Cytochrome p450 isozyme protein verified in the skin of southern hemisphere humpback whales (megaptera novaeangliae) : implications for biochemical biomarker assessment

Large mysticete whales represent a unique challenge for chemical risk assessment. Few epidemiological investigations are possible due to the low incidence of adult stranding events. Similarly their often extreme life-history adaptations of prolonged migration and fasting challenge exposure assumptions. Molecular biomarkers offer the potential to complement information yielded through tissue chemical analysis, as well as providing evidence of a molecular response to chemical exposure. In this study we confirm the presence of cytochrome P450 reductase (CPR) and cytochrome P450 isoenzyme 1A1 (CYP1A1) in epidermal tissue of southern hemisphere humpback whales (Megaptera novaeangliae). The detection of CYP1A1 in the integument of the humpback whale affords the opportunity for further quantitative non-destructive investigations of enzyme activity as a function of chemical stress.

Detoxification enzyme activities (CYP1A1 and GST) in the skin of humpback whales as a function of organochlorine burdens and migration status

The activities of glutathione-s-transferase (GST) and cytochrome P-450 1A1 (CYP1A1) enzymes were measured in freshly extracted epidermis of live-biopsied, migrating, southern hemisphere humpback whales (Megaptera novaeangliae). The two quantified enzyme activities did not correlate strongly with each other. Similarly, neither correlated strongly with any of the organochlorine compound groups previously measured in the superficial blubber of the sample biopsy core, likely reflecting the anticipated low levels of typical aryl-hydrocarbon receptor ligands. GST activity did not differ significantly between genders or between northward (early migration) or southward (late migration) migrating cohorts. Indeed, the inter-individual variability in GST measurements was relatively low. This observation raises the possibility that measured activities were basal activities and that GST function was inherently impacted by the fasting state of the sampled animals, as seen in other species. These results do not support the implementation of CYP1A1 or GST as effective biomarkers of organochlorine contaminant burdens in southern hemisphere populations of humpback whales as advocated for other cetacean species. Further investigation of GST activity in feeding versus fasting cohorts may, however, provide some insight into the fasting metabolism of these behaviourally adapted populations.

Feeding Ecology of Humpback Whales in Continental Shelf Waters near Cordell Bank, California

Daytime feeding behavior of humpback whales (Megaptera novaeangliae) in Gulf of the Farallones, California, and adjacent waters was observed during autumn of 1988 to 1990. Bodega Canyon, Cordell Bank, and the Farallon Islands were the primary sites of feeding activity. Fecal samples of whales and zooplankton tows contained euphausiids exclusively, dominated by Thysanoessa spinifera (79%), with lesser amounts of Euphausia pacifica (14%), Nyctiphanes simplex (4%), and Nematoscelis difficilis (3%). In 1988 and 1990, whales also were infrequently observed feeding on small schooling fish, presumably Pacific herring (Clupea pallasii), northern anchovy (Engraulis mordax), and juvenile rockfish (Sebastes spp.). Feeding was the most common behavior observed (52%), and less frequently traveling (23%), milling (21 %), and resting (4%). Whales used different methods to consume euphausiid prey at the surface (0-10 m), in shallow water (11-60 m), and deep water (61-140 m). Humpback whales fed at the surface 56% of time in 1988 and 32% of time in 1990, using primarily lateral lunges to capture swarms of euphausiids. In 1989, no surface feeding was observed; however, deep, long-duration dives were followed by extended surface intervals with many respirations. These 1989 observations coincided with increased prey depth as indicated by depth sounder records of diving whales and prey scattering layers. In 1989, increased prey depth and associated feeding behaviors were strongly associated with unusually high surface temperatures, calm seas, and changes in water circulation. Environmental conditions in 1989 triggered the most intense and wide-spread occurrence of red tide in this region since 1980. Red tide samples collected throughout this period contained Alexandrium (=Gonyaulax) catenella and Noctiluca scintillans. Surface feeding was observed only in 1988 and 1990, when surface prey were available and red tides were very limited in extent, duration, and intensity. Annual variations in humpback whale feeding behavior were related to prey availability which is affected by corresponding environmental conditions. (PDF contains 94 pages)

