocating toothed whale species. Nonetheless, it has been hypo- thesized that the soft bodies of squid provide echoes too weak to be detected by toothed whale biosonars, and that only the few hard parts of the squid body may generate significant backscatter. We measured the acoustic backscatter from the common squid Loligo pealeii for signals similar to toothed whale echolocation clicks using an energy detector to mimic the mammalian auditory system. We show that the dorsal target strengths of L. pealeii with mantle lengths between 23 and 26 cm fall in the range from -38 to -44 dB, and that the pen, beak and lenses do not contribute significantly to the backscatter. Thus, the muscular mantle and fins of L. pealeii constitute a sufficient sonar target for individual biosonar detection by toothed whales at ranges between 25 and 325 m, depending on squid size, noise levels, click source levels, and orientation of the ensonified squid. While epipelagic squid must be fast and muscular to catch prey and avoid visual predators, it is hypothesized that some deep-water squid may have adopted passive acoustic crypsis, with a body of low muscle mass and low metabolism that will render them less con- spicuous to echolocating predators.