COSEWIC Assessment and Status Report on the Humpback Whale in Canada-2003

  1. Table of Contents
  2. Assessment Summary
  3. Executive Summary
  4. Species Information
  5. Distribution
  6. Habitat
  7. General Biology
  8. Population Sizes and Trends
  9. Limiting Factors and Threats
  10. Special Significance of the Species
  11. Existing Legal Protection
  12. Technical Summary: North Pacific population
  13. Technical summary: Western North Atlantic population
  14. Acknowledgements
  15. Literature Cited
  16. The Author

COSEWIC status reports are working documents used in assigning the status of wildlife species suspected of being at risk. This report may be cited as follows:

Please note: Persons wishing to cite data in the report should refer to the report (and cite the author(s)); persons wishing to cite the COSEWIC status will refer to the assessment (and cite COSEWIC). A production note will be provided if additional information on the status report history is required.

COSEWIC 2003. COSEWIC assessment and update status report on the humpback whale Megaptera novaeangliae in Canada. Committee on the Status of Endangered Wildlife in Canada. Ottawa. viii + 25 pp.

Baird, R.W. 2003. Update COSEWIC status report on the humpback whale Megaptera novaeangliae in Canada in COSEWIC assessment and update status report on the humpback whale Megaptera novaeangliae in Canada. Committee on the Status of Endangered Wildlife in Canada. Ottawa. 1-25 pp.

Previous report (s):

Whitehead, H. 1985. Update COSEWIC status report on the humpback whale Megaptera novaeangliae in Canada. Committee on the Status of Endangered Wildlife in Canada. Ottawa. 23 pp.

Hay, K. 1982. COSEWIC status report on the humpback whale Megaptera novaeangliae in Canada. Committee on the Status of Endangered Wildlife in Canada. Ottawa. 22 pp.

Également disponible en français sous le titre Rapport du COSEPAC sur la situation du rorqual à bosse (Megaptera novaeangliae)au Canada - Mise à jour

Cover illustration:
Humpback Whale -- Illustration, Judie Shore, Richmond Hill, Ontario

©Her Majesty the Queen in Right of Canada, 2003
Catalogue No.CW69-14/233-2002E-IN
ISBN 0-662-32922-8

Assessment Summary

Common name: Humpback Whale (North Pacific Population)

Scientific name: Megaptera Novaeangliae

Status: Threatened

Reason for designation: Heavily reduced by whaling, the North Pacific population appears to be increasing. The number of animals that use British Columbia waters is probably in the low hundreds. The high-level of feeding ground fidelity suggests that if animals are exterminated from a particular area, it is unlikely that it will be rapidly repopulated from other areas. Two extirpated British Columbia populations have shown no sign of rescue. Humpbacks are occasionally entangled in fishing gear, though the number entangled is not thought to threaten or limit the population. In summary, humpback whales that use British Columbia waters appear to be well below historical numbers and have not returned to some portions of their former range.

Canadian occurrence: Pacific Ocean

Status history: The Western North Atlantic and North Pacific populations were given a single designation of Threatened in April 1982. Split into two populations in April 1985 (Western North Atlantic population and North Pacific population). North Pacific population was designated Threatened in 1985. Status re-examined and confirmed Threatened in May 2003. Last assessment based on an update status report.

Common name: Humpback Whale (Western North Atlantic Population)

Scientific name: Megaptera Novaeangliae

Status: Not at Risk

Reason for designation: Although heavily reduced by whaling, this well-studied population seems to have regrown to at least a substantial proportion of its pre-whaling size. The population does face threats (including entanglement in fishing gear, habitat degradation on breeding grounds, possible resumption of commercial whaling), but neither the North Atlantic population nor any of its breeding sub-populations is at risk from current activity levels or levels that may reasonably be foreseen in the next few years.

Canadian occurrence: Atlantic Ocean

Status history: The Western North Atlantic and North Pacific populations were given a single designation of Threatened in April 1982. Split into two populations in April 1985 (Western North Atlantic population and North Pacific population). Western North Atlantic population was designated Special Concern in 1985. Status re-examined and de-listed (Not at Risk) in May 2003. Last assessment based on an update status report.

Executive Summary
Description

Humpback whales are one of the larger cetaceans, typically reaching lengths of about 13 m for males and 14 m for females. Humpbacks are easily recognizable, with their dark dorsal colouration, large size, extremely long pectoral flippers (to nearly one-third of the body length), tendency to raise their tail flukes above the surface when they dive, and frequent aerial behaviour.

Distribution

Humpback whales are found in tropical, temperate and sub-polar waters world-wide. In Canada, humpbacks are found on both the east and west coasts, extending north to Labrador in the east and into northwestern Alaska in the west. Humpbacks have long been considered a coastal species, though recent acoustic evidence shows that some individuals do use offshore areas during periods when they are not expected to be migrating. Like most other baleen whales, humpback whales exhibit seasonal migrations from high-latitude feeding areas in summer to low-latitude breeding and calving areas in winter, thus Canadian waters are used primarily for feeding.

Habitat

In the North Pacific, humpback whales feed from California north along the west coast of North America into the Bering Sea. Calving occurs in three broad areas, in the western Pacific south of Japan to the northern Philippines, in the main Hawaiian Islands, and off Mexico and central America south to Panama. Acoustic and movement evidence suggests that some exchange occurs between these breeding populations. However, individuals show considerable fidelity to feeding areas, that is, a number of discrete stocks can be defined based on feeding areas. Humpback whales in British Columbia are thought to belong to at least two distinct feeding stocks; those identified off southern British Columbia show little interchange with those seen off northern British Columbia. Humpback whales identified in the southern province show only a very low level of interchange with those seen off California/Oregon/Washington and should probably not be considered part of this stock for management purposes. Those in the northern province show some interchange with whales identified in SE Alaska that are considered part of the “Central North Pacific” stock. There has been relatively little effort to determine the boundary of these stocks in British Columbia. Humpback whales from British Columbia have been identified in all three breeding regions in the North Pacific (Japan, Hawaii, and Mexico). Historically in the North Atlantic there were two breeding areas, in the West Indies and off the Cape Verde Islands, off western Africa. Today most humpbacks from both the western and eastern North Atlantic appear to use the West Indies for calving/breeding, though small numbers (most likely from the eastern Atlantic) may breed in the Cape Verdes. Three feeding stocks are recognized from eastern Canada: the Gulf of Maine (which includes animals from the Scotian Shelf), the Gulf of St. Lawrence, and Newfoundland/Labrador (which includes animals from the Strait of Belle Isle). Some interchange between these areas occurs, and juvenile whales from all stocks may mix in mid-latitude feeding areas off the mid-Atlantic states of the U.S.

General Biology

Gestation lasts 11 to 12 months, and a single calf is born every one to three years (average of 2.38 years in the N. Atlantic). Calves are born primarily between December and April in the northern hemisphere. Life expectancy is unknown, although maximum longevity is known to be at least 48 years. In the North Atlantic, mean age at attainment of sexual maturity for females is five years, thus females give birth (on average) at six years of age.

Population Size and Trends

The most recent population estimates available are from 1992/1993 for both the North Atlantic (11,570 individuals) and North Pacific (6,010 individuals), and both estimates are known to under-estimate true population sizes. A number of estimates for feeding stocks within these basins are also available, although reliable estimates for Canadian feeding stocks are not available. Populations in both the Atlantic and Pacific appear to be growing and may be approaching pre-whaling levels in the Atlantic.

Limiting Factors and Threats

Potential threats to humpback whales in Canadian waters today include a reduction in their prey base, incidental mortality in fisheries, ship strikes, and disturbance or injury in association with vessel traffic and/or high-intensity underwater sounds, though none of these threats are thought to currently jeopardize the growth of either population.

Existing Protection

Commercial harvesting of humpback whales has been banned by the International Whaling Commission in the North Pacific since 1965 and in the North Atlantic since 1955. The only hunting that is currently permitted is for small numbers of animals for Aboriginal subsistence purposes (St. Vincent and the Grenadines are authorized to catch two per year). Within Canada, management of humpbacks falls under the responsibility of the Department of Fisheries and Oceans. The Marine Mammal Regulations state that "no person should disturb a marine mammal except when under the authorities of these regulations”.

