Soviet-American Cooperative Research on Marine Mammals. Volume 1- Pinnipeds: Under Project V.6. Marine Mammals, of the US-USSR Agreement on Cooperation in the Field of Environmental Protection

Some 25 to 30 yr ago, when we as students were beginning our respective careers and were developing for the first time our awareness of marine mammals in the waters separating western North America from eastern Asia, we had visions of eventually bridging the communication gap which existed between our two countries at that time. Each of us was anxious to obtain information on the distribution, biology, and ecological relations of "our" seals and walruses on "the other side," beyond our respective political boundari~s where we were not permitted to go to study them. We were concerned that the resource management practices on the other side of the Bering and Chukchi Seas, implemented in isolation, on a purely unilateral basis, might endanger the species which we had come to know and were striving to conserve. At once apparent to both of us was the need for free exchange of biological information between our two countries and, ultimately, joint management of our shared resources. In a small way, we and others made some initial efforts to generate that exchange by personal correspondence and through vocal interchange at the annual meetings of the North Pacific Fur Seal Commission. By the enabling Agreement on Cooperation in the Field of Environmental Protection, reached between our two countries in 1972, our earlier visions at last came true. Since that time, within the framework of the Marine Mammal Project under Area V of that Agreement, we and our colleagues have forged a strong bond of professional accord and respect, in an atmosphere of free intercommunication and mutual understanding. The strength and utility of this arrangement from the beginning of our joint research are reflected in the reports contained in this, the first compendium of our work. The need for a series of such a compendia became apparent to us in 1976, and its implementation was agreed on by the regular meeting of the Project in La Jolla, Calif., in January 1977. Obviously, the preparation and publication of this first volume has been excessively delayed, in part by continuing political distrust between our governments but mainly by increasing demands placed on the time of the contributors. In this period of growing environmental concern in both countries, we and our colleagues have been totally immersed in other tasks and have experienced great difficulty in drawing together the works presented here. Much of the support for doing so was provided by the State of Alaska, through funding for Organized Research at the University of Alaska-Fairbanks. For its ultimate completion in publishable form we wish to thank Helen Stockholm, Director of Publications, Institute of Marine Science, University of Alaska, and her staff, especially Ruth Hand, and the numerous referees narned herein who gave willingly oftheir time to review each ofthe manuscripts critically and to provide a high measure of professionalism to the final product. (PDF file contains 110 pages.)

Trends and Potential Interactions Between Pinnipeds and Fisheries of New England and the U.S. West Coast

Long-term trends in the abundance and distribution of several pinniped species and commercially important fisheries of New England and the contiguous U.S. west coast are reviewed, and their actual and potential interactions discussed. Emphasis is on biological interactions or competition. The pinnipeds include the western North Atlantic stock of harbor seals, Phoca vitulina concolor; western North Atlantic gray seals, Halochoerus grypus; the U.S. stock of California sea lions, Zalophus californianus californianus; the eastern stock of Steller sea lions, Eumetopias jubatus; and Pacific harbor seals, Phoca vitulina richardii. Fisheries included are those for Atlantic cod, Gadus morhua; silver hake, Merluccius bilinearis; Atlantic herring, Clupea harengus; the coastal stock of Pacific whiting, Merluccius productus; market squid, Loligo opalescens; northern anchovy, Engraulis mordax; Pacific her-ring, Clupea pallasi; and Pacific sardine, Sardinops sagax. Most of these pinniped populations have grown exponentially since passage of the U.S. Marine Mammal Protection Act in 1972. They exploit a broad prey assemblage that includes several commercially valuable species. Direct competition with fisheries is therefore possible, as is competition for the prey of commercially valuable fish. The expanding pinniped populations, fluctuations in commercial fish biomass, and level of exploitation by the fisheries may affect this potential for competition. Concerns over pinnipeds impacting fisheries (especially those with localized spawning stocks or at low biomass levels) are more prevalent than concerns over fisheries’ impacts on pinnipeds. This review provides a framework to further evaluate potential biological interactions between these pinniped populations and the commercial fisheries with which they occur.

