BILATERAL DIRECTIONAL ASYMMETRY OF THE APPENDICULAR SKELETON OF THE HARBOR PORPOISE (PHOCOENA PHOCOENA)

Directional asymmetry ( DA ) of the lengths, diameters, and masses of the scapula, humerus, radius, and ulna were analyzed on a sample of 213 harbor porpoises from Denmark and West Greenland. The levels of DA were consistent across yearlings and older animals, mature and immature animals, sexes, and populations. All investigated variables showed significant DA favoring the right side. For the pooled sample, DA of lengths ranged from 0.2% SE 0.1 (ulna) to 1.2% SE 0.1 (scapula). DA s of diameters were 1.5% SE 0.2 for the humerus, 0.6% SE 0.1 for the radius, and 1.3% SE 0.2 for the ulna. DA of mass ranged from 2.8% SE 0.5 (humerus) to 4.3% SE 0.7 (ulna). The humerus and ulna had significantly larger mean diameter/ length ratios on the right side than the left, making them more robust. The large DA of scapula length indicates larger muscle mass associated with the right flipper, while the generally more robust right humerii and ulnae may be designed for higher levels of mechanical stress. These DA s and the examples of lateralized behavior recorded in cetaceans, point to the existence of lateralized use of the flippers at the population level in harbor porpoises and possibly other cetacean species.     

DIVING BEHAVIOR AND PERFORMANCE OF HARBOR PORPOISES, PHOCOENA PHOCOENA, IN FUNKA BAY, HOKKAIDO, JAPAN

In order to monitor the diving behavior of free-ranging cetaceans, microdataloggers, with pre-programmed release mechanisms, were attached to the dorsal fins of two female harbor porpoises ( Phocoena phocoena ) in Funka Bay, Hokkaido, Japan, in 1994. The two loggers were successfully recovered and a total of 141 h of diving data (depth and water temperature in 4,671 dives) was obtained. Both porpoises dived almost continuously, rarely exhibiting long-term rest at the surface. Maximum dive depths were 98.6 m and 70.8 m, respectively, with more than 70% of diving time at 20 m or less. Most shallow dives were V-shaped with no bottom time. The V-shaped dives were significantly shallower in dive depth and shorter in dive duration than U-shaped dives. Descent rate was not constant during a dive. The deeper the dive depths, the faster the mean descent and initial descent rates. This suggests that porpoises have anticipated the depth to which they will dive before initiating the dive itself.     

HEMATOLOGY VALUES OF WILD HARBOR PORPOISES (PHOCOENA PHOCOENA) FROM THE BAY OF FUNDY,CANADA

Clinical hematology values were determined for 29 harbor porpoises (Phocoena phocoena) released from herring weirs in the Bay of Fundy, Canada. Erythrocyte values exhibited narrow ranges, but there was a high degree of individual variability in counts of white blood cells. Total white cell counts ranged from 2.6 to 15.5 ± 10⁹/liter, with an overall mean of 6.5 ± 2.7 ± 10±/liter. There were significant differences among reproductive classes in mean values of total red blood cells, hematocrit (HCT), mean cell volume (MCV), mean cell hemoglobin (MCHC) and total monocyte count. Wild harbor porpoises had fewer white blood cells, lower relative total number of neutrophils and lymphocytes, and higher percentages of monocytes and eosinophils than reported in the literature for captive porpoises. Compared to published values for other odontocetes, the hemograms of harbor porpoises were most similar to those of Pacific white-sided dolphins (Lagenorhynchus acutus). These hematology data represent a baseline from free-ranging harbor porpoises that can be used as a reference for long-term monitoring of the health of this population and as a tool for rehabilitation facilities.     