The Committee on the Status of Endangered Wildlife in Canada (COSEWIC) determines the national status of wild species, subspecies, varieties, and nationally significant populations that are considered to be at risk in Canada. Designations are made on all native species for the following taxonomic groups: mammals, birds, reptiles, amphibians, fish, lepidopterans, molluscs, vascular plants, lichens, and mosses.

COSEWIC comprises representatives from each provincial and territorial government wildlife agency, four federal agencies (Canadian Wildlife Service, Parks Canada Agency, Department of Fisheries and Oceans, and the Federal Biosystematic Partnership), three nonjurisdictional members and the co-chairs of the species specialist groups. The committee meets to consider status reports on candidate species.

Species: Any indigenous species, subspecies, variety, or geographically defined population of wild fauna and flora.

Extinct (X): A species that no longer exists.

Extirpated (XT): A species no longer existing in the wild in Canada, but occurring elsewhere.

Endangered (E): A species facing imminent extirpation or extinction.

Threatened (T): A species likely to become endangered if limiting factors are not reversed.

Special Concern (SC)*: A species of special concern because of characteristics that make it particularly sensitive to human activities or natural events.

Not at Risk (NAR)**: A species that has been evaluated and found to be not at risk.

Data Deficient (DD)***: A species for which there is insufficient scientific information to support status designation.

* Formerly described as "Vulnerable" from 1990 to 1999, or "Rare" prior to 1990.
** Formerly described as "Not In Any Category", or "No Designation Required."
*** Formerly described as "Indeterminate" from 1994 to 1999 or "ISIBD" (insufficient scientific information on which to base a designation) prior to 1994.

The Committee on the Status of Endangered Wildlife in Canada (COSEWIC) was created in 1977 as a result of a recommendation at the Federal-Provincial Wildlife Conference held in 1976. It arose from the need for a single, official, scientifically sound, national listing of wildlife species at risk. In 1978, COSEWIC designated its first species and produced its first list of Canadian species at risk. Species designated at meetings of the full committee are added to the list.

Environment Environnement
Canada Canada

Canadian Wildlife Service canadien
Service de la faune

The Canadian Wildlife Service, Environment Canada, provides full administrative and financial support to the COSEWIC Secretariat.

Species Information
Name, Classification and Taxonomy

The humpback whale, Megaptera novaeangliae (Borowski, 1781) is the only member of the genus Megaptera. No subspecies are recognized. Differences in timing of reproduction, as well as pigmentation pattern differences (Pike 1953; Rosenbaum et al. 1995), suggest that northern and southern hemisphere populations are reproductively isolated. However, southeastern Pacific humpbacks may overwinter in tropical waters of the northern hemisphere, in areas also used by northern hemisphere humpbacks (Stone et al. 1990; Acevedo and Smultea 1995; Florez-Gonzalez et al. 1998), suggesting the potential for genetic exchange between northern and southern hemisphere populations (Baker et al. 1993). Based on a review of available genetic data, Baker and Palumbi (1997) note that migration between ocean basins probably occurs once or twice per generation.

Description

The humpback whale (Figure 1) is one of the larger cetaceans, with adult males and adult females typically reaching lengths of about 13 m and 14 m, respectively (Chittleborough 1965; Stevick 1999). Maximum recorded length is 17.4 meters (Chittleborough 1965). Humpbacks are easily recognizable, with their dark dorsal colouration, large size, extremely long pectoral flippers (to nearly one-third of the body length), tendency to raise their tail flukes above the surface when they dive, and frequent aerial behaviour (Figure 1). Males and females are generally similar in appearance, though females have a hemispherical lobe in the genital region (Glockner 1983), and the underside of the tail flukes in males tends to be whiter than in females (Pike 1953; Allen et al. 1994), though this latter feature cannot be confidently used to determine sex.

Distribution

Humpback whales can be found in tropical, temperate and sub-polar waters around the world. In Canada, humpbacks are found on both the east and west coasts. On the east coast their range extends north to Labrador, while on the west coast their range extends north to northwest Alaska (Whitehead 1987). Like most other baleen whales, humpback whales exhibit seasonal migrations from high-latitude feeding areas in summer to low-latitude breeding and calving areas in winter (with the possible exception of the Arabian Sea population – Whitehead 1985; Mikhalev 1997), thus Canadian waters are used primarily for feeding. Such migrations suggest strong seasonal differences in distribution. However, in some high-latitude feeding areas in particular, it is clear that some humpbacks remain well into the breeding season, or perhaps individual variability in timing of migrations results in the presence of some individuals in high latitude areas during all months of the year (Straley 1990; Brown et al. 1995; Craig and Herman 1997; S. Todd, personal communication). In addition, there


Figure 1. Top. Two humpback whales, showing the characteristic profile of the back (left) and the tail flukes being raised above the surface on a dive (right). Bottom. Humpback whale breaching, showing the characteristically long flippers. Photos by the author.

is some evidence that juveniles may remain in mid-latitude areas for feeding, while the majority of individuals have moved to lower-latitudes (Swingle et al. 1993; Laerm et al. 1997; Barco et al. 2002). Humpback whales have long been characterized as a coastal species, though recent acoustic evidence shows that some individuals do use offshore areas during periods when they are not expected to be migrating (C. Clark, personal communication). Although the occurrence of feeding in traditional breeding areas is rare, some feeding behaviour has been observed (Salden 1989; Baraff et al. 1991; Gendron and Urban 1993). Movements between feeding and breeding grounds can be both direct (Mate et al. 1998) and rapid (Gabriele et al. 1996), though relatively little is known about migratory routes or behaviour during migration (Mate et al. 1998; Norris et al. 1999).

Habitat

In the North Pacific, humpback whales feed from California north along the west coast of North America through to the Aleutians (Figure 2). Calving occurs in three broad areas, in the western Pacific south of Japan south to the northern Philippines, in the windward Hawaiian Islands, and off the coast of Mexico and central America south to Panama. Acoustic and movement evidence suggests that some exchange between these breeding populations occurs (Payne and Guinee 1983; Baker et al. 1986; Helweg et al. 1990; Darling and Cerchio 1993; Darling et al. 1996; Calambokidis et al. 1997, 2000, 2001; Salden et al. 1999). However, both genetic and sighting data between feeding and breeding areas suggest that individuals show considerable fidelity to feeding areas, that is, a number of discrete stocks can be defined based on feeding areas (Darling and Jurasz 1983; Baker et al. 1986; Calambokidis et al. 1996, 1997; Waite et al. 1999). Baker and Palumbi (1997) note that in the North Pacific, long-term migratory exchange between some feeding populations (e.g., southeastern Alaska and central California) may be less than one individual per generation. The U.S. National Marine Fisheries Service has used this evidence to manage three discrete feeding sub-stocks in the North Pacific (see references above and Barlow et al. 1998; Hill and DeMaster 1998). They list two of these as found in British Columbia; the “California/Oregon/Washington and Mexico” stock extending to southern British Columbia to feed and the “Central North Pacific” stock feeding from northern British Columbia west to Kodiak Island, Alaska. The stock structure of humpback whales in British Columbia is more complex and not as clear.

There is some evidence that whales that feed off British Columbia represent two feeding aggregations separate from those to the north and south. Humpback whales that feed off California, Oregon, and southern Washington show a high rate of interchange among areas within this range but very low rates of interchange with the whales that feed off northern Washington and southern British Columbia, and no interchange with whales off northern British Columbia (Calambokidis et al. 1996, 2001). Additionally, no interchange has been found to date between humpback whales that have been identified off southern and northern British Columbia (Calambokidis et al. 2001; G. Ellis, personal communication). While about a half-dozen matches have been found between whales of northern British Columbia and those off southeastern Alaska, this match rate appears fairly low (Ellis and Straley, unpublished data; Calambokidis et al. 2001).