A method to improve size estimates of walleye pollock (Theragra chalcogramma) Atka mackerel (Pleurogrammus monopterygius) consumed by pinnipeds: digestion correction factors applied to bones and otoliths recovered in scats

The lengths of otoliths and other skeletal structures recovered from the scats of pinnipeds, such as Steller sea lions (Eumetopias jubatus), correlate with body size and can be used to estimate the length of prey consumed. Unfortunately, otoliths are often found in too few scats or are too digested to usefully estimate prey size. Alternative diagnostic bones are frequently recovered, but few bone-size to prey-size correlations exist and bones are also reduced in size by various degrees owing to digestion. To prevent underestimates in prey sizes consumed techniques are required to account for the degree of digestion of alternative bones prior to estimating prey size. We developed a method (using defined criteria and photo-reference material) to assign the degree of digestion for key cranial structures of two prey species: walleye pollock (Theragra chalcogramma) and Atka mackerel (Pleurogrammus monopterygius). The method grades each structure into one of three condition categories; good, fair or poor. We also conducted feeding trials with captive Steller sea lions, feeding both fish species to determine the extent of erosion of each structure and to derive condition-specific digestion correction factors to reconstruct the original sizes of the structures consumed. In general, larger structures were relatively more digested than smaller ones. Mean size reduction varied between different types of structures (3.3−26.3%), but was not influenced by the size of the prey consumed. Results from the observations and experiments were combined to be able to reconstruct the size of prey consumed by sea lions and other pinnipeds. The proposed method has four steps: 1) measure the recovered structures and grade the extent of digestion by using defined criteria and photo-reference collection; 2) exclude structures graded in poor condition; 3) multiply measurements of structures in good and fair condition by their appropriate digestion correction factors to derive their original size; and 4) calculate the size of prey from allometric regressions relating corrected structure measurements to body lengths. This technique can be readily applied to piscivore dietary studies that use hard remains of fish.

Historic and recent occurrences of pinnipeds in the Archipelago of the Azores

The Archipelago of the Azores (Portugal) is located between 378 and 418N and 258 and 318W and crosses the Mid-Atlantic Ridge. It is the most isolated archipelago in the Atlantic, situated 1600 km west of mainland Portugal and 3500 km from the eastern coast of the United States of America. At present, the only population of seals occurring in the Portuguese territory is found on Desertas Islands, Archipelago of Madeira, where a colony of 24 Mediterranean monk seals, Monachus monachus (Hermann, 1779), still persists (Pires and Neves 2001). Nonetheless, historical accounts reported by Frutuoso (1983) dating from the early to late 1500s mention sightings of ‘‘sea wolves’’ (the old Portuguese folk term for the Mediterranean monk seal) at several sites along the Azorean Island of Santa Maria. Little is known about the occurrence of monk seals in this area over the past five centuries, but the species certainly did not escape deliberate killing by the first settlers. While the early monk seal reports by Frutuoso (1983) are the only reports referring to the presence of colonies of seals in the Azores, more recently several sightings and strandings of vagrant seals of other species have been noted.

Variance in male reproductive success and sexual size dimorphism in pinnipeds: testing an assumption of sexual selection theory

The theory of evolution by sexual selection for sexual size dimorphism (SSD) postulates that SSD primarily reflects the adaptation of males and females to their different reproductive roles. For example, competition among males for access to females increases male body size because larger males are better able to maintain dominant status than smaller males. Larger dominant males sire most offspring while smaller subordinate males are unsuccessful, leading to skew in reproductive success. Therefore, species with male-biased SSD are predicted to have greater variance in male reproductive success than those in which both sexes are similar in size. We tested this prediction among the Pinnipedia, a mammalian group with a great variation in SSD. From a literature review, we identified genetic estimates of male reproductive success for 10 pinniped taxa (eight unique species and two subspecies of a ninth species) that range from seals with similarly sized males and females to species in which males are more than four times as large as females. We found no support for a positive relationship between variance in reproductive success and SSD among pinnipeds after excluding the elephant seals Mirounga leonina and Mirounga angustirostris, which we discuss as distinctive cases. Several explanations for these results are presented, including the revival of one of Darwin's original ideas. Darwin proposed that natural selection may explain SSD based on differences in energetic requirements between sexes and the potential for sexual niche segregation. Males may develop larger bodies to exploit resources that remain unavailable to females due to the energetic constraints imposed on female mammals by gestation and lactation. The importance of this alternative explanation remains to be tested.

