Echolocation-based foraging by harbor porpoises and sperm whales, including effects of noise and acoustic propagation

Submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy at the Massachusetts Institute of Technology and the Woods Hole Oceanographic Institution September 2008.In this thesis, I provide quantitative descriptions of toothed whale echolocation and foraging behavior, including assessment of the effects of noise on foraging behavior and the potential influence of ocean acoustic propagation conditions on biosonar detection ranges and whale noise exposure. In addition to presenting some novel basic science findings, the case studies presented in this thesis have implications for future work and for management. In Chapter 2, I describe the application of a modified version of the Dtag to studies of harbor porpoise echolocation behavior. The study results indicate how porpoises vary the rate and level of their echolocation clicks during prey capture events; detail the differences in echolocation behavior between different animals and in response to differences in prey fish; and show that, unlike bats, porpoises continue their echolocation buzz after the moment of prey capture. Chapters 3-4 provide case studies that emphasize the importance of applying realistic models of ocean acoustic propagation in marine mammal studies. These chapters illustrate that, although using geometric spreading approximations to predict communication/target detection ranges or noise exposure levels is appropriate in some cases, it can result in large errors in other cases, particularly in situations where refraction in the water column or multi-path acoustic propagation are significant. Finally, in Chapter 5, I describe two methods for statistical analysis of whale behavior data, the rotation test and a semi-Markov chain model. I apply those methods to test for changes in sperm whale foraging behavior in response to airgun noise exposure. Test results indicate that, despite the low-level exposures experienced by the whales in the study, some (but not all) of them reduced their buzz production rates and altered other foraging behavior parameters in response to the airgun exposure.Work presented in this thesis was supported by a National Science Foundation Graduate Research Fellowship, the WHOI Ocean Life Institute (Grant Numbers 32031300 and 25051351), the Office of Naval Research, the U.S. Department of the Interior Minerals Management Service (Cooperative Agreement Numbers 1435-01-02-CA-85186 and NA87RJ0445; WHOI Grant Number 15205601), the Industry Research Funding Coalition, and the WHOI/MIT Joint Program in Oceanography/Applied Ocean Science & Engineering (including a Fye Teaching Fellowship)

Dumb-bell galaxies in southern clusters: Catalog and preliminary statistical results

The dominant galaxy of a rich cluster is often an object whose formation and evolution is closely connected to the dynamics of the cluster itself. Hoessel (1980) and Schneider et al. (1983) estimate that 50 percent of the dominant galaxies are either of the dumb-bell type or have companions at projected distances less than 20 kpc, which is far in excess of the number expected from chance projection (see also Rood and Leir 1979). Presently there is no complete sample of these objects, with the exception of the listing of dumb-bell galaxies in BM type I and I-II clusters in the Abell statistical sample of Rood and Leir (1979). Recent dynamical studies of dumb-bell galaxies in clusters (Valentijn and Casertano, 1988) still suffer from inhomogeneity of the sample. The fact that it is a mixture of optically and radio selected objects may have introduced an unknown biases, for instance if the probability of radio emission is enhanced by the presence of close companions (Stocke, 1978, Heckman et al. 1985, Vettolani and Gregorini 1988) a bias could be present in their velocity distribution. However, this situation is bound to improve: a new sample of Abell clusters in the Southern Hemisphere has been constructed (Abell et al., 1988 hereafter ACO), which has several advantages over the original northern catalog. The plate material (IIIaJ plates) is of better quality and reaches fainter magnitudes. This makes it possible to classify the cluster types with a higher degree of accuracy, as well as to fainter magnitudes. The authors therefore decided to reconsider the whole problem constructing a new sample of dumb-bell galaxies homogeneously selected from the ACO survey. Details of the classification criteria are given

First direct measurements of behavioural responses by Cuvier's beaked whales to mid-frequency active sonar