There are also differences in the migratory destinations of whales in these different feeding areas. Humpback whales that feed off California, Oregon, and Washington primarily winter off Mexico and Central America (Steiger et al. 1991; Calambokidis et al. 2000, 2001, Urban et al. 2000) and those that feed of southeastern Alaska primarily winter in Hawaii (Darling and Jurasz 1983; Baker et al. 1986; Calambokidis et al. 2001). In contrast to this, humpback whales that feed off both southern and northern British Columbia migrate to several wintering grounds without a clear preference. This has included migrations from both regions of British Columbia to Mexico (including both the mainland and the offshore Revillagigedos), Hawaii, and the breeding island of Ogasawara off Japan (Darling and Jurasz 1983; Darling et al. 1996; Urban et al. 2000; Calambokidis et al. 2001).

Figure 2. Humpback whale primary feeding and breeding areas (a)North Atlantic. (b) North Pacific. Feeding areas are represented by light stipple and breeding areas by dark shading. Broken arrows indicate undocumented or poorly documented migrations. In most regions, actual travel routes are not known, and arrows merely connect migratory endpoints. Map reproduced from Stevick et al. (2002).

In the North Atlantic, humpbacks from both the western and eastern North Atlantic appear to use the West Indies for calving/breeding (Whitehead 1982; Martin et al. 1984; Katona and Beard 1990; Larsen et al. 1996; Stevick et al. 1998, 2003), although individuals from the eastern North Atlantic are less likely to overwinter in the West Indies (Stevick et al. 2003). Small numbers of humpbacks have been observed in the eastern tropical Atlantic during winter, off the Cape Verde Islands (Reiner et al. 1996), though it is not known which feeding population(s) these individuals are from. This area appears historically to have been a major wintering/breeding area (Reeves et al. 2002), and today may represent a secondary wintering/breeding ground for eastern North Atlantic humpbacks (Reiner et al. 1996; Hazevoet and Wenzel 2000). As with the North Pacific, considerable evidence for segregation of feeding stocks in the North Atlantic also exists, based on genetic results and photo-identification data (Clapham and Mayo 1987; Katona and Beard 1990; Clapham et al. 1993; Palsboll et al. 1995; Larsen et al. 1996; Waring et al. 2000). In the western North Atlantic four feeding stocks are recognized, three of which are found in Canadian waters. These four stocks are the “Gulf of Maine” (which includes animals from the Bay of Fundy and the Scotian Shelf; Clapham et al. 2001), the “Gulf of St. Lawrence”, “Newfoundland and Labrador”, and “western Greenland” (Katona and Beard 1990; Waring et al. 2000). Considerable research has been undertaken on the Gulf of Maine stock; however, in recent years less research has been undertaken on other populations. Based on genetic markers, animals from the Strait of Belle Isle appear to be from the Newfoundland/Labrador stock (P. Palsboll, personal communication). Based on photo-identification, there is a higher level of exchange of individuals between the Gulf of St. Lawrence and the Newfoundland/Labrador populations than for other populations, thus it is possible that they should be considered one stock (Anonymous 2001). Juvenile individuals from the Gulf of Maine and Newfoundland stocks do mix in mid-latitude areas during the winter months (Barco et al. 2002). The International Whaling Commission recently undertook a comprehensive review of North Atlantic humpbacks, and concluded that the appropriate unit for management is that of the feeding sub-stock (Anonymous 2001). There is evidence, however, of some spatial structuring within feeding stocks (Robbins et al. 2001), that is, there are apparently preferred feeding areas for individuals within these broader regions.

General Biology
Reproduction

Gestation lasts 11 to 12 months (Chittleborough 1958), and a single calf is born. In the northern hemisphere calves are born over a diffuse period lasting from December through April, with the peak number of births in January and February. Mating has never been positively documented, though mating-related activities (e.g., males escorting females and defending access from other males) also occurs over this same time period. Most calves are weaned at about a year of age, though a few stay with their mother for two years (Clapham 1992), and some are weaned at less than a year (Baraff and Weinrich 1993). In the North Atlantic, mean age at attainment of sexual maturity for females is five years (Clapham 1992), thus females give birth (on average) at six years of age. There is some suggestion that North Pacific humpbacks may have a later average age at first birth, though more research is necessary to confirm this (Gabriele 1992; Straley et al. 1993; J. Straley, personal communication). Humpback whales may reproduce annually, though two or three-year calving intervals are more common (Clapham and Mayo 1990; Weinrich et al. 1993; Straley et al. 1994). Barlow and Clapham (1997) estimated a mean inter-birth interval for North Atlantic females of 2.38 years, a youngest age at first parturition of 5 years, and a calf survival rate of 0.875, giving a population growth rate of 1.065. Rates of increase of east Australian humpbacks are on the order of 10% per year (Paterson et al. 1994). Longevity is unknown, but the oldest animal recorded appears to be a 48-year-old individual from western Australia (Chittleborough 1965). However, humpbacks likely live much longer, as Chittleborough’s (1965) sample was of whales taken after a considerable period of whaling, when most of the largest (and thus oldest) whales had probably already been killed.

Population Sizes and Trends

All humpback populations were drastically reduced due to commercial whaling; some evidence suggests that perhaps 90-95% of the world-wide population was killed (Johnson and Wolman 1984). A number of estimates from both the North Pacific and North Atlantic have been presented over the past 30 years (summarized by Whitehead 1987). Each method of estimating population size carries with it a number of biases, making comparisons between estimates difficult.

There are no rigorous pre-exploitation population estimates for either the North Pacific or North Atlantic. For the North Pacific, the pre-exploitation population has been reported at approximately 15,000 individuals (Rice 1978). The post-whaling population (1966) size has been reported from 1,200 to 1,400 individuals (Gambell 1975; Johnson and Wolman 1984), though the techniques used to produce these estimates were extremely imprecise (Calambokidis et al. 1997). Estimates from the late 1970s and early 1980s for portions of the North Pacific ranged from about 1,000-2,000 individuals (Darling and Morowitz 1986; Baker and Herman 1987). Calambokidis et al. (1997) produced the most recent estimate for the North Pacific (from 1992-1993) of 6,010 animals (SE = 474). They discuss a number of directional biases that should be taken into account, which suggest that the estimate of about 6,000 animals underestimates the true abundance (which might be closer to 8,000 individuals). Based on photo-identification results, population estimates for humpbacks off California have increased at an average annual rate of 8% between 1988 and 1998 (Calambokidis et al. 1999). Off Hawaii, based on aerial survey results from 1977-80 and from 1990, Mobley et al. (1999a) show evidence of a local increase in numbers on the Hawaiian wintering grounds (see also Cerchio 1998). Aerial surveys in Hawaii in 1993, 1995 and 1998 show further evidence of an increase in numbers, at an annual rate of 7% (Mobley et al. 1999b).

No estimates exist for numbers that use British Columbia waters, though they are probably in the hundreds. Numbers present today are only a small fraction of those found historically (Pike and MacAskie 1969; Merilees 1985; Gregr et al. 2000), though off some areas of British Columbia (e.g., the Queen Charlotte Islands) they are the most frequently reported baleen whale (Ford et al. 1994). At least 5,638 humpback whales were killed off British Columbia from 1908 to 1967, with the highest catches prior to 1917 (Gregr et al. 2000). Declines in catch rates after 1917 indicated a depletion in these populations and this followed a different trend from the declines noted off California and Alaska, indicating the B.C. whales were part of a distinct feeding aggregation (Gregr et al. 2000). Humpback whales have not returned to all of the areas they were once common. Commercial whaling for humpback whales in British Columbia in the early 1900s including hunting of humpback whales in inside waters well into the winter. A whaling station operated at Page’s Lagoon from 1907-09 and was abandoned once the supply of whales was exhausted (Tonnessen and Johnsen 1982; Webb 1988). Humpback whales remain uncommon today in the Strait of Georgia and other inside waters in this region. Off southern British Columbia and northern Washington, 115 unique humpback whales have been identified from 1990 to 2000; the entire population may not be much larger than this because only a third of the whales identified in 2000 had not been previously seen (J. Calambokidis, unpublished data). Off northern British Columbia, where analysis of a long-term dataset is still underway, the number of humpback whales appears somewhat larger with over 500 unique individuals identified from 1989 to 2001 (DFO, unpublished. data; J. Ford, personal communication).