Accuracy of ARGOS locations of pinnipeds at-sea estimated using Fastloc GPS

Background: ARGOS satellite telemetry is one of the most widely used methods to track the movements of free-ranging marine and terrestrial animals and is fundamental to studies of foraging ecology, migratory behavior and habitat-use. ARGOS location estimates do not include complete error estimations, and for many marine organisms, the most commonly acquired locations (Location Class 0, A, B, or Z) are provided with no declared error estimate.
Methodology/Principal Findings: We compared the accuracy of ARGOS locations to those obtained using Fastloc GPS from the same electronic tags on five species of pinnipeds: 9 California sea lions (Zalophus californianus), 4 Galapagos sea lions (Zalophus wollebaeki), 6 Cape fur seals (Arctocephalus pusillus pusillus), 3 Australian fur seals (A. p. doriferus) and 5 northern elephant seals (Mirounga angustirostris). These species encompass a range of marine habitats (highly pelagic vs coastal), diving behaviors (mean dive durations 2–21 min) and range of latitudes (equator to temperate). A total of 7,318 ARGOS positions and 27,046 GPS positions were collected. Of these, 1,105 ARGOS positions were obtained within five minutes of a GPS position and were used for comparison. The 68^th percentile ARGOS location errors as measured in this study were LC-3
0.49 km, LC-2 1.01 km, LC-1 1.20 km, LC-0 4.18 km, LC-A 6.19 km, LC-B 10.28 km.
Conclusions/Significance: The ARGOS errors measured here are greater than those provided by ARGOS, but within the range of other studies. The error was non-normally distributed with each LC highly right-skewed. Locations of species that make short duration dives and spend extended periods on the surface (sea lions and fur seals) had less error than species like elephant seals that spend more time underwater and have shorter surface intervals. Supplemental data (S1) are provided allowing the creation of density distributions that can be used in a variety of filtering algorithms to improve the quality of ARGOS tracking data.

Navigating uncertain waters: a critical review of inferring foraging behaviour from location and dive data in pinnipeds

In the last thirty years, the emergence and progression of biologging technology has led to great advances in marine predator ecology. Large databases of location and dive observations from biologging devices have been compiled for an increasing number of diving predator species (such as pinnipeds, sea turtles, seabirds and cetaceans), enabling complex questions about animal activity budgets and habitat use to be addressed. Central to answering these questions is our ability to correctly identify and quantify the frequency of essential behaviours, such as foraging. Despite technological advances that have increased the quality and resolution of location and dive data, accurately interpreting behaviour from such data remains a challenge, and analytical methods are only beginning to unlock the full potential of existing datasets. This review evaluates both traditional and emerging methods and presents a starting platform of options for future studies of marine predator foraging ecology, particularly from location and two-dimensional (time-depth) dive data. We outline the different devices and data types available, discuss the limitations and advantages of commonly-used analytical techniques, and highlight key areas for future research. We focus our review on pinnipeds - one of the most studied taxa of marine predators - but offer insights that will be applicable to other air-breathing marine predator tracking studies. We highlight that traditionally-used methods for inferring foraging from location and dive data, such as first-passage time and dive shape analysis, have important caveats and limitations depending on the nature of the data and the research question. We suggest that more holistic statistical techniques, such as state-space models, which can synthesise multiple track, dive and environmental metrics whilst simultaneously accounting for measurement error, offer more robust alternatives. Finally, we identify a need for more research to elucidate the role of physical oceanography, device effects, study animal selection, and developmental stages in predator behaviour and data interpretation.