Most marine mammal­ strandings coincident with naval sonar exercises have involved Cuvier's beaked whales (Ziphius cavirostris). We recorded animal movement and acoustic data on two tagged Ziphius and obtained the first direct measurements of behavioural responses of this species to mid-frequency active (MFA) sonar signals. Each recording included a 30-min playback (one 1.6-s simulated MFA sonar signal repeated every 25 s); one whale was also incidentally exposed to MFA sonar from distant naval exercises. Whales responded strongly to playbacks at low received levels (RLs; 89–127 dB re 1 µPa): after ceasing normal fluking and echolocation, they swam rapidly, silently away, extending both dive duration and subsequent non-foraging interval. Distant sonar exercises (78–106 dB re 1 µPa) did not elicit such responses, suggesting that context may moderate reactions. The observed responses to playback occurred at RLs well below current regulatory thresholds; equivalent responses to operational sonars could elevate stranding risk and reduce foraging efficiency.Publisher PDFPeer reviewe

The Development of the Vascular System in Quail Embryos: A Combination of Microvascular Corrosion Casts and Immunohistochemical Identification

Although vascular casts, obtained by injection with methacrylates, are frequently used to investigate the adult vascular system, little data are available for embryonic stages. In this paper we use MercoxR in quail embryos in the period of 2 to 7 days after incubation. The microvascular corrosion casts were evaluated in the scanning electron microscope (SEM) with special attention to the development and remodelling of the large arteries and veins. Our results show that the remodelling of the large arteries and veins together with their developing tributary vessels can be visualized from very early embryonic stages onwards. However, complete replication of a developing vascular system depends on diameter and regularity of the lumen. In the stages investigated, the vascular lumen, even of the largest vessels, is still very irregular. Detailed cellular characteristics like nuclear impressions of endothelial cells, as often seen in adult material, were seldom found in the embryos. To examine whether blind-ending sprouts are completely or incompletely replicated in a developing vascular system, additional series of quail embryos were stained immunohistochemically with a monoclonal antibody (MB1) specific for endothelial and hemopoietic cells. It seems that a plexus consisting of endothelial precursors (endothelial cells lacking a lumen) is present in the developing organ before the formation of a lumen and assembly into vessels, which are connected to an adjacent artery or vein. Expansion of the vascular system may in part be due to incorporation of these endothelial precursors in the wall of existing vessels

Beluga whale (Delphinapterus leucas) acoustic foraging behavior and applications for long term monitoring

This paper is not subject to U.S. copyright. The definitive version was published in Castellote, M., Mooney, A., Andrews, R., Deruiter, S., Lee, W.-J., Ferguson, M., & Wade, P. Beluga whale (Delphinapterus leucas) acoustic foraging behavior and applications for long term monitoring. Plos One, 16(11), (2021): e0260485, https://doi.org/10.1371/journal.pone.0260485.Cook Inlet, Alaska, is home to an endangered and declining population of 279 belugas (Delphinapterus leucas). Recovery efforts highlight a paucity of basic ecological knowledge, impeding the correct assessment of threats and the development of recovery actions. In particular, information on diet and foraging habitat is very limited for this population. Passive acoustic monitoring has proven to be an efficient approach to monitor beluga distribution and seasonal occurrence. Identifying acoustic foraging behavior could help address the current gap in information on diet and foraging habitat. To address this conservation challenge, eight belugas from a comparative, healthy population in Bristol Bay, Alaska, were instrumented with a multi-sensor tag (DTAG), a satellite tag, and a stomach temperature transmitter in August 2014 and May 2016. DTAG deployments provided 129.6 hours of data including foraging and social behavioral states. A total of 68 echolocation click trains ending in terminal buzzes were identified during successful prey chasing and capture, as well as during social interactions. Of these, 37 click trains were successfully processed to measure inter-click intervals (ICI) and ICI trend in their buzzing section. Terminal buzzes with short ICI (minimum ICI <8.98 ms) and consistently decreasing ICI trend (ICI increment range <1.49 ms) were exclusively associated with feeding behavior. This dual metric was applied to acoustic data from one acoustic mooring within the Cook Inlet beluga critical habitat as an example of the application of detecting feeding in long-term passive acoustic monitoring data. This approach allowed description of the relationship between beluga presence, feeding occurrence, and the timing of spawning runs by different species of anadromous fish. Results reflected a clear preference for the Susitna River delta during eulachon (Thaleichthys pacificus), Chinook (Oncorhynchus tshawytscha), pink (Oncorhynchus gorbuscha), and coho (Oncorhynchus kisutch) salmon spawning run periods, with increased feeding occurrence at the peak of the Chinook and pink salmon runs.Project funding was provided by Georgia Aquarium, the Marine Mammal Laboratory of the Alaska Fisheries Science Center (MML/AFSC). Tagging was funded by the NOAA Fisheries Office of Science and Technology’s Ocean Acoustics Program. DTAG data analysis was funded by the U.S. Marine Mammal Commission grant #16-239. Funding for collecting and analyzing Cook Inlet beluga acoustic data in Susitna Delta was provided by the National Marine Fisheries Service Section 6 Office to the Alaska Department of Fish and Game. This publication is partially funded by the Cooperative Institute for Climate, Ocean, and Ecosystem Studies (CICOES), University of Washington, under NOAA Cooperative Agreement NA15OAR4320063, Contribution No. 2021-1145