For the North Atlantic, estimates from the mid-1800s (after some whaling had already been undertaken) were of between about 4,000 and 6,000 individuals (Reeves and Mitchell 1982; Breiwick et al. 1983; Whitehead 1987). Post-whaling estimates (from the late 1960s or early 1970s) were of between about 800 and 1,300 individuals (Mitchell 1973; Winn et al. 1975). Stevick et al. (2001) produced an estimate of 11,570 individuals (CV=0.069), for the entire North Atlantic. This estimate is known to be negatively biased (Anonymous 2001). No more recent basin-wide estimates are available. Two recent estimates are available for the Gulf of Maine. A mark-recapture estimate from 1992/93 for the Gulf of Maine indicated a population of 652 individuals (CV = 0.15; Waring et al. 2000), while a 1999 line-transect estimate for the same population was of 816 individuals (CV = 0.45; Palka 2000). Estimates for the remainder of eastern Canadian waters are problematic due to spatially uneven sampling (Anonymous 2001). For the 1992/93 period there is an estimate from eastern Canada of 2,509 individuals (CV = 0.077), though this estimate is almost certainly negatively biased due to heterogeneity in sampling (Anonymous 2001). For the West Indies breeding population, data from 1979 through 1993 suggest an annual population growth of 3% (Stevick et al. 2001).

Limiting Factors and Threats
Natural Mortality

There are a number of potential sources of natural mortality, including predation, parasitism, disease, biotoxins, and accidental beaching or entrapment. Approximately one third of humpbacks in the western North Atlantic have been reported to have scars originating from killer whale (Orcinus orca) attacks (Katona et al. 1980). Clapham (2000) notes that the vast majority of individuals with killer whale scars have such scars the first time they are seen, suggesting that attacks occur primarily on calves (see also Naessig 1999). Predation by killer whales (Jefferson et al. 1991; Florez-Gonzalez et al. 1994), false killer whales (Pseudorca crassidens - Hoyt 1983), or large sharks (particularly on calves on the breeding grounds – Glockner-Ferrari and Ferrari 1997; Mazzuca et al. 1998) appear to be relatively infrequent. However, Corkeron and Connor (2000) argue that avoidance of predation by killer whales in high-latitude areas may be partially responsible for the migratory behavior exhibited by this species. In terms of parasites, while humpback whales externally carry cyamid lice, it is unlikely they contribute to mortality in any way. Little is known about internal parasites or disease processes in humpback whales (Clapham 2000). Humpbacks generally do not mass strand (although S. Todd, personal communication, reports a stranding of three individuals off Newfoundland); most single stranded animals likely strand due to debilitation from disease (or wash up after being killed in fishing gear or from ship strikes, see below). One mass mortality event, associated with consumption of prey contaminated with biotoxins, has been reported from the east coast of the United States (Geraci et al. 1989), though these individuals stranded over a wide area and did not strand all at the same time. Ice entrapment occurs occasionally (Tomilin 1967; S. Todd, personal communication).

Anthropogenic Influences

Potentially negative interactions with humans fall under two broad categories. Some impacts may have acute (immediate) effects on individuals or a population, such as directed takes (whaling), entanglement in fishing gear, collisions with vessels, or exposure to acute pollutants (e.g., oil spills). Immunotoxic effects due to accumulation of persistent toxic chemicals may also have an acute impact by increasing susceptibility to diseases, thus causing an increase in mortality. Besides these acute impacts, there are a number of less tangible, longer-term, potentially negative human influences, including a reduction in reproductive rates due to immunotoxic effects, reduced prey availability due to human activities, and disturbance or displacement by vessel traffic or other sources of underwater sounds. While each of these is treated independently below, it should also be taken into account that cumulative impacts of all of these factors could be important (or in the case of longer-term stressors, synergistic interactions between impacts could occur; Whitehead et al. 2000).

Historically the primary source of direct mortality was commercial whaling, which was officially banned in the 1960s. However, there have been reports of substantial illegal whaling outside Canada since that date (Yablokov 1994; Mikhalev 1997). Whaling is strictly regulated by the International Whaling Commission, with only small numbers allowed to be killed for Aboriginal subsistence purposes. For populations that use Canadian waters, the only known hunting that currently occurs is up to two animals per year from the North Atlantic population, taken in St. Vincent. Reiner et al. (1996) report one animal taken off the Cape Verde Islands in 1988, and small numbers were taken in an Aboriginal fishery in West Greenland until 1985 (IWC 1986).

More animals are killed incidentally in fisheries each year than are killed through directed hunting (Volgenau et al. 1995; Mazzuca et al. 1998). Precise estimates of annual mortality for all Canadian populations from this source are not available. Off Newfoundland, estimates of annual mortality in fishing gear in the early 1990s ranged from 3 to 17, with a mean of 8 individuals (Volgenau et al. 1995), though previous entanglement (and mortality) rates, associated with the inshore cod (Gadus morhua) fishery, were higher (Lien 1994). Reported entanglements in fishing gear off Newfoundland from 2000 through 2002 ranged from 11-22 each year, and known mortalities ranged from only 0-5 per year (Ledwell et al. 2000; Ledwell and Huntington 2001, 2002). However, some animals were last seen towing gear and may have died as a result (see above citations). Types of fishing gear/activities associated with entanglements off Newfoundland in recent years include herring nets, crab pots, whelk pots, capelin traps, cod gillnets, and unspecified gillnets and ropes/buoys (Ledwell et al. 2000; Ledwell and Huntington 2001). Fishing effort in recent years in Newfoundland has shifted offshore (J. Lien, W. Ledwell, personal communications), and entanglements in offshore areas may not be reported as frequently as entanglements in inshore areas, thus the total number of animals entangled (and killed) each year is likely greater than suggested by the above references. In British Columbia there were only four records of entanglement in fishing gear reported between 1987 and 1995 (in prawn trap gear, salmon gillnet, and lines associated with traps), none of which were known to definitely result in mortality (Langelier et al. 1990; Guenther et al. 1995). In March 1999 a juvenile humpback was killed in British Columbia after entangling with a herring roe pond net (R. Palm, personal communication). Ship strikes are also a source of mortality and injury, and in some areas may be more important than mortality in fishing gear (Wiley et al. 1995; Barlow et al. 1998; Mazzuca et al. 1998; Laist et al. 2001).

In terms of indirect impacts, the biggest potential threat to humpbacks is probably disturbance at their primary breeding sites, since these areas are limited in size, and in the Pacific, at least, are relatively near-shore and thus perhaps more subject to human influences. Disturbance or disruption of behaviour might arise from whale watching (Bauer 1997), industrial activities (e.g., seismic exploration), or other high-intensity underwater sounds (e.g., oceanographic experiments or acoustic harassment devices used in aquaculture operations (Todd et al. 1996; Nitta 1997), or military sonars (Miller et al. 2000)). Seismic activity may affect humpback whales (perhaps by displacement, disturbance, or physiological damage), but specifics are unknown. Off eastern Canada, there is overlap between seismic activity and humpback whale habitat on the Grand Banks and Scotian Shelf. However, other important areas of humpback whale habitat are not currently subject to seismic activities: inshore waters off Newfoundland and Labrador, and southwest Nova Scotia.

Because of their rather low trophic level, accumulation of persistent anthropogenic toxins is not thought to be a threat to this species (though see O’Shea et al. 1999).

Reduction of their prey base is a matter for concern; one of the primary prey species in the North Pacific, Pacific herring (Clupea harengus) is currently considered a candidate for listing as “Threatened” under the U.S. Endangered Species Act. The principal prey of humpback whales in the western North Atlantic is capelin (Mallotus villosus). There is some controversy about the current status of capelin stocks off eastern Canada (DFO 2000; Rose and O’Driscoll 2002). Capelin seem to have moved south off Newfoundland as waters cooled in the 1990s. Some indices suggest that capelin abundance has been low since about 1990; other analyses suggest that there have been relatively strong capelin year classes since about 1992 (see above citations).

Special Significance of the Species

Their tendency to inhabit nearshore areas in many parts of the world, including Canadian waters, has brought humpbacks into regular contact with humans. They are probably the most acrobatic of the large whales, and in both eastern and western Canada humpbacks are the focus of a number of commercial whale watching operations. Perhaps the feature for which they are most well known is their long and complex “song” (Payne and McVay 1971), which has received considerable attention both from the scientific community and the general public.

