The growing industry of whale-watching is allowing increasing numbers of people access to whales in their natural environment, and constitutes a non-consumptive use of the whales compared to whaling. At the same time, questions are often raised about the hidden effects of whale-watching on the whales. A population of gray whales (Eschrichtius robustus) which spends the summer feeding in Clayoquot Sound, on the West Coast of Vancouver Island, is regularly observed by whale-watchers from the nearby tourist centre of Tofino. Concern among whale-watching business operators and tourists about the possible effects of whale-watching on the feeding whales was heightened in the years preceding this study by an apparent northward movement of the whales, taking them farther from the Tofino, the point of departure for whale-watching tours. This study attempts to explain this apparent trend by finding out whether the whales' short-term behaviour is affected by the presence of whale-watching vessels, and by examining their short and long-term behaviour in the wider context of some of the features of their environment. Whales were observed from a small research vessel for three feeding seasons, in five locations within the area known as Clayoquot Sound. The whales' ventilations were recorded continuously and their location and the number of whale-watching vessels present was recorded at regular intervals. The whales' benthic prey was sampled in all three seasons and their planktonic prey in 1995. A series of variables were calculated from the ventilation data and compared to the number of vessels. The whales' dive behaviour was correlated much more strongly with feeding location than with vessel number. Even with these two factors taken into account, much of the variation in their behaviour remains unaccounted for. Because of this, although the behavioural change in the presence of vessels is statistically significant, there is reason to doubt whether it is biologically significant. The effects of feeding location are probably a composite of the effects of depth, prey type and other factors which are difficult to measure. Of the two main components, prey type appears to have a greater effect than depth on gray whale behaviour. Interaction exists between the effects of site and those of whale-watch vessels, meaning that the effects of vessels are different at different sites. The general pattern is that the effects of vessel presence are more pronounced in shallow sites than in deep, although there are some exceptions to this trend. Gray whale prey shows considerable variation in location, density and composition from year to year. The long-term patterns of gray whale habitat use more closely resemble a prey-selection-driven pattern than a pattern of avoidance of whale-watch vessels. The relatively small influence of vessel numbers on gray whale feeding behaviour suggests that the current guidelines in place for whale-watching vessels are effective in limiting disturbance of the whales. The variable nature of their prey supply suggests that gray whales use all the sites in Clayoquot Sound, and that the availability of a diverse selection of prey is necessary for their success in the tertiary feeding grounds. / Graduate

The enzymatic organization of muscle tissue usually is examined in only a select few muscles of any one animal species. However, because the functional demands placed on individual muscles can vary so widely from muscle to muscle, it is inappropriate to generalize findings from one or two muscles to muscle tissue in general. The differences or similarities in metabolic machinery between skeletal muscles of a wide functional range provides crucial information with respect to a particular animals' whole body metabolism. Nowhere is this understanding more important than in the diving marine mammal which must operate as a closed system (with respect to oxygen supply) while submerged. The goals of this thesis are: 1) to provide a broad body of information on the metabolic organization of a large cross-section of marine mammal muscles, both functionally and with regard to location, 2) to assess the implications of the enzyme differences between muscles to the diving habit, and 3) to compare the metabolic organization of skeletal muscle among several species of marine mammal with different diving abilities and habits. A series of 13 enzymes were measured in 21 skeletal muscles of the harbor seal, Phoca vitulina. In addition, 23 enzyme activity ratios were calculated and analyzed for these muscles. A similar analysis of 22 muscles from fin whales, Balaenoptera physalis. was conducted --including 7 key enzymes and 15 activity ratios. Overall, both the maximum activities and the enzyme activity ratios are consistent with the idea that marine mammal muscle is typical mammalian muscle, exhibiting few significant differences from terrestrial species with respect to catabolic enzymes. The only obvious exception to this in the species examined is observed with fin whale locomotory muscle which has extremely high activities of lactate dehydrogenase (over 2000 units/gm wet wt at 25°C) due to an apparent scaling phenomenon. Tight control of this high potential glycolytic flux is indicated by pyruvate kinase activities that scale downward. Comparisons of enzyme relationships between muscles of harbor seals seem to indicate a very aerobically poised metabolic make-up. This is especially true with respiratory and locomotory muscles, which also show a high tendency to utilize fat. This pattern of enzyme activities and activity ratios in the locomotory muscles of harbor seal is evidence that muscle contractile activity while diving is powered primarily through oxidative pathways and largely based on fat as fuel. The majority of non-locomotory muscles appear to be more able to function anaerobically utilizing carbohydrate. This pattern may correlate with circulatory redistributions while diving that preferentially fuel the locomotory muscles with oxygen, leaving the inactive muscles significantly more hypoperfused and, therefore, candidates for energy saving O₂ sparing (metabolic depression). Fin whales exhibit an opposite pattern, with enzyme profiles more typical of "white" muscle. Unlike harbor seals, the locomotory muscles of fin whales are consistently the least oxidatively poised of the muscles examined. This apparently more anaerobic nature of fin whale muscle is possibly complicated by scaling adaptations, but appears to be a real phenomenon. The examination of three to four skeletal muscles from each of three additional phocid seal species from Antarctica, leopard seals (Hydrurga leptonyx). crab-eater seals (Lobodon carcinophagus). and Weddell seals (Leptonychotes weddelli) confirm that the harbor seal pattern of enzyme profiles is fairly consistent among phocid seals. By these criteria skeletal muscles of phocid seals (particularly the locomotory and respiratory muscles) appear to be designed for sustained aerobic metabolism during diving regardless of the habits or diving capabilities of the seal. / Science, Faculty of / Zoology, Department of / Graduate