The effect of predation (current and historical) by humpback whales (Megaptera novaeangliae) on fish abundance near Kodiak Island, Alaska

Humpback whales (Megaptera novaeangliae) are significant marine consumers. To examine the potential effect of predation by humpback whales, consumption (kg of prey daily) and prey removal (kg of prey annually) were modeled for a current and historic feeding aggregation of humpback whales off northeastern Kodiak Island, Alaska. A current prey biomass removal rate was modeled by using an estimate of the 2002 humpback whale abundance. A historic rate of removal was modeled from a prewhaling abundance estimate (population size prior to 1926). Two provisional humpback whale diets were simulated in order to model consumption rate. One diet was based on the stomach contents of whales that were commercially harvested from Port Hobron whaling station in Kodiak, Alaska, between 1926 and 1937, and the second diet, based on local prey availability as determined by fish surveys conducted within the study area, was used to model consumption rate by the historic population. The latter diet was also used to model consumption by the current population and to project a consumption rate if the current population were to grow to reach the historic population size. Models of these simulated diets showed that the current population likely removes nearly 8.83

Historical Catches of Humpback Whales, Megaptera novaeangliae, in the North Atlantic Ocean: Estimates of Landings and Removals

Whaling for humpback whales, Megaptera novaeangliae, in the North At- lantic Ocean has occurred in various forms (e.g. for local subsistence, for oil to be sold commercially, using hand harpoons and deck-mounted cannons, using oar-driven open boats and modern powered catcher boats) from the early 1600’s to the present. Several previous attempts to estimate the total numbers of humpback whales removed were considered close to comprehensive, but some uncertainties remained. Moreover, the statistical uncertainty was not consistently presented with the previous estimates. Therefore, we have pursued several avenues of additional data collection and conducted further analyses to close outstanding data gaps and address remaining issues. Our new estimates of landings and total removals of humpback whales from the North Atlantic are 21,476 (SE=214) and 30,842 (SE=655), respectively. These results include statistical uncertainty, reflect new data and improved analysis methods, and take account of some fisheries for which estimates had not been made previously. The new estimates are not sufficiently different from previous ones to resolve the major inconsistencies and discrepancies encountered in efforts to determine the conservation status of humpback whale populations in the North Atlantic.

Commercial Whaling, Especially for Gray Whales, Eschrichtius robustus, and Humpback Whales, Megaptera novaeangliae, at California and Baja California Shore Stations in the 19th Century (1854–1899)

Shore whaling along North America’s California and Baja California coasts during 1854–99 was ancillary to the offshore and alongshore American whale fishery, which had begun in the North Pacific in the early 1800’s and was flourishing by the 1840’s. From its inception at Monterey, Calif., in the mid 1850’s, the shore fishery, involving open boats deployed from land to catch and tow whales for processing, eventually spread from Monterey south to San Diego and Baja California and north to Crescent City near the California–Oregon border. It had declined to a relict industry by the 1880’s, although sporadic efforts continued into the early 20th century. The main target species were gray whales, Eschrichtius robustus, and humpback whales, Megaptera novaeangliae, with the valuable North Pacific right whale, Eubalaena japonica, also pursued opportunistically. Catch data are grossly incomplete for most stations; no logbooks were kept for these operations as they were for high-seas whaling voyages. Even when good information is available on catch levels, usually as number of whales landed or quantity of oil produced, it is rarely broken down by species. Therefore, we devised methods for extrapolation, interpolation, pro rationing, correction, and informed judgment to produce time series of catches. The resulting estimates of landings from 1854 to 1899 are 3,150 (SE = 112) gray whales and 1,637 (SE = 62) humpback whales. The numbers landed should be multiplied by 1.2 to account for hunting loss (i.e. whales harpooned or shot but not recovered and processed).