Existing Legal Protection

Two factors are important in the legal protection of a species, the system that is in place to prohibit or regulate hunts or other threats, and how effective the system is, in terms of whether monitoring and enforcement is sufficient. Where information is available, each of these is discussed below.

International

Two international conventions are relevant to the protection of humpback whales, the Convention on International Trade in Endangered Species of Wild Fauna and Flora 1973 (CITES) and the International Convention for the Regulation of Whaling 1946 (administered by the International Whaling Commission - IWC).

All species of cetaceans are listed by CITES under one of two appendices. Appendix I includes species threatened with extinction (and which may be affected by trade), while Appendix II includes species which may become threatened with extinction unless trade is regulated, as well as species which must be subject to regulation in order that trade in threatened species of similar appearance may be controlled (Klinowska 1991). Humpback whales, because of the large numbers of animals taken in whaling operations and the subsequent population declines, are listed in Appendix I of CITES, thus all trade is banned between countries which are Parties to CITES. As of October 1998 there were 144 Parties to CITES (including Canada), leaving approximately 90 countries world-wide which were not members (CITES Secretariat statistics).

Whaling of humpback whales is regulated by the IWC. Commercial harvesting has been banned in the North Pacific since 1965 and in the North Atlantic since 1955. Forty-one animals were killed for scientific purposes off eastern Canada between 1966 and 1971 (Mitchell 1973), and Greenland was permitted to kill 10 per year up until 1985 for Aboriginal subsistence purposes (IWC 1986). The only takes that are currently permitted are small numbers of animals for Aboriginal subsistence purposes (St. Vincent and the Grenadines are authorized to catch two per year). Canada is not currently a member of the IWC, having withdrawn in 1982 (IWC 1982).

National

Canada: Within Canada, management of humpback whales falls under the responsibility of the Department of Fisheries and Oceans. Until 1993 humpbacks were covered under the "Cetacean Protection Regulations" (under the Fisheries Act of Canada of 1867). These regulations prohibited "hunting" without a licence. "Hunting" was defined as "to chase, shoot at, harpoon, take, kill, attempt to take or kill, or to harass cetaceans in any manner". No scheme, however, was in place to enforce such regulations, and Aboriginal hunting could be undertaken without a licence. In 1993, the federal government consolidated various marine mammal regulations, including the Cetacean Protection Regulations, under the new "Marine Mammal Regulations". These regulations stated that "no person should disturb a marine mammal except when under.... the authorities of these regulations", with "marine mammal" defined as all species listed under a particular appendix. Currently, hunting of humpback whales could occur if a "Fishing Licence" was obtained (except for Aboriginals, who could hunt without a licence), and fees for such licences are low ($5). However, no such licences have been issued, and issuance is at the discretion of the federal Minister of Fisheries and Oceans. It is unlikely any would be issued in Canada due to widespread public interest in these animals.

The 1997 Oceans Act provides for the establishment of marine protected areas (MPAs) in federal waters. One of the specific justifications listed for establishing MPAs is to conserve and protect marine mammals and their habitats. However, as with other federal legislation regarding marine mammals, establishment of marine protected areas and exclusion of activities which might jeopardize humpback whales or other marine mammals is at the discretion of the Minister of Fisheries and Oceans, rather than mandated. Regardless, there are general concerns about the efficacy of using MPAs to “protect” cetaceans (Phillips 1996; Whitehead et al. 2000), due primarily to the large range of most species and the lack of boundaries in the marine environment. Whitehead et al. (2000) note that most marine protected areas have provided little or no change in the level of threats faced by cetaceans in an area.

Other Countries

Given that humpback whales that utilize Canadian waters also move between a number of different countries (potentially including at least Norway, Iceland, Greenland, Ireland, the United Kingdom, the United States, Cuba, Bermuda, the Dominican Republic, Puerto Rico, the Virgin Islands, St. Vincent, Mexico, a number of Central American countries, the Philippines and Japan), protective measures in these countries are directly relevant to the conservation of the species in Canada. In the United States, interactions between humans and humpback whales are managed though the Marine Mammal Protection Act of 1972, the Endangered Species Act of 1973, the Marine Protection, Research and Sanctuaries Act of 1974 (through their presence in the Hawaiian Islands Humpback Whale National Marine Sanctuary as well as the Stellwagen Bank National Marine Sanctuary and several others), as well as a variety of other federal and state (in the State of Hawaii) regulations.

Technical Summary: North Pacific population

Megaptera novaeangliae
Humpback whale Rorqual à bosse
North Pacific population Population du Pacifique nord
Range of Occurrence in Canada: Pacific Ocean

Extent and Area information

· extent of occurrence (EO)(km²) >20,000 km2
· specify trend (decline, stable, increasing, unknown) Stable
· are there extreme fluctuations in EO (> 1 order of magnitude)? No
· area of occupancy (AO) (km²) > 20,000 km2
· specify trend (decline, stable, increasing, unknown)
Decline from pre-whaling.
Currently stable or increasing.
· are there extreme fluctuations in AO (> 1 order magnitude)? No
· number of extant locations N/A
· specify trend in # locations (decline, stable, increasing, unknown) N/A
· are there extreme fluctuations in # locations (>1 order of magnitude)?
· habitat trend: specify declining, stable, increasing or unknown trend in area, extent or N/Aquality of habitat Stable

Population information

· generation time (average age of parents in the population) (indicate years, months, days, etc.) ~20 years
· number of mature individuals (capable of reproduction) in the Canadian population (or, specify a range of plausible values) 500-1,000
· total population trend: specify declining, stable, increasing or unknown trend in number of mature individuals Past decline, current increase
· if decline, % decline over the last/next 10 years or 3 generations, whichever is greater (or specify if for shorter time period) >50% over 60 years
· are there extreme fluctuations in number of mature individuals (> 1 order of magnitude)? No
· is the total population severely fragmented (most individuals found within small and relatively isolated (geographically or otherwise) populations between which there is little exchange, i.e., < 1 successful migrant / year)? Some separation between 2 sub-populations.
· list each population and the number of mature individuals in each
Southern BC: ~100
Northern BC: 500+
· specify trend in number of populations (decline, stable, increasing, unknown) Unknown
· are there extreme fluctuations in number of populations (>1 order of magnitude)? No

Threats

Entrapment in fishing gear, ship strikes.

Rescue Effect (immigration from an outside source) Moderate

· does species exist elsewhere (in Canada or outside)? Yes
· status of the outside population(s)? Increasing
· is immigration known or possible? Yes
· would immigrants be adapted to survive here? Unknown
· is there sufficient habitat for immigrants here? Unknown

Quantitative Analysis Not undertaken

Technical summary: Western North Atlantic population

Megaptera novaeangliae
Humpback whale Rorqual à bosse
Western North Atlantic population Population de l’Atlantique nord-ouest
Range of Occurrence in Canada: Atlantic Ocean

Extent and Area information

· extent of occurrence (EO)(km²) Throughout the North Atlantic Ocean; >20,000 km2
· specify trend (decline, stable, increasing, unknown) Stable
· are there extreme fluctuations in EO (> 1 order of magnitude)? No
· area of occupancy (AO) (km²) > 20,000 km2
· specify trend (decline, stable, increasing, unknown) Stable
· are there extreme fluctuations in AO (> 1 order magnitude)? No
· number of extant locations N/A
· specify trend in # locations (decline, stable, increasing, unknown) N/A
· are there extreme fluctuations in # locations (>1 order of magnitude)? N/A
· habitat trend: specify declining, stable, increasing or unknown trend in area, extent or quality of habitat Concern over habitat degradation in breeding grounds in Caribbean.