VARIABILITY IN THE DIET OF HARBOR PORPOISES (PHOCOENA PHOCOENA) IN SCOTTISH WATERS 1992–2003

Interannual, seasonal, and regional variation in the diet of porpoises, Phocoena phocoena , in Scottish (UK) waters was studied using stomach contents of animals stranded between 1992 and 2003. Most samples came from the east coast (including many porpoises killed by bottlenose dolphins), with smaller numbers from the west coast and from Shetland. The most important prey types, in terms of contribution by number and mass, were whiting ( Merlangius merlangus ) and sand eels (Ammodytidae). Multivariate analysis confirmed the existence of regional, seasonal, and interannual variation in diet, as well as differences ( i. e. , biases) related to cause of death. These differences were further explored using univariate analyses. Sand eels were more important in the summer months (quarters 2 and 3) and on the east coast, whereas gadids were more important in winter and in the Shetland area. Some, but not all, observed trends in the numerical importance and size of prey taken were consistent with trends in abundance and size of fish taken during research trawl surveys. There was some evidence that porpoises <1 yr old took more gobies (Gobiidae) and shrimps than did older porpoises. Clupeids (herring Clupea harengus and sprat Sprattus sprattus ) formed a relatively small proportion of the diet, but their importance varied from year to year. Although possible methodological biases prevent firm conclusions, it appears that the importance of clupeids in porpoise diet may have decreased since the 1960s, mirroring the decline in North Sea herring abundance. The recovery of the North Sea herring stock in recent years is not as yet reflected in porpoise diet.     

EFFECT OF THE SOUND GENERATED BY AN ACOUSTIC HARASSMENT DEVICE ON THE RELATIVE ABUNDANCE AND DISTRIBUTION OF HARBOR PORPOISES (PHOCOENA PHOCOENA) IN RETREAT PASSAGE, BRITISH COLUMBIA

We describe an experiment conducted to assess the impact of the sound generated by an acoustic harassment device (AHD) on the relative abundance and distribution of harbor porpoises ( Phocoena phocoena ) in Retreat Passage, British Columbia. During control periods when the AHD was inactive, the mean number of porpoises observed in the study area was 0.39 for broad area scans conducted with the naked eye and 0.48 for narrow sector scans conducted with binoculars. Abundance declined precipitously when the AHD was activated, to 0.007 porpoises per broad area scan and 0.018 per narrow sector scan. The mean number of porpoise resightings while tracking their movements also declined from 12.2 to 13.6 per sighting during control periods to 1.1–1.9 per sighting when the AHD was activated, which suggested that the few porpoises that ventured into the study area spent less time within it when the AHD was activated. The effect of the AHD diminished with distance. No porpoises were observed within 200 m of the AHD when it was activated. The number of sightings and resightings observed when it was activated was less than 0.2% of the number expected had there been no AHD effect at a range of 200–399 m, 1.4% the number expected at a range of 400–599 m, varied between 2.5% and 3.3% of the number expected at a range of 600–2,499 m, and was 8.1% the number expected at a range of 2,500–3,500 m, which suggested that the impact of the AHD extended beyond our maximum sighting range of 3.5 km.     

ECHOLOCATION RATES OF TWO HARBOR PORPOISES (PHOCOENA PHOCOENA)

The rate of occurrence of click trains of two harbor porpoises ( Phocoena phocoena ) were counted during 14 nights. We developed an echolocation signal detection system that was harnessed to a porpoise and activated a light when the animal emitted an echolocation signal. This device, referred to as a click-light, detects echolocation signals above 150 dB re 1 μPa in the 28–180 kHz range. Echolocation rates, i.e., occurrences of click trains, changed frequently, ranging from 0 to 25 per minute. Echolocation rates were affected by feeding, individual difference, and enclosure type such as the net enclosure and the pool. The porpoise echolocation rates seemed to show acclimation.     

GEOGRAPHICAL VARIATION IN METRIC SKULL CHARACTERS AMONG PROPOSED SUBPOPULATIONS AND STOCKS OF HARBOR PORPOISE, PHOCOENA PHOCOENA, IN THE WESTERN NORTH ATLANTIC

A total of 433 skulls of harbor porpoises, Phocoena phocoena , were examined to evaluate geographical variation within the western North Atlantic, using analysis of covariance and discriminant analysis based on 45 measurements of skull components. The relationship between condylobasal length and body length was sigmoid in both sexes. Condylobasal length is generally less in adult males than in adult females.
Despite some overlap, morphological differences between different regions and even adjacent local stocks could be recognized by ANCOVA and discriminant analysis. The percentage of correct classification by discriminant functions was 65.2% for the four regional samples: (1) North Canadian Atlantic (which includes Gulf of St. Lawrence and Newfoundland), (2) Bay of Fundy, (3) Gulf of Maine-New York, and (4) Maryland-South Carolina. It was 65.9% for the four local geographical areas (10s of km) within the Bay of Fundy; Quoddy-region porpoises revealed a closer relationship with the southwestern Nova Scotia population than with other adjacent stocks. The results at this level imply that gene flow is restricted to some degree even among closely adjacent geographical units. Management of this species should aim, therefore, for conservation of a variety of local stocks.     