Catches of Humpback Whales, Megaptera novaeangliae, by the Soviet Union and Other Nations in the Southern Ocean, 1947–1973

From 1947 to 1973, the U.S.S.R. conducted a huge campaign of illegal whaling worldwide. We review Soviet catches of humpback whales, Megaptera novaeangliae, in the Southern Ocean during this period, with an emphasis on the International Whaling Commission’s Antarctic Management Areas IV, V, and VI (the principal regions of illegal Soviet whaling on this species, south of Australia and western Oceania). Where possible, we summarize legal and illegal Soviet catches by year, Management Area, and factory fleet, and also include information on takes by other nations. Soviet humpback catches between 1947 and 1973 totaled 48,702 and break down as follows: 649 (Area I), 1,412 (Area II), 921 (Area III), 8,779 (Area IV), 22,569 (Area V), and 7,195 (Area VI), with 7,177 catches not currently assignable to area. In all, at least 72,542 humpback whales were killed by all operations (Soviet plus other nations) after World War II in Areas IV (27,201), V (38,146), and VI (7,195). More than one-third of these (25,474 whales, of which 25,192 came from Areas V and VI) were taken in just two seasons, 1959–60 and 1960–61. The impact of these takes, and of those from Area IV in the late 1950’s, is evident in the sometimes dramatic declines in catches at shore stations in Australia, New Zealand, and at Norfolk Island. When compared to recent estimates of abundance and initial population size, the large removals from Areas IV and V indicate that the populations in these regions remain well below pre-exploitation levels despite reported strong growth rates off eastern and western Australia. Populations in many areas of Oceania continue to be small, indicating that the catches from Area VI and eastern Area V had long-term impacts on recovery.

Distribution and abundance of humpback whales (Megaptera novaeangliae) and other marine mammals off the northern Washington coast

We examined the summer distribution of marine mammals off the northern Washington coast based on six ship transect surveys conducted between 1995 and 2002, primarily from the NOAA ship McArthur. Additionally, small boat surveys were conducted in the same region between 1989 and 2002 to gather photographic identification data on humpback whales (Megaptera novaeangliae) and killer whales (Orcinus orca) to examine movements and population structure. In the six years of ship survey effort, 706 sightings of 15 marine mammal species were made. Humpback whales were the most common large cetacean species and were seen every year and a total of 232 sightings of 402 animals were recorded during ship surveys. Highest numbers were observed in 2002, when there were 79 sightings of 139 whales. Line-transect estimates for humpback whales indicated that about 100 humpback whales inhabited these waters each year between 1995 and 2000; in 2002, however, the estimate was 562 (CV= 0.21) whales. A total of 191 unique individuals were identified photographically and mark recapture estimates also indicated that the number of animals increased from under 100 to over 200 from 1995 to 2002. There was only limited interchange of humpback whales between this area and feeding areas off Oregon and California. Killer whales were also seen on every ship survey and represented all known ecotypes of the Pacific Northwest, including southern and northern residents, transients, and offshore-type killer whales. Dall’s porpoise (Phocoenoides dalli) were the most frequently sighted small cetacean; abundance was estimated at 181−291 individuals, except for 2002 when we observed dramatically higher numbers (876, CV= 0.30). Northern fur seals (Callorhinus ursinus) and elephant seals (Mirounga angustirostris) were the most common pinnipeds observed. There were clear habitat differences related to distance offshore and water depth for different species.