Population information

· generation time (average age of parents in the population) (indicate years, months, days, etc.) ~20 years
· number of mature individuals (capable of reproduction) in the Canadian population (or, specify a range of plausible values) Total number, all ages: >2500. Mature individuals likely <1500
· total population trend: specify declining, stable, increasing or unknown trend in number of mature individuals Increasing
· if decline, % decline over the last/next 10 years or 3 generations, whichever is greater (or specify if for shorter time period) N/A
· are there extreme fluctuations in number of mature individuals (> 1 order of magnitude)? No
· is the total population severely fragmented (most individuals found within small and relatively isolated (geographically or otherwise) populations between which there is little exchange, i.e., < 1 successful migrant / year)? No
· list each population and the number of mature individuals in each N/A
· specify trend in number of populations (decline, stable, increasing, unknown) N/A
· are there extreme fluctuations in number of populations (>1 order of magnitude)? N/A

Threats

Habitat degradation on breeding grounds in the Caribbean, entrapment in fishing gear, ship strikes

Rescue Effect (immigration from an outside source) Moderate

· does species exist elsewhere (in Canada or outside)? Yes
· status of the outside population(s)? Increasing
· is immigration known or possible? Yes
· would immigrants be adapted to survive here? Unknown
· is there sufficient habitat for immigrants here? Unknown

Quantitative Analysis Not undertaken

Acknowledgements

This report was undertaken under contract to the Committee on the Status of Endangered Wildlife in Canada, funded by the Canadian Wildlife Service, Environment Canada. Bob Campbell administered the contract. I would like to thank John Calambokidis, Sal Cerchio, Chris Clark, Graeme Ellis, John Ford, Christine Gabriele, Jon Lien, Joe Mobley, Per Palsboll, Jan Straley, and Hal Whitehead for providing references, unpublished information, and/or for helpful comments. Phil Clapham, Peter Corkeron, Sascha Hooker, Sean Todd, Hal Whitehead and several anonymous reviewers made a number of helpful suggestions on versions of the manuscript.

Literature Cited

Acevedo, A., and M.A. Smultea. 1995. First records of humpback whales including calves at Golfo Dulce and Isla del Coco, Costa Rica, suggesting geographical overlap of northern and southern hemisphere populations. Marine Mammal Science 11:554-560.

Allen, J.M., H.C. Rosenbaum, S.K. Katona, P.J. Clapham and D.K. Mattila. 1994. Regional and sexual differences in fluke pigmentation of humpback whales (Megaptera novaeangliae) from the North Atlantic Ocean. Canadian Journal of Zoology 72:274-279.

Anonymous. 2001. Report of the Scientific Committee, London 2001. International Whaling Commission, Cambridge, U.K.

Baker, C.S., and L.M. Herman. 1987. Alternative population estimates of humpback whales (Megaptera novaeangliae) in Hawaiian waters. Canadian Journal of Zoology 65:2818-2821.

Baker, C.S., and S.R. Palumbi. 1997. The genetic structure of whale populations: implications for management. Society for Marine Mammalogy Special Publication 3:117-146.

Baker, C.S., L.M. Herman, A. Perry, W.S. Lawton, J.M. Straley, A.A. Wolman, G.D. Kaufman, H.E. Winn, J.D. Hall, J.M. Reinke and J. Ostman. 1986. Migratory movement and population structure of humpback whales (Megaptera novaeangliae) in the central and eastern North Pacific. Marine Ecology Progress Series 41:103-114.

Baker, C.S., A. Perry, J.L. Bannister, M.T. Weinrich, R.B. Abernathy, J. Calambokidis, J. Lien, R.H. Lambertsen, J. Urban, O. Vasquez, P.J. Clapham, A. Alling, S.J. O’Brien and S.R. Palumbi. 1993. Abundant mitochondrial DNA variation and world-wide population structure in humpback whales. Proceedings of the National Academy of Sciences 90:8239-8343.

Barco, S.G., W.A. McLellan, J.M. Allen, R.A. Asmutis-Silvia, R. Mallon-Day, E.M. Meagher, D.A. Pabst, J. Robbinns, R.E. Seton, W.M. Swingle, M.T. Weinrich, and P.J. Clapham. 2002. Population identity of humpback whales (Megaptera novaeangliae) in the waters of the US mid-Atlantic sates. Journal of Cetacean Research and Management 4:135-141.

Barlow, J., and P.J. Clapham. 1997. A new birth-interval approach to estimating demographic parameters of humpback whales. Ecology 78:535-546.

Barlow, J., P.S. Hill, K.A. Forney and D.P. DeMaster. 1998. U.S. Pacific marine mammal stock assessments: 1998. NOAA Technical Memorandum SWFSC-258.

Baraff, L., and M.T. Weinrich. 1993. Separation of humpback whale mothers and calves on a feeding ground in early autumn. Marine Mammal Science 9:431-434.

Baraff, L.S., P.J. Clapham, D.K. Mattila and R.S. Bowman. 1991. Feeding behavior of a humpback whale in low-latitude waters. Marine Mammal Science 7:197-202.

Bauer, G.B. 1997. An overview of the effects of vessel approaches on humpback whale behavior in Hawaii. NOAA Technical Memorandum NMFS-OPR-11:68-73.

Breiwick, J.M., E. Mitchell and R.R. Reeves. 1983. Simulated population trajectories for northwest Atlantic humpback whales, 1865-1980. In Abstracts of the Fifth Biennial Conference on the Biology of Marine Mammals, Boston, MA.

Brown, M.R., P.J. Corkeron, P.T. Hale, K.W. Schultz and M.M. Bryden. 1995. Evidence of sex-segregated migration in the humpback whale (Megaptera novaeangliae). Proceedings of the Royal Society of London B 259:229-234.

Calambokidis, J., G.H. Steiger, J.R. Evenson, K.R. Flynn, K.C. Balcomb, D.E. Claridge, P. Bloedel, J.M. Straley, C.S. Baker, O. von Ziegesar, M.E. Dahlheim, J.M. Waite, J.D. Darling, G. Ellis, and G.A. Green. 1996. Interchange and isolation of humpback whales off California and other North Pacific feeding grounds. Marine Mammal Science 12:215-226.

Calambokidis, J., G.H. Steiger, J.M. Straley, T.J. Quinn, L.M. Herman, S. Cerchio, D.R. Salden, M. Yamaguchi, F. Sato, J. Urban, J. Jacobsen, O. von Ziegesar, K.C. Balcomb, C.M. Gabriele, M.E. Dahlheim, N. Higashi, S. Uchida, J.K.B. Ford, Y. Miyamura, P.L. de Guevara, S.A. Mizroch, L. Schlender, and K. Rasmussen. 1997. Abundance and population structure of humpback whales in the North Pacific basin. Final Report under Contract No. 5ABNF500113 to the Southwest Fisheries Science Center, National Marine Fisheries Service, La Jolla, CA.

Calambokidis, J., T. Chandler, K. Rasmussen, G.H. Steiger and L. Schlender. 1999. Humpback and blue whale photo-identification research off California, Oregon and Washington in 1998. Report prepared for Southwest Fisheries Science Center, Olympic Coast National Marine Sanctuary and University of California, Santa Cruz, CA.

Calambokidis, J., G.H. Steiger, K. Rasmussen, J. Urban R., K.C. Balcomb, P. L. de Guevara P., M. Salinas Z., J.K. Jacobsen, C.S. Baker, L.M. Herman, S. Cerchio and J.D. Darling. 2000. Migratory destinations of humpback whales that feed off California, Oregon and Washington. Marine Ecology Progress Series 192:295-304.

Calambokidis, J., G.H Steiger, J.M Straley, L.M. Herman, S. Cerchio, D.R. Salden, J. Urbán R., J.K. Jacobsen, O. von Ziegesar, K.C. Balcomb, C.M. Gabriele, M.E. Dahlheim, S. Uchida, G. Ellis, Y. Miyamura, P. Ladrón de Guevara P., M. Yamaguchi, F. Sato, S.A. Mizroch, L. Schlender, K. Rasmussen, J. Barlow and T.J. Quinn II. 2001. Movements and population structure of humpback whales in the North Pacific. Marine Mammal Science 17:769-794.

Campbell, R.R. 1996. Rare and endangered fish and marine mammals of Canada: COSEWIC fish and marine mammal subcommittee status reports: X. Canadian Field-Naturalist 110:454-461.

Cerchio, S. 1998. Estimates of humpback whale abundance off Kauai, 1989 to 1993: evaluating biases associated with sampling the Hawaiian Islands breeding assemblage. Marine Ecology Progress Series 175:23-34.