REACTIONS OF CAPTIVE HARBOR PORPOISES (PHOCOENA PHOCOENA) TO PINGER-LIKE SOUNDS

Pingers on gill nets can reduce bycatch of harbor porpoises. If harbor porpoises habituate to pingers, the effect may be reduced or lost. Two captive harbor porpoises were exposed to three sound types. All sounds were in the frequency band from 100 kHz to 140 kHz, 200 ms long, and presented once per 4 s. The source level was 153 dB re 1 μPa RMS at 1 m. Each session consisted of a 10‐min presound, a 5‐min sound, and a 10‐min postsound period. Behavior was recorded on video and on dataloggers placed on the dorsal fin of one animal. The loggers recorded heart rate, swimming speed, dive duration, and depth. The animals responded most strongly to the initial presentations of a sound. Surface time decreased, the heart rate dropped below the normal bradycardia, and echolocation activity decreased. The reactions of both animals diminished rapidly in the following sessions. Should the waning of responsiveness apply to wild animals, porpoises may adapt to the sounds but still avoid nets, or the bycatch may increase after some time. The success of long‐term use of pingers may then depend on the variety of sounds and rates of exposure.     

BLOOD CHEMISTRY OF WILD HARBOR PORPOISES PHOCOENA PHOCOENA (L.)

Blood chemistry values were measured from 31 harbor porpoises ( Phocoena phocoena ) released from herring weirs in the Bay of Fundy, Canada. Sodium, phosphorus, calcium, chloride, magnesium, total protein, albumin, globulin, urea, cholesterol, serum osmolality, and alanine aminotransferase levels fell within ranges reported for captive harbor porpoises and other odontocetes. Glucose, potassium, creatine kinase, aspartate aminotransferase, hemoglobin, thyroxine, bilirubin, and alkaline phosphatase levels were generally higher than those reported for captive odontocetes. Cortisol and creatinine levels were significantly higher ( P = 0.0356 and 0.0174, respectively) in porpoises that were handled for longer periods and fitted with electronic tags (mean cortisol value = 314 ± 107 nmol/L, mean creatinine value = 94 ± 14 μmol/L) than those receiving roto-tags (mean cortisol value = 222 ± 82 nmol/L, mean creatinine value = 75 ± 15 μmol/L). A more controlled study of duration of handling, time spent out of water, degree of manipulation, and method of tag attachment is required to identify which factors are responsible for the stress response.     

LIFE IN THE FAST LANE: THE LIFE HISTORY OF HARBOR PORPOISES FROM THE GULF OF MAINE

We describe the life history of harbor porpoises in the Gulf of Maine by examining 239 animals killed in gill net fisheries and comparing these findings with the results of previous studies from the Bay of Fundy. Most female porpoises matured at age three and became pregnant each year thereafter. Reproduction was strictly seasonal, with ovulation, conception, and parturition occurring in the spring and early summer. The oldest specimen in the sample was 17 yr of age, but most individuals were younger than 12. The findings are similar to those of earlier studies from the Bay of Fundy and support the hypothesis that these animals form a single population. Harbor porpoises represent one end of a continuum of odontocete life histories that spans a wide diversity of strategies. In comparison with other, large odontocetes, harbor porpoises mature at an earlier age, reproduce more frequently, and live for shorter periods.     