Data collected from Pribilof far seals, Callorhinus ursinus, on land (1911-89) and at sea (1958-74) are analyzed to establish biological relationships and distinguish ecological patterns that are relevant to understanding and managing northern fur seal populations. The thesis follows the development of the fur seal from conception and birth through to sexual maturity and finally to a synthesis of the earlier material in terms of population regulation, management, and reasons for the decline of the Pribilof herd. Growth curves show that male fetuses grow faster and larger than female fetuses, and that fetal size is influenced by the age, size, and reproductive history of the mother. Juvenile and adult fur seals experience pronounced seasonal increases and decreases in body length and mass. Rapid gains in mass and growth occur during a brief 1-3 month period as the population migrates through the coastal waters of northern British Columbia and Alaska on its way to the Pribilof Islands. Body mass is gradually lost during the rest of the year while fasting on land and wintering along the coasts of Washington, Oregon, and California. The timing of migration and pupping is highly synchronized from year to year and may be related to the effect of climatic conditions on pup survival during the breeding season. Predictions from a thermal budget developed for pups and the results of a seasonal decomposition of weather patterns on the Pribilof Islands show that the synchronism of births in early July corresponds to the start of three months of conditions that are optimal for growth and survival of pups. Long term fluctuations are noted in pup mass and subadult growth rates which may be related to underlying, large scale natural changes in prey abundance. Changes in the physiological condition (body growth) and vital rates (survival and reproduction) are analyzed for the period 1911-89 as the population increased and decreased. Few density dependent relationships could be demonstrated. Two hypotheses concerning the current decline of the Pribilof population are reviewed and a new, third hypothesis is proposed. The thesis also examines biases in data collection related to the effects of tagging and the handling of fur seals and outlines some directions for future research. / Science, Faculty of / Zoology, Department of / Graduate

Organochlorines are ubiquitous pollutants of the marine environment. These lipid-soluble and highly persistent compounds are found in detectable amounts in almost all marine organisms, and accumulate in the lipid tissues of marine animals. This bioaccumulation leads to biomagnification of these contaminants in higher trophic levels. Near the top of many marine food chains are found the marine mammals, in whose blubber high levels of organochlorine residues have been measured. The most commonly occurring of these pollutants in these animals are the polychlorinated biphenyls (PCB's) and p,p-DDE, a metabolite of the insecticide DDT. These substances have been shown to cause disruptions in the endocrine, immune, and reproductive systems, and are passed from mother to offspring through the placenta and by lactation. Presence and levels of residues of these compounds are, therefore, monitored in marine mammals to provide an indication of the health of a given population and the environment in which they live. Such monitoring is generally done with the use of gas chromatography (GC). High performance liquid chromatography (HPLC) is little used due to the poor ultraviolet (UV) absorbance properties of many of the organochlorines. PCB's and p,p-DDE do absorb UV well enough at concentrations usually encountered in marine mammals to permit the use of HPLC for detection and semiquantification of these substances. A method was developed for the screening of blubber of marine mammals for total PCB's and p,p-DDE using HPLC. The method was applied to the detection and approximation of levels of these two organochlorines in marine mammals from the east and west coasts of the United States. Geographical differences in levels of the two pollutants were found, indicating differences in primary feeding ranges. Evidence of placental transfer of these two organochlorines was also found. Especially high residue levels were found in the blubber of stranded killer whales, indicating that acquisition of high pollutant burdens is still a problem in these top predators. It was concluded that HPLC can be used to screen marine mammals for PCB's and p,p-DDE, and that residue levels determined can be useful in investigating species range, pollutant burdens, and health of populations.