Chittleborough, R.G. 1958. The breeding cycle of the female humpback whale, Megaptera nodosa (Bonnaterre). Australian Journal of Marine and Freshwater Research 9:1-18.

Chittleborough, R.G. 1965. Dynamics of two populations of the humpback whale. Australian Journal of Marine and Freshwater Research 16:13-128.

Clapham, P.J. 1992. The attainment of sexual maturity in humpback whales. Canadian Journal of Zoology 70:1470-1472.

Clapham, P.J. 2000. The humpback whale: seasonal feeding and breeding in a baleen whale. In Cetacean societies: field studies in behavior. Edited by J. Mann, R. Connor, P. Tyack and H. Whitehead. University of Chicago Press, Chicago, IL.

Clapham, P.J., and C.A. Mayo. 1987. Reproduction and recruitment of individually identified humpback whales, Megaptera novaeangliae, observed in Massachusetts Bay, 1979-1985. Canadian Journal of Zoology 65:2853-2863.

Clapham, P.J., and C.A. Mayo. 1990. Reproduction of humpback whales (Megaptera novaeagliae) observed in the Gulf of Maine. Reports of the International Whaling Commission Special Issue 12:171-175.

Clapham, P.J., D.K. Mattila and P.J. Palsboll. 1993. High-latitude-are composition of humpback whale competitive groups in Samana Bay: further evidence for panmixis in the North Atlantic population. Canadian Journal of Zoology 71:1065-1066.

Clapham, P., J. Barlow, T. Cole, D. Mattila, R. Pace, D. Palka, and J. Robbins. 2001. Stock definition, abundance and demographic parameters of humpback whales from the Gulf of Maine. International Whaling Commission Document SC/53/NAH10.

Corkeron, P.J., and R.C. Connor. 2000. Why do baleen whales migrate? Marine Mammal Science 15:1228-1245.

Craig, A.S., and L.M. Herman. 1997. Sex differences in site fidelity and migration of humpback whales (Megaptera novaeangliae) to the Hawaiian islands. Canadian Journal of Zoology 75:1923-1933.

Darling, J.D., and S. Cerchio. 1993. Movement of a humpback whale (Megaptera novaeangliae) between Japan and Hawaii. Marine Mammal Science 9:84-89.

Darling, J.D. and C.M. Jurasz. 1983. Migratory destinations of North Pacific humpback whales (Megaptera novaeangliae). In Communication and behavior of whales. Edited by R. Payne. AAAS Selected Symposium 76. Washington, DC.

Darling, J.D., and H. Morowitz. 1986. Census of “Hawaiian” humpback whales (Megaptera novaeangliae) by individual identification. Canadian Journal of Zoology 64:105-111.

Darling, J.D., J. Calambokidis, K.C. Balcomb, P. Bloedel, K. Flynn, A. Mochizuki, K. Mori, F. Sato, H. Suganuma and M. Yamaguchi. 1996. Movement of a humpback whale (Megaptera novaeanagliae) from Japan to British Columbia and return. Marine Mammal Science 12:281-287.

DFO. 2000. Capelin in Subarea2 + Div. 3KL Update. DFO Science Stock Status Report B2-02 (2001).

Forez-Gonzalez, L., J.J. Capella and H.C. Rosenbaum. 1994. Attack of killer whales (Orcinus orca) on humpback whales (Megaptera novaeangliae) on a South American Pacific breeding ground. Marine Mammal Science 10:218-222.

Florez-Gonzalez, L., J. Capella, B. Haase, G.A. Bravo, F. Felix and T. Gerrodette. 1998. Changes in winter destinations and the northernmost record of southeastern Pacific humpback whales. Marine Mammal Science 14:189-196.

Ford, J.K.B., K. Heise, L.G. Barrett-Lennard and G.M. Ellis. 1994. Killer whales and other cetaceans of the Queen Charlotte Islands/Haida Gwaii. Report to the Gwaii Haanas National Park Reserve, Parks Canada.

Gabriele, C.M. 1992. Population characteristics of humpback whales in Glacier Bay and adjacent waters: 1992. Report to the National Park Service, Glacier Bay National Park and Preserve, Gustavus, AK.

Gabriele, C.M., J.M. Straley, L.M. Herman and R.J. Coleman. 1996. Fastest documented migration of a North Pacific humpback whale. Marine Mammal Science 12:457-464.

Gambell, R. 1975. World whale stocks. Mammal Review 6:41-53.

Gendron, D., and J. Urban. 1993. Evidence of feeding by humpback whales (Megaptera novaeangliae) in the Baja California breeding ground, Mexico. Marine Mammal Science 9:76-81.

Geraci, J.R., D.M. Anderson, R.J. Timperi, D.J. St. Aubin, G.A. Early, J.H. Prescott and C.A. Mayo. 1989. Humpback whale (Megaptera novaeangliae) fatally poisoned by dinoflagellate toxin. Canadian Journal of Fisheries and Aquatic Sciences 46:1195-1198.

Glockner, D.A. 1983. Determining the sex of humpback whales, Megaptera novaeangliae, in their natural environment. In Communication and behavior of whales. Edited by R. Payne. AAAS Selected Symposium 76. Washington, DC.

Glockner-Ferrari, D.A., and M.J. Ferrari. 1997. Overview of the reproductive parameters of the North Pacific humpback whale based on long-term studies. NOAA Technical Memorandum NMFS-OPR-11:15-28.

Gregr, E.J., L. Nichol, J.K.B. Ford, G. Ellis, and A.W. Trites. 2000. Migration and population structure of northeastern Pacific whales off coastal British Columbia: An analysis of commercial whaling records from 1908-1967. Marine Mammal Science 16:699-727.

Guenther, T.J., R.W. Baird, R.L. Bates, P.M. Willis, R.L. Hahn and S.G. Wischniowski. 1995. Strandings and fishing gear entanglements of cetaceans off the west coast of Canada in 1994. International Whaling Commission Document SC/47/O6.

Hay, K.A. 1985. Status of the humpback whale, Megaptera novaeangliae, in Canada. Canadian Field-Naturalist 99:425-432.

Hain, J.H.W., S.L. Ellis, R.D. Kenney, P.J. Clapham, B.K. Gray, M.T. Weinrich and I.G. Babb. 1995. Apparent bottom feeding by humpback whales on Stellwagen Bank. Marine Mammal Science 11:464-479.

Hazevoet, C.J., and F.W. Wenzel. 2000. Whales and dolphins (Mammalia, Cetacea) of the Cape Verde Islands, with special reference to the humpback whale Megaptera novaeangliae (Borowski, 1781). Contributions to Zoology 69:197-211.

Helweg, D.A., L.M. Herman, S. Yamamoto and P.H. Forestell. 1990. Comparison of songs of humpback whales (Megaptera novaeangliae) recorded in Japan, Hawaii and Mexico during the winter of 1989. Scientific Reports of the Cetacean Research Institute 1:1-20.

Hill, P.S., and D.P. DeMaster. 1998. Alaska marine mammal stock assessments, 1998. NOAA Technical Memorandum NMFS-AFSC-97.

Hoyt, E. 1983. Great winged whales: combat and courtship rites among humpbacks, the ocean’s not-so-gentle giants. Equinox 10:25-47.

IUCN. 1996. 1996 IUCN red list of threatened animals. IUCN, Gland, Switzerland.

IWC. 1982. International Whaling Commission Report 1980-1981. Report of the International Whaling Commission 32:3-7.

IWC. 1986. Report of the sub-committee on protected species and aboriginal subsistence whaling. Report of the International Whaling Commission 36:95-111.

Jefferson, T.A., P.J. Stacey and R.W. Baird. 1991. A review of killer whale interactions with other marine mammals: predation to co-existence. Mammal Review 21:151-180.

Johnson, J.H., and A.A. Wolman. 1984. The humpback whale, Megaptera novaeangliae. Marine Fisheries Review 46:30-37.

Jurasz, C.M., and V.P. Jurasz. 1979. Feeding modes of the humpback whale, Megaptera novaeangliae, in southeast Alaska. Scientific Reports of the Whales Research Institute 31:69-83.