THE ABUNDANCE OF HARBOR SEALS IN THE GULF OF ALASKA

The abundance of harbor seals ( Phoca vitulina richardii ) has declined in recent decades at several Alaska locations. The causes of these declines are unknown, but there is concern about the status of the populations, especially in the Gulf of Alaska. To assess the status of harbor seals in the Gulf of Alaska, we conducted aerial surveys of seals on their haul-out sites in August-September 1996. Many factors influence the propensity of seals to haul out, including tides, weather, time of day, and time of year. Because these "covariates" cannot simultaneously be controlled through survey design, we used a regression model to adjust the counts to an estimate of the number of seals that would have been ashore during a hypothetical survey conducted under ideal conditions for hauling out. The regression, a generalized additive model, not only provided an adjustment for the covariates, but also confirmed the nature and shape of the covariate effects on haul-out behavior. The number of seals hauled out was greatest at the beginning of the surveys (mid-August). There was a broad daily peak from about 1100–1400 local solar time. The greatest numbers were hauled out at low tide on terrestrial sites. Tidal state made little difference in the numbers hauled out on glacial ice, where the area available to seals did not fluctuate with the tide. Adjusting the survey counts to the ideal state for each covariate produced an estimate of 30,035 seals, about 1.8 times the total of the unadjusted counts (16,355 seals). To the adjusted count, we applied a correction factor of 1.198 from a separate study of two haul-out sites elsewhere in Alaska, to produce a total abundance estimate of 35,981 (SE 1,833). This estimate accounts both for the effect of covariates on survey counts and for the proportion of seals that remained in the water even under ideal conditions for hauling out.     

TRENDS IN ABUNDANCE OF ALASKA HARBOR SEALS, 1983–2001

We estimated trends in abundance of harbor seals ( Phoca vitulina richardsii ) using over dispersed, multinomial models and counts obtained during aerial surveys conducted during 1983–2001 in the Ketchikan, Sitka, Kodiak, and Bristol Bay areas of Alaska. Harbor seal numbers increased significantly at 7.4%/yr during 1983–1998 and 5.6%/yr during 1994–1998 in the Ketchikan area, and 6.6%/yr during 1993–2001 in the Kodiak area. Counts were stable (trends not significant) during 1984–2001 (0.7%/yr) and 1995–2001 (-0.4%/yr) in Sitka, and during 1998–2001 (-1.3%/yr) in Bristol Bay. The influence of covariates ( e.g. , survey date, tide height) on trend estimates was significant and varied among areas and across years, demonstrating the need to include covariates in statistical analyses to accurately estimate trend. Our increasing trend estimate for Kodiak represents the first documented increase in harbor seal numbers over a relatively expansive area in the Gulf of Alaska. However, the trend for the Gulf of Alaska stock is equivocal due to the continued decline in Prince William Sound. Similarly, the trend for the Southeast Alaska stock is equivocal based on our increasing (Ketchikan) and stable (Sitka) trend estimates, and a recent decline reported for Glacier Bay. The Bering Sea stock appears stable after a period of possible decline.     

SKIN DENSITY AND ITS INFLUENCE ON BUOYANCY IN THE MANATEE (TRICHECHUS MANATUS LATIROSTRIS), HARBOR PORPOISE (PHOCOENA PHOCOENA), AND BOTTLENOSE DOLPHIN (TURSIOPS TRUNCATUS)

This study investigated how skin contributes to buoyancy control in the Florida manatee ( Trichechus manatus latirostris ), harbor porpoise ( Phocoena phocoena ), and bottlenose dolphin ( Tursiops truncatus ). Manatees are shallow divers and control their position in the water column hydrostatically. The two cetaceans are relatively deep divers that control their buoyancy hydrodynamically. Although the cetacean skin had been hypothesized to lower total body density ( e. g. , Dearolf et al. 2000, Nowacek et al. 2001), its buoyant force had not been calculated. The density of manatee skin, and its contribution to buoyancy, was unknown. Skin densities of 27 manatees, five harbor porpoises, and five bottlenose dolphins were measured volumetrically. Skin mass and density were used to calculate buoyant force. Harbor porpoise (952 kg/m³) and bottlenose dolphin (969 kg/m³) skins were less dense than seawater, and added 9 and 25 N of positive buoyant force, respectively, to total body buoyancy. By contrast, manatee skin (1,121 kg/m³) contributed 56 N of negative buoyant force, which equaled 70% of the negative buoyant force of their dense, pachyosteosclerotic ribs. Calculation of buoyant forces of the skeleton, skin and lungs demonstrates that the manatee is positively buoyant at the surface and negatively buoyant at depths of less than 10 m.     