Behavioral responses of fin whales to military mid-frequency active sonar

Funding. Primary funding for the SOCAL-BRS project was initially provided by the US Navy’s Chief of Naval Operations Environmental Readiness Division and subsequently by the US Navy’s Living Marine Resources (LMR) Program. Additional support for environmental sampling and logistics was also provided by the Office of Naval Research, Marine Mammal Program.The effect of active sonars on marine mammal behavior is a topic of considerable interest and scientific investigation. Some whales, including the largest species (blue whales, Balaenoptera musculus), can be impacted by mid-frequency (1-10 kHz) military sonars. Here we apply complementary experimental methods to provide the first experimentally controlled measurements of behavioral responses to military sonar and similar stimuli for a related endangered species, fin whales (Balaenoptera physalus). Analytical methods include: (1) Principal Component Analysis paired with Generalized Additive Mixed Models; (2) Hidden Markov Models; and (3) structured expert elicitation using response severity metrics. These approaches provide complementary perspectives on the nature of potential changes within and across individuals. Behavioral changes were detected in five of 15 whales during controlled exposure experiments (CEEs) using mid-frequency active sonar (MFAS) or pseudorandom noise (PRN) of similar frequency, duration, and source and received level. No changes were detected during six control (no noise) sequences. Overall responses were more limited in occurrence, severity, and duration than in blue whales and were less dependent upon contextual aspects of exposure and more contingent upon exposure received level. Quantifying the factors influencing marine mammal responses to sonar is critical in assessing and mitigating future impacts.Publisher PDFPeer reviewe

Mechanisms behind surface modification of polypropylene film using an atmospheric-pressure plasma jet

Plasma treatments are common for increasing the surface energy of plastics, such as polypropylene (PP), to create improved adhesive properties. Despite the significant differences in plasma sources and plasma properties used, similar effects on the plastic film can be achieved, suggesting a common dominant plasma constituent and underpinning mechanism. However, many details of this process are still unknown. Here we present a study into the mechanisms underpinning surface energy increase of PP using atmospheric-pressure plasmas. For this we use the effluent of an atmospheric-pressure plasma jet (APPJ) since, unlike most plasma sources used for these treatments, there is no direct contact between the plasma and the PP surface; the APPJ provides a neutral, radical-rich environment without charged particles and electric fields impinging on the PP surface. The APPJ is a RF-driven plasma operating in helium gas with small admixtures of O2 (0-1%), where the effluent propagates through open air towards the PP surface. Despite the lack of charged particles and electric fields on the PP surface, measurements of contact angle show a decrease from 93.9° to 70.1° in 1.4 s and to 35° in 120 s, corresponding to a rapid increase in surface energy from 36.4 mN m-1 to 66.5 mN m-1 in the short time of 1.4 s. These treatment effects are very similar to what is found in other devices, highlighting the importance of neutral radicals produced by the plasma. Furthermore, we find an optimum percentage of oxygen of 0.5% within the helium input gas, and a decrease of the treatment effect with distance between the APPJ and the PP surface. These observed effects are linked to two-photon absorption laser-induced fluorescence spectroscopy (TALIF) measurements of atomic oxygen density within the APPJ effluent which show similar trends, implying the importance of this radical in the surface treatment of PP. Analysis of the surface reveals a two stage mechanism for the production of polar bonds on the surface of the polymer: a fast reaction producing carboxylic acid, or a similar ketone, followed by a slower reaction that includes nitrogen from the atmosphere on the surface, producing amides from the ketones