Understanding the physiological response of marine mammals to anthropogenic stressors can inform marine ecosystem conservation strategies. Stress stimulates release of glucocorticoid (GC) hormones, which increase energy substrate availability while suppressing energy-intensive processes. Exposure to repeated stressors can potentially affect an animal’s ability to respond to and recover from subsequent challenges. To assess the endocrine response of a marine mammal to repeated stressors, we administered adrenocorticotropic hormone (ACTH) to free-ranging juvenile northern elephant seals (Mirounga angustirostris; n=7) once daily for four days. ACTH administration induced significant, but transient (<24 h) elevation in circulating cortisol levels (p < 0.0001). These increases did not vary in magnitude between the first ACTH challenge on day 1 and the last challenge on day 4. In contrast, aldosterone levels remained elevated above baseline for at least 24 hours after each ACTH injection (p < 0.001), and responses were greater on day 4 than day 1 (p < 0.01). Total triiodothyronine (tT3) levels were decreased on day 4 relative to day 1 (p < 0.01), while reverse triiodothyronine (rT3) concentrations increased relative to baseline on days 1 and 4 (p < 0.001) in response to ACTH, indicating a suppression of thyroid hormone secretion. There was no effect of ACTH on the sex steroid dehydroepiandrosterone (DHEA). These results suggest that elephant seals are able to mount adrenal responses to multiple ACTH challenges. However, repeated stress results in facilitation of aldosterone secretion and suppression of tT3, which may impact osmoregulation and metabolism. We propose that aldosterone and tT3 are informative additional indicators of repeated stress in marine mammals.

Ocean is home to a large population of marine mammals such as dolphins and whales and concerns over anthropogenic activities in the regions close to their habitants have been increased. Therefore the ability to detect the presence of these species in the field, to analyze and classify their vocalization patterns for signs of distress and distortion of their communication calls will prove to be invaluable in protecting these species. The objective of this research is to investigate methods that automatically detect and classify vocalization patterns of marine mammals. The first work performed is the classification of bottlenose dolphin calls by type. The extraction of salient and distinguishing features from recordings is a major part of this endeavor. To this end, two strategies are evaluated with real datasets provided by Woods Hole Oceanographic Institution: The first strategy is to use contour-based features such as Time-Frequency Parameters and Fourier Descriptors and the second is to employ texture-based features such as Local Binary Patterns (LBP) and Gabor Wavelets. Once dolphin whistle features are extracted for spectrograms, selection of classification procedures is crucial to the success of the process. For this purpose, the performances of classifiers such as K-Nearest Neighbor, Support Vector Machine, and Sparse Representation Classifier (SRC) are assessed thoroughly, together with those of the underlined feature extractors. / Includes bibliography. / Dissertation (Ph.D.)--Florida Atlantic University, 2014. / FAU Electronic Theses and Dissertations Collection

<p>Sound is a key sensory modality for Hawaiian spinner dolphins. Like many other marine animals, these dolphins rely on sound and their acoustic environment for many aspects of their daily lives, making it is essential to understand soundscape in areas that are critical to their survival. Hawaiian spinner dolphins rest during the day in shallow coastal areas and forage offshore at night. In my dissertation I focus on the soundscape of the bays where Hawaiian spinner dolphins rest taking a soundscape ecology approach. I primarily relied on passive acoustic monitoring using four DSG-Ocean acoustic loggers in four Hawaiian spinner dolphin resting bays on the Kona Coast of Hawai‛i Island. 30-second recordings were made every four minutes in each of the bays for 20 to 27 months between January 8, 2011 and March 30, 2013. I also utilized concomitant vessel-based visual surveys in the four bays to provide context for these recordings. In my first chapter I used the contributions of the dolphins to the soundscape to monitor presence in the bays and found the degree of presence varied greatly from less than 40% to nearly 90% of days monitored with dolphins present. Having established these bays as important to the animals, in my second chapter I explored the many components of their resting bay soundscape and evaluated the influence of natural and human events on the soundscape. I characterized the overall soundscape in each of the four bays, used the tsunami event of March 2011 to approximate a natural soundscape and identified all loud daytime outliers. Overall, sound levels were consistently louder at night and quieter during the daytime due to the sounds from snapping shrimp. In fact, peak Hawaiian spinner dolphin resting time co-occurs with the quietest part of the day. However, I also found that humans drastically alter this daytime soundscape with sound from offshore aquaculture, vessel sound and military mid-frequency active sonar. During one recorded mid-frequency active sonar event in August 2011, sound pressure levels in the 3.15 kHz 1/3rd-octave band were as high as 45.8 dB above median ambient noise levels. Human activity both inside (vessels) and outside (sonar and aquaculture) the bays significantly altered the resting bay soundscape. Inside the bays there are high levels of human activity including vessel-based tourism directly targeting the dolphins. The interactions between humans and dolphins in their resting bays are of concern; therefore, my third chapter aimed to assess the acoustic response of the dolphins to human activity. Using days where acoustic recordings overlapped with visual surveys I found the greatest response in a bay with dolphin-centric activities, not in the bay with the most vessel activity, indicating that it is not the magnitude that elicits a response but the focus of the activity. In my fourth chapter I summarize the key results from my first three chapters to illustrate the power of multiple site design to prioritize action to protect Hawaiian spinner dolphins in their resting bays, a chapter I hope will be useful for managers should they take further action to protect the dolphins.</p> / Dissertation