Micronekton of the North Pacific

1. INTRODUCTION 1.1 Working Group History 2. SPECIES COMPOSITION AND DISTRIBUTION PATTERNS RELATED TO WATER MASSES 2.1 Mesopelagic Fishes 2.1.1 Dominant families 2.1.2 Large-scale feeding and/or spawning migration or expatriation? 2.1.3 Definition of water masses 2.1.4 Species composition 2.2 Crustacean Micronekton 2.2.1 Euphausiids 2.2.2 Mysids and decapods 2.3 Cephalopod Micronekton 2.3.1 Family Enoploteuthidae 2.3.2 Family Gonatidae 2.3.3 Family Onychoteuthidae 2.3.4 Family Pyroteuthidae 2.3.5 Other cephalopods 3. VERTICAL DISTRIBUTION PATTERNS 3.1 Mesopelagic Fishes 3.1.1 Significance of diel vertical migration 3.1.2 DVM patterns 3.1.3 Ontogenetic change in DVM patterns 3.2 Crustacean Micronekton 3.3 Cephalopod Micronekton 4. BIOMASS PATTERNS 4.1 Micronektonic Fish 5. LIFE HISTORY 5.1 Fish Micronekton 5.1.1 Age and growth 5.1.2 Production 5.1.3 Reproduction 5.1.4 Mortality 5.2 Crustacean Micronekton 5.2.1 Age and growth 5.2.2 Production 5.2.3 Reproduction and early life history 5.2.4 Mortality 5.3 Cephalopod Micronekton 5.3.1 Age and growth 5.3.2 Production 5.3.3 Reproduction and early life history 5.3.4 Mortality 6. ECOLOGICAL RELATIONS 6.1 Feeding Habits 6.1.1 Fish micronekton 6.1.2 Crustacean micronekton 6.1.3 Cephalopod micronekton 6.2 Estimating the Impact of Micronekton Predation on Zooplankton 6.2.1 Predation by micronektonic fish 6.3 Predators 6.3.1 Cephalopods 6.3.2 Elasmobranchs 6.3.3 Osteichthyes 6.3.4 Seabirds 6.3.5 Pinnipeds 6.3.6 Cetaceans 6.3.7 Human consumption 6.4 Predation Rate 6.5 Ecosystem Perspectives 6.6 Interactions between Micronekton and Shallow Topographies 7. SAMPLING CONSIDERATIONS 7.1 Net Trawling 7.1.1 Sampling gears 7.1.2 Sampling of surface migratory myctophids 7.1.3 Commercial-sized trawl sampling 7.1.4 Sampling of euphausiids and pelagic decapods 7.2 Acoustic Sampling 7.2.1 Acoustic theory and usage 7.3 Video Observations (Submersible and ROV) 8. SUMMARY OF PRESENT STATE OF KNOWLEDGE 8.1 Fish Micronekton 8.2 Crustacean Micronekton 8.3 Cephalopod Micronekton 9. RECOMMENDATIONS 10. REFERENCES 11. APPENDICES (122 page document)

Food habits of California Sea Lions (Zalophus californianus) and their impact on Salmonid Fisheries in Monterey Bay, California