DIEL VARIATION IN ECHOLOCATION BEHAVIOR OF WILD HARBOR PORPOISES

The echolocation rate and behavior of wild harbor porpoises were studied using a harbor porpoise click detector (POD) deployed close to the bottom at 40 m depth in Scottish waters, UK, April—June 2001. Echolocation variables were compared among four diel phases; morning, day, evening, and night. The echolocation encounter rate, the minimum interclick interval per train, and the proportion of echolocation click trains with a minimum interclick interval below 10 msec were all significantly higher at night than during the day. The variation in echolocation rate implies that porpoises increased their echolocation rate and/or visited the depth of the POD more often at night than during the day. Further, the changes in minimum interclick interval per train suggest that they used their echolocation for foraging or investigating objects at a close range to a higher extent, and acoustically explored the environment at greater distances at night than during the day.     

RADIOTAGGING STUDIES OF BELUKHA WHALES (DELPHINAPTERUS LEUCAS) IN BRISTOL BAY, ALASKA

Research was conducted in Bristol Bay, Alaska, to determine the applicability of radiotagging to studies of behavior, distribution and movements of belukha whales. Backpack-style VHF transmitters were attached to two belukhas by pinning through the dorsal ridge. Both packages were shed after about 2 wk due to migration of the pin through the tissue. Movements of radio-tagged whales were essentially local within Kvichak Bay. Three basic respiration patterns were identified: surfacings that were grouped into breathing periods separated by longer dives; surfacings that did not occur during restricted breathing periods; and long-to-very-long surfacings separated by short-to-very-short dives. These patterns were interpreted as representing traveling, feeding and feeding or resting in very shallow water. Surface and dive interval data were used to calculate a correction factor of 2.75, which could then be applied to aerial survey counts to estimate the total number of belukha whales in the study area. Modifications to radio packages are necessary in order to increase retention time.     

Distribution,abundance, and density of harbor porpoises in Hood Canal,Washington

Harbor porpoises (Phocoena phocoena) are the only cetaceans routinely sighted in Hood Canal, a narrow fjord that comprises the western edge of Puget Sound, Washington, USA. Harbor porpoises are sensitive to anthropogenic sounds, including noise from recreational and commercial vessel traffic, and the United States Navy, which conducts military training and testing within Hood Canal that can include underwater sound sources. This study was funded as part of the Navy monitoring program to assess potential impacts of naval activities on cetaceans. We conducted vessel-based line-transect surveys for harbor porpoises in Hood Canal in 2022–2023 to derive seasonal estimates of abundance and density. We carried out surveys over 37 days and surveyed the entire canal twice per season totaling 2,176 km of on-effort track line. We recorded 809 on-effort harbor porpoise groups and 1,385 individuals. Seasonal abundance estimates were lowest in winter (308 animals, 95% CI = 189–503) and gradually increased through spring and summer to a peak of 1,336 animals (95% CI = 826–2,160) in fall. Overall porpoise density was highest in central Hood Canal, an area that includes a designated United States Navy training range, though porpoise sightings were notably absent in a 21-km² area adjacent to the naval submarine base within this otherwise high-density region. Though we collected only a single year of data, these results suggest that harbor porpoise abundance in Hood Canal increased significantly since it was last estimated (2013–2015). The notable seasonal fluctuation of harbor porpoise abundance suggests Hood Canal may host a larger percentage of the overall Washington Inland Waters stock during the fall season, raising important management considerations.     

Morphology and microanatomy of harbor porpoise (Phocoena phocoena) dorsal fin tubercles