<p>Effective conservation and management of top predators requires a comprehensive understanding of their distributions and of the underlying biological and physical processes that affect these distributions. The Mid-Atlantic Bight shelf break system is a dynamic and productive region where at least 32 species of cetaceans have been recorded through various systematic and opportunistic marine mammal surveys from the 1970s through 2012. My dissertation characterizes the spatial distribution and habitat of cetaceans in the Mid-Atlantic Bight shelf break system by utilizing marine mammal line-transect survey data, synoptic multi-frequency active acoustic data, and fine-scale hydrographic data collected during the 2011 summer Atlantic Marine Assessment Program for Protected Species (AMAPPS) survey. Although studies describing cetacean habitat and distributions have been previously conducted in the Mid-Atlantic Bight, my research specifically focuses on the shelf break region to elucidate both the physical and biological processes that influence cetacean distribution patterns within this cetacean hotspot. </p><p>In Chapter One I review biologically important areas for cetaceans in the Atlantic waters of the United States. I describe the study area, the shelf break region of the Mid-Atlantic Bight, in terms of the general oceanography, productivity and biodiversity. According to recent habitat-based cetacean density models, the shelf break region is an area of high cetacean abundance and density, yet little research is directed at understanding the mechanisms that establish this region as a cetacean hotspot. </p><p>In Chapter Two I present the basic physical principles of sound in water and describe the methodology used to categorize opportunistically collected multi-frequency active acoustic data using frequency responses techniques. Frequency response classification methods are usually employed in conjunction with net-tow data, but the logistics of the 2011 AMAPPS survey did not allow for appropriate net-tow data to be collected. Biologically meaningful information can be extracted from acoustic scattering regions by comparing the frequency response curves of acoustic regions to theoretical curves of known scattering models. Using the five frequencies on the EK60 system (18, 38, 70, 120, and 200 kHz), three categories of scatterers were defined: fish-like (with swim bladder), nekton-like (e.g., euphausiids), and plankton-like (e.g., copepods). I also employed a multi-frequency acoustic categorization method using three frequencies (18, 38, and 120 kHz) that has been used in the Gulf of Maine and Georges Bank which is based the presence or absence of volume backscatter above a threshold. This method is more objective than the comparison of frequency response curves because it uses an established backscatter value for the threshold. By removing all data below the threshold, only strong scattering information is retained.</p><p>In Chapter Three I analyze the distribution of the categorized acoustic regions of interest during the daytime cross shelf transects. Over all transects, plankton-like acoustic regions of interest were detected most frequently, followed by fish-like acoustic regions and then nekton-like acoustic regions. Plankton-like detections were the only significantly different acoustic detections per kilometer, although nekton-like detections were only slightly not significant. Using the threshold categorization method by Jech and Michaels (2006) provides a more conservative and discrete detection of acoustic scatterers and allows me to retrieve backscatter values along transects in areas that have been categorized. This provides continuous data values that can be integrated at discrete spatial increments for wavelet analysis. Wavelet analysis indicates significant spatial scales of interest for fish-like and nekton-like acoustic backscatter range from one to four kilometers and vary among transects. </p><p>In Chapter Four I analyze the fine scale distribution of cetaceans in the shelf break system of the Mid-Atlantic Bight using corrected sightings per trackline region, classification trees, multidimensional scaling, and random forest analysis. I describe habitat for common dolphins, Risso’s dolphins and sperm whales. From the distribution of cetacean sightings, patterns of habitat start to emerge: within the shelf break region of the Mid-Atlantic Bight, common dolphins were sighted more prevalently over the shelf while sperm whales were more frequently found in the deep waters offshore and Risso’s dolphins were most prevalent at the shelf break. Multidimensional scaling presents clear environmental separation among common dolphins and Risso’s dolphins and sperm whales. The sperm whale random forest habitat model had the lowest misclassification error (0.30) and the Risso’s dolphin random forest habitat model had the greatest misclassification error (0.37). Shallow water depth (less than 148 meters) was the primary variable selected in the classification model for common dolphin habitat. Distance to surface density fronts and surface temperature fronts were the primary variables selected in the classification models to describe Risso’s dolphin habitat and sperm whale habitat respectively. When mapped back into geographic space, these three cetacean species occupy different fine-scale habitats within the dynamic Mid-Atlantic Bight shelf break system. </p><p>In Chapter Five I present a summary of the previous chapters and present potential analytical steps to address ecological questions pertaining the dynamic shelf break region. Taken together, the results of my dissertation demonstrate the use of opportunistically collected data in ecosystem studies; emphasize the need to incorporate middle trophic level data and oceanographic features into cetacean habitat models; and emphasize the importance of developing more mechanistic understanding of dynamic ecosystems.</p> / Dissertation