In the ocean commercial troll and recreational salmon fishery in Monterey Bay California, California sea lions (Zalophus califomianus) will swim near or follow fishing boats and will depredate fish once hooked. The objectives of the study were to determine the percentage of salmon taken by pinnipeds in commercial and recreational fisheries, identify relative importance of prey items seasonally consumed by sea lions, and determine the proportion of salmonids in the sea lion diet on a seasonal basis. From April 1997 through September 1998, 1041 hours of onboard and dockside surveys of the commercial and recreational salmon fisheries were conducted at the three ports in Monterey Bay, California. Sea lions depreadated 7.9 % of the fish hooked in the commercial fishery in 1997 and 28.6 % in 1998,8.4 % (1997) and 18.3 % (1998) of the CPFV fishery, and 15.6 % (1997) and 17.5 % (1998) of the private skiff fishery. Increased depredation rates in both the commercial and recreational salmon fisheries in 1998 were most likely the result of the large EI Nino Southern Oscillation event that occurred in 1997-1998 during which a greater number of sea lions were present in central California. Prey hardparts identified in sea lion fecal samples collected in Monterey Bay indicated that schooling fishes were the predominant prey fish species, such as market squid (Loligo opalescens), Pacific sardine (Sardinops caeruleus), northern anchovy (Engraulis mordax), and rockfish (Sebastes sp.). Sea lions consumed similar prey species in the summer and fall 1997, winter 1997-98, and spring 1998 (PSI> 70.0) with market squid and northern anchovy being the dominant prey species. However, prey composition changed significantly during the summer 1998 and fall 1998 (PSI < 46.0) because of the increased importance of sardine and rockfish in the diet and the decreased importance of market squid. This report does not intend to imply that salmonids are not a prey species for pinnipeds in the Monterey Bay region, but highlights the difficulties encountered in establishing the role of salmonids in the pinniped diet when analyzing fecal samples. (PDF contains 38 pages).

A method for age determination of dolphins.

The use of growth layers in teeth as an indicator of age in odnotocetes and pinnipeds was suggested by Laws (1954) and since then the method has been used extensively in both marine and non-marine mammals. Dentinal growth layers are groups (growth layer groups) of repetitive alternating bands which in cross-section are similar to growth rings in trees. The most commonly used methods for counting growth layer groups (GLGs) are by undecalcified longitudinal thin sections (150 um) or decalcified and stained thin sections (10-30 um). In longitudinal sections viewed with light microscopy, GLGs appear as opaque and translucent cones nestled one inside another, with the oldest dentine Iying adjacent to the enamel, and the newest layer borderinq the pulp cavity.

Spatial and seasonal relationships between Pacific harbor seals (Phoca vitulina richardii) and their prey, at multiple scales

Knowing where pinnipeds forage is vital to managing and protecting their populations, and for assessing potential interactions with fisheries. We assessed the spatial relationship between the seasonal distribution of Pacific harbor seals (Phoca vitulina richardii) outfitted with satellite transmitters and the seasonal distributions of potential harbor seal prey species in San Francisco Bay, California. Pearson’s correlation coefficients were calculated between the number of harbor seal locations in an area of the San Francisco Bay and the abundance of specific prey species in the same area. The influence of scale on the analyses was assessed by varying the scale of analysis from 1 to 10 km. There was consistency in the prey species targeted by harbor seals year-round, although there were seasonal differences between the most important prey species. The highest correlations between harbor seals and their prey were found for seasonally abundant benthic species, located within about 10 km of the primary haul-out site. Probable foraging habitat for harbor seals was identified, based on areas with high abundances of prey species that were strongly correlated with harbor seal distribution. With comparable local data inputs, this approach has potential application to pinniped management in other areas, and to decisions about the location of marine reserves designed to protect these species.

Application of two methods for determining diet of northern fur seals (Callorhinus ursinus)

Examination of hard parts recovered from scats (feces) is currently the most common method for determining the diet of pinnipeds. However, large or sharp prey remains may be spewed (regurgitated) biasing prey composition and size estimations in diet studies based on scats. Percent frequency of occurrence (FO%) and age or size of selected prey remains recovered from northern fur seal (Callorhinus ursinus) scat (n=3444) and spew samples (n=267) collected from rookeries on St. George Island and St. Paul Island, Alaska, between 1990 and 2000 were compared to determine if a bias in prey composition and age or size estimations existed between scats and spews. Overall prey composition was similar between sample type and location, but the relative FO% of primary prey (≥5%) varied by sample type and location. Age or size estimates of walleye pollock (Theragra chalcogramma) and of two species of gonatid squids (Gonatopsis borealis and Berryteuthis magister) were significantly larger in spews than in scats. Observed differences in FO% and estimated age or size of prey species whose remains were found in scats and spews likely result from size-selective digestion of prey remains. Scats were biased toward smaller prey remains, whereas spews were biased toward larger prey remains and cephalopod beaks. The percent overlap between age classes of walleye pollock caught by the commercial trawl fishery and age classes of walleye pollock consumed by northern fur seals varied noticeably between sample types for both islands (scats: St. George=15. 5%; St. Paul=4.1%; spews: St. George=94.6%; St. Paul=89.6%). These results demonstrate that the inclusion of multiple sampling methods allows for a more accurate assessment of northern fur seal prey occurrence and prey age and size.