Assessing the Impact of Shipping Vessel Noise on Humpback Whales

The ocean is a hub of noise. Bioacoustic noise, noise from precipitation and wind, and noise from oceanic shelf slides and other geologic processes have occurred consistently as marine species have evolved over time. However, with the discovery of oceanic oil and gas reserves, submarine systems, ship propulsion and the emergence of global trade, anthropogenic sources of sound have added significant quantities of sound to the oceanic system. Shipping has been found to be the largest input of low-frequency anthropogenic noise and Humpback Whales (Megaptera novaengliae), known to be the most vocal marine species, have an auditory sensitivity that falls within the range of frequencies emitted by shipping vessels. As Humpback Whales are heavily dependent on vocalizations, for reasons relatively unconfirmed, a better understanding of why they sing and how their communication is being impacted by vessel noise is critical. Evaluating existing literature both on Humpback behavior and communication, the mechanics of their communication, sound emissions from modern ships, oceanic sound transmission, and studies regarding Humpback's exposure to other sources of low-frequency anthropogenic noise, it is clear that more research is needed to draw any causational conclusions between vessel noise and detrimental impacts on Humpback Whales. With a projected increase in global consumption and vessel traffic, there is an urgent need for further research exploring shipping noise impacts and behavioural alterations of Humpbacks. Existing research has shown changes in Humpback communication when exposed to low-frequency sonar noise, however few studies have been conducted on their communication when in close proximity to shipping vessels. In order for the impacts to be properly assessed, preliminary understanding of humpback communication, their auditory thresholds and more studies between vessel noise exposure and Humpback Whale behavior must be conducted.

Fluctuating abundance of humpback whales (Megaptera novaeangliae) in a calving ground off coastal Brazil

The humpback whale (Megaptera novaeangliae) population that uses Abrolhos Bank, off the east coast of Brazil as a breeding ground is increasing. To describe temporal changes in the relative abundance of humpback whales around Abrolhos, seven years (1998-2004) of whale count data were collected during July through to November. During one-hour-scans, observers determined group size within 9.3 km (5 n.m.) of a land-based observing station. A total Of 930 scans, comprising 7996 sightings of adults and 2044 calves were analysed using generalized linear models that included variables for time of day, day of the season, years and two-way interactions as possible predictors. The pattern observed was the gradual build-up and decline in whale counts within seasons. Patterns and peaks of adult and calf counts varied among years. Although fluctuation was observed, there was generally an increasing trend in adult counts among years. Calf counts increased only in 2004. These fluctuations may have been caused by some environmental conditions in humpback whales` summering grounds and also by changes in spatial-temporal concentrations in Abrolhos Bank. The general pattern observed within the study area mirrored what was observed in the whole Abrolhos Bank. Knowledge of the consistency with which humpback whales use this important nursing area should prove beneficial for designing future monitoring programmes especially related to whale watching activities around Abrolhos Archipelago.

Catches of Humpback Whales, Megaptera novaeangliae, by the Soviet Union and Other Nations in the Southern Ocean, 1947–1973

The great whales of the Southern Ocean were extensively exploited by modern whaling methods, with the first catches made in the Falkland Islands Dependencies region of IWC Management Area II in 1904 (Tønnesson and Johnsen, 1982; Hart, 2006). Exploitation went through several phases. Populations of humpback whales, Megaptera novaeangliae, and blue whales, Balaenoptera musculus, around South Georgia crashed around the time of World War I, and further exploitation occurred in other regions into the 1930’s. There was a hiatus in whaling during World War II, but large-scale catches resumed in Antarctic waters after 1945.

Catches of Humpback Whales in the Southern Ocean, 1947-1973

We review catches of humpback whales (Megaptera novaeangliae) in the Southern Ocean during the period following World War II, with an emphasis on Areas IV, V and VI (the principal regions of illegal Soviet whaling on this species). Where possible, we summarize legal and illegal Soviet catches by year, Area and factory fleet, and also include information on takes by other nations. Soviet humpback catches between 1947 and 1973 totaled 48702 and break down as follows: 649 (Area I), 1412 (Area II), 921 (Area III), 8779 (Area IV), 22569 (Area V) and 7195 (Area VI), with 7177 catches not assignable to area. In all, at least 72542 humpback whales were killed by all operations (Soviet plus other nations) after World War 2 in Areas IV (27201), V (38146) and VI (7195). More than a third of these (25474 whales, of which 25192 came from Areas V and VI) were taken in just two seasons, 1959/60 and 1960/61. The impact of these takes, and of those from Area IV in the late 1950's, is evident in the sometimes dramatic declines in catches at shore stations in Australia, New Zealand and Norfolk Island. When compared to recent estimates of abundance, the large removals from Areas IV and V indicate that the populations in these regions remain well below pre-exploitation levels despite reported strong growth rates off eastern and western Australia. Populations in many areas of Oceania continue to be small, indicating that the catches from Area VI and eastern Area V had long-term impacts on recovery.