Wavelets have been used in numerous geophysical studies but few have examined their applicability to underwater acoustic signals. Wavelet transforms can remove noise from a given time series and allow data analysis at multiple levels of resolution. This unique ability is exercised as a feasible application to the signals in this thesis: a reflected scattered signal from a swimbladder-bearing fish, alewife (Alosa pseudoharengus), and several Odontocetes vocalizations. Both studies reveal that wavelet-based techniques show potential in providing viable information for these acoustic signals despite the lack of statistical analysis. The alewife portion shows a reasonable first order approximation to the absolute target strength and to the time delay correlation caused by the spatial separation of scattering features in the fish. The marine mammal application shows a possible real time method to estimate the mammal's range using the root mean square (RMS) energy of the decomposed signal. Because of wavelet function mismatch, both studies conclude that more extensive research is necessary to develop these techniques into systematic processes.

Human impacts on ocean ecosystems are driving declines in marine biodiversity, including marine mammals. Comprehensive spatial data are vital for making informed management decisions that may aid species recovery and facilitate the sustainable use of ocean ecosystems. However, marine mammal studies are often data limited, thereby restricting possible research questions. Developing novel analytical approaches and incorporating unconventional datasets can expand the scope of analysis by increasing the information content of existing data sources. The goal of our research is to support conservation and management of marine mammals in British Columbia (BC), Canada, through the application of advanced spatial statistical methodology to characterize spatial distribution and density patterns and provide assessments of data uncertainty. Our first objective is to generate statistical models to map spatially continuous predictions of marine mammal distributions and densities within BC’s north coast and apply methodology from spatial statistics to identify hotspots of elevated use. We use species observations collected from systematic line transect surveys previously adjusted to generate estimates of density per nautical mile of transect. We predict the distribution and density patterns of nine marine mammal species by employing a species-habitat model to relate species densities to environmental covariates using a generalized additive model. We use spatial statistical hotspot analysis (Getis-Ord Gi*statistic) and an aspatial threshold approach to identify hotspots of high density. Our analysis reveals that hotspots selected using a top percentage threshold produced smaller and more conservative hotspots than those generated using the Gi*statistic. The Gi*statistic demonstrates a robust and objective technique for quantifying spatial hotspots and offers an alternative method to the commonly applied aspatial threshold measure. We find that maps show agreement with prior research and hotspots align with ecologically important areas previously identified by expert opinion. Our second objective is to apply map comparison techniques to compare cetacean density maps from disparate data collection methods (systematic surveys and citizen science) to evaluate the information content of each map product and quantify similarities and differences. Discrepancies are quantified by performing image differencing techniques on the rank order values of each map surface. We subsequently use the Gi*statistic to isolate regions where extreme differences occur. To assess similarities, a Gi*statistic is applied to both maps to locate spatially explicit areas of high cetacean density. Where clusters of high density values in both maps overlap we infer higher confidence that the datasets are representing a true ecological signal, while areas of difference we recommend as targeted locations for future sampling effort. We contextualize map similarities and differences using a dataset of human activity in the form of cumulative human effect scores. Overall, our analytical approach integrates novel spatial datasets from systematic surveys, citizen science, and remote sensing to provide updated information on cetacean distributions in BC. Our study generates geographic data products that fill knowledge gaps and results provide baseline information valuable for future decision-making. The methodology applied in this study can be generalized across species and locations to support spatial planning and conservation prioritization in both marine and terrestrial contexts. / Graduate / 2017-11-13