A rotation test for behavioural point-process data

Author Posting. © Elsevier B.V., 2008. This is the author's version of the work. It is posted here by permission of Elsevier B.V. for personal use, not for redistribution. The definitive version was published in Animal Behaviour 76 (2008): 1429-1434, doi:10.1016/j.anbehav.2008.06.016.A common problem in animal behavior is determining whether the rate at which a certain behavioural event occurs is affected by an environmental or other factor. In the example considered later in this paper, the event is a vocalization by an individual sperm whale and the factor is the operation or non-operation of an underwater sound source. A typical experiment to test for such effects involves observing animals during control and treatment periods and recording the times of the events that occur in each. In statistical terminology, the data arising from such an experiment – the times at which events of a specified type occur – represent a point process (Cox & Lewis 1978). Events in a point process are treated as having no duration. Although this is not strictly correct for behavioural events, the approximation is reasonable when the duration of events is small in relation to the interval between them.Funding for the sperm whale experiments was provided by the Office of Naval Research, the U.S. Department of the Interior Minerals Management Service Cooperative Agreements Nos. 1435-01-02-CA-321 85186 and NA87RJ0445, and the Industry Research Funding Coalition

Dynamic Analysis of Vascular Morphogenesis Using Transgenic Quail Embryos

Background: One of the least understood and most central questions confronting biologists is how initially simple clusters or sheet-like cell collectives can assemble into highly complex three-dimensional functional tissues and organs. Due to the limits of oxygen diffusion, blood vessels are an essential and ubiquitous presence in all amniote tissues and organs. Vasculogenesis, the de novo self-assembly of endothelial cell (EC) precursors into endothelial tubes, is the first step in blood vessel formation [1]. Static imaging and in vitro models are wholly inadequate to capture many aspects of vascular pattern formation in vivo, because vasculogenesis involves dynamic changes of the endothelial cells and of the forming blood vessels, in an embryo that is changing size and shape. Methodology/Principal Findings: We have generated Tie1 transgenic quail lines Tg(tie1:H2B-eYFP) that express H2B-eYFP in all of their endothelial cells which permit investigations into early embryonic vascular morphogenesis with unprecedented clarity and insight. By combining the power of molecular genetics with the elegance of dynamic imaging, we follow the precise patterning of endothelial cells in space and time. We show that during vasculogenesis within the vascular plexus, ECs move independently to form the rudiments of blood vessels, all while collectively moving with gastrulating tissues that flow toward the embryo midline. The aortae are a composite of somatic derived ECs forming its dorsal regions and the splanchnic derived ECs forming its ventral region. The ECs in the dorsal regions of the forming aortae exhibit variable mediolateral motions as they move rostrally; those in more ventral regions show significant lateral-to-medial movement as they course rostrally. Conclusions/Significance: The present results offer a powerful approach to the major challenge of studying the relative role(s) of the mechanical, molecular, and cellular mechanisms of vascular development. In past studies, the advantages of the molecular genetic tools available in mouse were counterbalanced by the limited experimental accessibility needed for imaging and perturbation studies. Avian embryos provide the needed accessibility, but few genetic resources. The creation of transgenic quail with labeled endothelia builds upon the important roles that avian embryos have played in previous studies of vascular development