Acoustic Detections of Singing Humpback Whales in Deep Waters Off The British Isles

From October 1996 through September 1998, we used bottom-mounted hydrophone arrays to monitor deep-water areas north and west of the British Isles for songs of humpback whales (Megaptera novaeangliae). Singing humpbacks were consistently detected between October and March from the Shetland- Faroe Islands south to waters west of the English Channel. Temporal and geographic patterns of song detections, and movements of individually tracked whales, exhibited a southwesterly trend over this period, but with no corresponding northward trend between April and September. These results, together with a review of historical data from this area, suggest that the offshore waters of the British Isles represent a migration corridor for humpbacks, at least some of which summer in Norwegian (and possibly eastern Icelandic) waters. The migratory destination of the detected animals remains unknown, but the limited data suggest that these whales are bound primarily for the West Indies rather than historical breeding areas off the northwestern coast of Africa. Humpbacks detected in British waters after early to mid- March probably do not undertake a full migration to the tropics. These data provide further evidence that singing is not confined to tropical waters in winter, but occurs commonly on migration even in high latitudes.

A Novel Behavior Observed in Humpback Whales on Wintering Grounds at Abrolhos Bank (Brazil)

We describe a novel behavior, termed “tail-up,” observed in humpback whales (Megaptera novaeangliae) on wintering grounds on Abrolhos Bank, Brazil. The behavior involves the whale positioned vertically in the water column with its tail in the air. Wirh the exception of calves, tail-up was observed in all social classes, and its frequency increased through the end of the season. Tail-ups were recorded in 144 (5.8%) of 2,465 groups of whales observed from a shore station, and in 297 (14.9%) of 1,996 groups observed from vessel surveys; biases in each method suggest that the true frequency lies between these sources. One hundred and fifty-two hours of continuous sampling showed that the duration of tail-up events lasted from a few seconds to 12 min and was longest in groups comprised of a single adult. The maximum duration of a recorded period that consistently included tail-up was 10 h; however, some individuals were observed to engage in the behavior at night and for four consecutive days. Tail-up movement speed did not vary by social class; however, it varied according to wind direction and speed. The characteristics of tail-up that we observed showed that it differed from the descriptions of similar behaviors in other cetacean species. The function of tail-up is unknown, but we suggest that it may be a multifunctional behavior.

Segregation of migration by feeding ground origin in North Atlantic humpback whales (Megaptera novaeangliae)

Results from a large-scale, capture–recapture study of humpback whales Megaptera novaeangliae in the North Atlantic show that migration timing is influenced by feeding ground origin. No significant differences were observed in the number of individuals from any feeding area that were re-sighted in the common breeding area in the West Indies. However, there was a relationship between the proportion (logit transformed) of West Indies sightings and longitude (r2 = 0.97, F1,3 = 98.27, P = 0.0022) suggesting that individuals feeding farther to the east are less likely to winter in the West Indies. A relationship was also detected between sighting date in the West Indies and feeding area. Mean sighting dates in the West Indies for individuals identified in the Gulf of Maine and eastern Canada were significantly earlier than those for animals identified in Greenland, Iceland and Norway (9.97 days, t179 = 3.53, P = 0.00054). There was also evidence for sexual segregation in migration; males were seen earlier on the breeding ground than were females (6.63 days, t105 = 1.98, P = 0.050). This pattern was consistently observed for animals from all feeding areas; a combined model showed a significant effect for both sex (F1 = 5.942, P = 0.017) and feeding area (F3 =4.756, P=0.0038). The temporal difference in occupancy of the West Indies between individuals from different feeding areas, coupled with sexual differences in migratory patterns, presents the possibility that there are reduced mating opportunities between individuals from different high latitude areas.