The unique pattern of small tubercles on the leading edge of the dorsal fins of harbor porpoises (Phocoena phocoena) has been widely noted in the literature, though their structure or function has never been conclusively identified. We examined external morphology and microanatomy of the tubercles for further understanding of the nature of the tubercles. Measurements were taken of height and peak‐to‐peak distance of the tubercles using scaled photographs. Mean tubercle height was standardized as a percentage of the dorsal fin height and ranged from 0.63 to 0.87%. Mean peak‐to‐peak distance ranged from 4.2 ± 2.0 to 5.6 ± 2.0 mm. The microstructure analysis of the dorsal fin leading edge, trailing edge and tubercles revealed an epidermal thickness of 0.7–2.7 mm with the thickest epidermis at the tubercular apex. The epidermis contained three distinct strata (=layers), including the stratum corneum, spinosum, and basale. The stratum corneum was significantly thickened in tubercles, over four times thicker than in the leading or trailing edge of the fin. The stratum spinosum, composed of lipokeratinocytes and lamellar oil bodies, was significantly thinner in the trailing edge than in the other two sites. There was no significant difference in the stratum basale among the three sites. Volume fraction of lipokeratinocytes was significantly higher at the sides of the leading edge and the apex of the tubercles, while volume fraction of lamellar oil bodies was significantly lower at the apex of the tubercles. Though the function of the tubercles is unknown, their position, hardened structure and increased epidermal stratum corneum suggest that they may have hydrodynamic importance. J. Morphol., 2011. © 2010 Wiley‐Liss, Inc.     

MAJOR DECLINE IN NUMBER OF HARBOR SEALS, PHOCA VITULINA RICHARDSI, ON TUGIDAK ISLAND, GULF OF ALASKA

Tugidak Island, located in the Gulf of Alaska, was once the site of one of largest local concentrations of harbor seals ( Phoca vitulina richardsi ) in the world. This population, which probably consisted of about 20,500 animals in the mid-1960s declined by about 85% between 1976 and 1988. The population appeared to decline more rapidly during the late 1970s than during the 1980s. Causes for the decline are not apparent. There appear to be both similarities and dissimilarities between this decline and recent declines in abundance of northern fur seals ( Callorhinus ursinus ) and Steller sea lions ( Eumetopias jubatus ) in the Bering Sea and Gulf of Alaska.     

LONG-TERM TRENDS IN HARBOR SEAL NUMBERS AT TUGIDAK ISLAND AND NANVAK BAY, ALASKA

We conducted land‐based counts of harbor seals (Phoca vitulina richardii) and collected related environmental data at Tugidak Island (Gulf of Alaska, 1994–2000) and Nanvak Bay (Bristol Bay, 1990–2000) to estimate population trends and identify factors influencing counts. At Tugidak Island, the seal population declined substantially during molting from 1976 through the 1980s, stabilized in the early 1990s, and increased at a moderate rate (3.4%/yr, CI: 1.0%–5.8%) from 1994 to 2000. Pups and all seals ashore during pupping increased at higher annual rates of 5.4% (CI: 2.2%–8.8%) and 8.3% (CI: 4.5%–12.3%) from 1994 to 2000 at Tugidak Island. At Nanvak Bay seals declined in abundance between 1975 and 1990 but increased during the 1990s at 9.2%/yr (CI: 7.2%–11.3%) during pupping and 2.1%/yr (CI: 0.6%–3.6%) during molting. Date and time‐of‐day were significant covariates in all analyses. Factors that led to declines at Tugidak Island and Nanvak Bay have seemingly abated sufficiently such that these populations are currently increasing, though still greatly reduced from the 1970s. Index sites are useful adjuncts to aerial surveys, providing survey‐related information not always available from aerial counts, which is useful in survey design and data analysis.     

PUPPING PHENOLOGY AND DEMOGRAPHY OF HARBOR SEALS (PHOCA VITULINA RICHARDSI)ON TUGIDAK ISLAND, ALASKA

The number of seals on shore at Tugidak Island (Gulf of Alaska) declined 72%–85% between 1976 and 1988 and increased during the 1990s. We compared pupping phenology and the ratio of pupping-period counts to molting-period counts between declining (1976–1979) and increasing (1994–1998) years, and examined the sex/age structure of seals ashore during the 1990s. In the 1970s the onset and peak of pupping occurred 6–18 d later than in the 1990s. Rate of pup abandonment was higher in 1978 than in the 1990s. Between 1994 and 1995, the maximum and mean number of seals ashore increased >50%, largely due to an increase in non-pups. From 1995 to 1998, the sex/age structure of seals ashore was similar among years. We observed three to four times as many seals during the molting period than during the pupping period in the 1970s, whereas similar numbers were ashore during these periods in the 1990s, perhaps reflecting changes in demography and/or haul-out behavior. Changes in pupping phenology and demography may reflect environmental changes, such as food availability, and when monitored in conjunction with population counts, may help us better interpret changes in the number of seals ashore.