Estimates of Marine Mammal, Sea Turtle, and Seabird Mortality in the California Drift Gillnet Fishery for Swordfish and Thresher Shark, 1996–2002

Estimates of incidental marine mammal, sea turtle, and seabird mortality in the California drift gillnet fishery for broadbill swordfish, Xiphias gladius, and common thresher shark, Alopias vulpinus, are summarized for the 7-year period, 1996 to 2002. Fishery observer coverage was 19% over the period (3,369 days observed/17,649 days fished). An experiment to test the effectiveness of acoustic pingers on reducing marine mammal entanglements in this fishery began in 1996 and resulted in statistically significant reductions in marine mammal bycatch. The most commonly entangled marine mammal species were the short-beaked common dolphin, Delphinus delphis; California sea lion, Zalophus californianus; and northern right whale dolphin, Lissodelphis borealis. Estimated mortality by species (CV and observed mortality in parentheses) from 1996 to 2002 is 861 (0.11, 133) short-beaked common dolphins; 553 (0.16, 103) California sea lions; 151 (0.25, 31) northern right whale dolphins; 150 (0.21, 27) northern elephant seals, Mirounga angustirostris; 54 (0.41, 10) long-beaked common dolphins, Delphinus capensis; 44 (0.53, 6) Dall’s porpoise, Phocoenoides dalli; 19 (0.60, 5) Risso’s dolphins, Grampus griseus; 11 (0.71, 2) gray whales, Eschrichtius robustus; 7 (0.83, 2) sperm whales, Physeter macrocephalus; 7 (0.96, 1) short-finned pilot whales, Globicephala macrorhychus; 12 (1.06, 1) minke whales, Balaenoptera acutorostrata; 5 (1.05, 1) fin whales, Balaenoptera physalus; 11 (0.68, 2) unidentified pinnipeds; 33 (0.52, 4) leatherback turtles, Dermochelys coriacea; 18 (0.57, 3) loggerhead turtles, Caretta caretta; 13 (0.73, 3) northern fulmars, Fulmarus glacialis; and 6 (0.86, 2) unidentified birds.

DNA extraction from formalin-fixed Franciscana tissues

The present paper reports the extraction of DNA from formalin-fixed Pontoporia blainvillei tissues. Following the Vachot and Monerot (1996) protocol, fragmented DNA (300-700bp) was extracted from more than 95% of liver and muscle samples. DNA yield in liver samples was significantly higher than in muscle samples (4.574 ± 1.169mg DNA/mg versus 0.808 ± 0.297mg DNA/mg). Similar results were obtained from nine other species of cetaceans and five species of pinnipeds. It is of special interest to have a method that allows the utilisation of museum specimens not originally preserved for genetic studies, which may include rarely available, declining or extinct species. SPANISH: El presente trabajo reporta la extracción de ADN a partir de tejidos formolizados de Pontoporia blainvillei. Siguiendo el protocolo de Vachot y Monerot (1996) se pudo extraer ADN degradado (300-700pb) en más del 95% de las muestras de hígado y músculo analizadas. El rendimiento en ADN fue significativamente mayor en muestras de hígado que en muestras de músculo (4.574 ± 1.169mg DNA/mg tejido húmedo versus 0.808 ± 0.297mg DNA/mg tejido húmedo). Resultados similares se obtuvieron en otras nueve especies de Cetáceos y cinco de Pinnípedos. Resulta de gran interés contar con un método que permita la utilización de especímenes depositados en museos y que no hayan sido originalmente colectados para estudios genéticos, incluyendo especies de difícil obtención, en franca declinación o extintas.