Katona, S.K., and J.A. Beard. 1990. Population size, migration and substock structure of the humpback whale (Megaptera novaeangliae) in the western North Atlantic Ocean. Report of the International Whaling Commission Special Issue 12:295-306.

Katona, S., P.M. Harcourt, J.S. Perkins and S. Kraus. 1980. Humpback whales in the western North Atlantic: a catalogue of identified individuals. Second Edition, College of the Atlantic, Bar Harbor, ME.

Katona, S., B. Baxter, O. Brazier, S. Kraus, J. Perkins and H. Whitehead. 1979. Identification of humpback whales by fluke photographs. In Behavior of marine animals, Volume 3: cetaceans. Edited by H.E. Winn and B.L. Olla. Plenum Press, New York, NY.

Klinowska, M. 1991. Dolphins, porpoises and whales of the world - the IUCN Red Data Book. IUCN, Gland, Switzerland.

Laerm, J., F. Wenzel, J.E. Craddock, D. Weinand, J. McGurk, M.J. Harris, G.A. Early, J.G. Mead, C.W. Potter and N.B. Barros. 1997. New prey species for northwestern Atlantic humpback whales. Marine Mammal Science 13:705-711.

Laist, D.W., A.R. Knowlton, J.G. Mead, A.S. Collet and M. Podesta. 2001. Collisions between ships and whales. Marine Mammal Science 17:35-75.

Langelier, K.M., P.J. Stacey and R.W. Baird. 1990. Stranded whale and dolphin program of B.C. – 1989 report. Wildlife Veterinary Report 3(1):10-11.

Larsen, A.H., J. Sigurjonsson, N. Oien, G. Vikingsson and P.J. Palsboll. 1996. Population genetic analysis of nuclear and mitochondrial loci in skin biopsies collected from central and northeastern North Atlantic humpback whales (Megaptera novaeangliae): population identity and migratory destinations. Proceedings of the Royal Society of London B 263:1611-1618.

Ledwell, W., J. Huntington, and J. Lien. 2000. Whale entrapments in fishing gear and other marine animal incidental entrapments during 2000. A report to the Habitat Stewardship Program of Environment Canada.

Ledwell, W., and J. Huntington. 2001. Whale entrapments in fishing gear, strandings and sightings of marine animals and summary of the entrapment program during 2001 – Newfoundland Region. A report to the Habitat Stewardship Program of Environment Canada.

Ledwell, W., and J. Huntington. 2002. Whale entrapments in fishing gear and a summary marine animal disentanglement assistance program in Newfoundland and Labrador during 2002. A report to the Habitat Stewardship Program of Environment Canada.

Lien, J. 1994. Entrapments of large cetaceans in passive inshore fishing gear in Newfoundland and Labrador (1979-1990). Report of the International Whaling Commission Special Issue 15:149-157.

Martin, A.R., S.K. Katona, D. Mattila, D. Hembree and T.D. Waters. 1984. Migration of humpback whales between the Caribbean and Iceland. Journal of Mammalogy 65:330-333.

Mate, B.R., R. Gisiner and J. Mobley. 1998. Local and migratory movements of Hawaiian humpback whales tracked by satellite telemetry. Canadian Journal of Zoology 76:863-868.

Mazzuca, L., S. Atkinson and E. Nitta. 1998. Deaths and entanglements of humpback whales, Megaptera novaeangliae, in the main Hawaiian islands, 1972-1996. Pacific Science 52:1-13.

Merilees, B. 1985. Humpbacks in our Strait. Waters (Vancouver Public Aquarium) 8:7-24.

Mikhalev, Y.A. 1997. Humpback whales Megaptera novaeangliae in the Arabian Sea. Marine Ecology Progress Series 149:13-21.

Miller, P.J.O., N. Biassoni, A. Samuels and P.L. Tyack. 2000. Whale songs lengthen in response to sonar. Nature 405:903.

Mitchell, E. 1973. Draft report on humpback whales taken under special scientific permit by eastern Canadian land stations, 1969-1971. Report of the International Whaling Commission 23:183-154.

Mobley, J.R., G.B. Bauer and L.M. Herman. 1999a. Changes over a ten-year interval in the distribution and relative abundance of humpback whales (Megaptera novaeangliae) wintering in Hawaiian waters. Aquatic Mammals 25:63-72.

Mobley, J.R., R.A. Grotefendt, P.H. Forestell, S. Spitz, E. Brown, G.B. Bauer and A.S. Frankel. 1999b. Population estimate for “Hawaiian” humpback whales: results of 1993-98 aerial surveys. In Abstracts of the 13th Biennial Conference on the Biology of Marine Mammals, Wailea, HI.

Naessig, P.J. 1999. Levels and origin of predatory scarring on humpback whales in east Australian waters. In Abstracts of the 13th Biennial Conference on the Biology of Marine Mammals, Wailea, HI.

Nitta, E.T. 1997. An overview of the responsibilities and activities of the National Marine Fisheries Service – Section 7 consultations, approach regulations and research. NOAA Technical Memorandum NMFS-OPR-11:74-80.

Norris, T.F., M. McDonald and J. Barlow. 1999. Acoustic detection of singing humpback whales (Megaptera novaeangliae) in the eastern North Pacific during their northbound migration. Journal of the Acoustical Society of America 106:506-514.

O’Shea, T.J., R.R. Reeves and A.K. Long. 1999. Marine mammals and persistent ocean contaminants: proceedings of the Marine Mammal Commission workshop, Keystone, Colorado, 12-15 October 1998. Marine Mammal Commission, Bethesda, Maryland.

Palka, D. 2000. Abundance of the Gulf of Maine/Bay of Fundy harbor porpoise based on shipboard and aerial surveys during 1999. NOAA/NMFS/NEFSC Ref. Doc. 00-07.

Palsboll, P.J., P.J. Clapham, D.K. Mattila, F. Larsen, R. Sears, H.R. Siegismund, J. Sigurjonsson, O. Vasquez and P. Arctander. 1995. Distribution of mtDNA haplotypes in North Atlantic humpback whales: the influence of behaviour on population structure. Marine Ecology Progress Series 116:1-10.

Paterson, R., P. Paterson and D.H. Cato. 1994. The status of humpback whales Megaptera novaeangliae in east Australia thirty years after whaling. Biological Conservation 70:135-142.

Payne, R., and S. McVay. 1971. Songs of humpback whales. Science 173:585-597.

Payne, R., and L.N. Guinee. 1983. Humpback whale (Megaptera novaeangliae) songs as an indicator of “stocks”. In Communication and behavior of whales. Edited by R. Payne. AAAS Selected Symposium 76. Washington, DC.

Phillips, C. 1996. Conservation in practice: agreements, regulations, sanctuaries and action plans. In The conservation of whales and dolphins. Edited by M.P. Simmonds and J.D. Hutchinson. John Wiley and Sons, Ltd. Chichester, UK.

Pike, G.C. 1953. Colour pattern of the humpback whales from the coast of British Columbia. Journal of the Fisheries Research Board of Canada 171:1-54.

Pike, G.C., and I.B. MacAskie. 1969. Marine mammals of British Columbia. Fisheries Research Board of Canada Bulletin 171.

Reeves, R., and E. Mitchell. 1982. How many humpacks – then and now? The log of Mystic Seaport 34:71-83.

Reeves, R.R., P.J. Clapham, and S.E. Wetmore. 2002. Humpback whale (Megaptera novaeangliae) occurrence near the Cape Verde Islands, based on American 19th century whaling records. Journal of Cetacean Research and Management 4:235-253.

Reiner, F., M.E. dos Santos and F.W. Wenzel. 1996. Cetaceans of the Cape Verde archipelago. Marine Mammal Science 12:434-443.

Rice, D.W. 1978. The humpback whale in the North Pacific: distribution, exploitation, and numbers. In Report on a workshop on problems related to humpback whales (Megaptera novaeangliae) in Hawaii. Edited by K.S. Norris and R.R. Reeves. U.S. Marine Mammal Commission Report 77/03.

Robbins, J., M. Bérubé, P.J. Clapham, N. Friday, and D. Mattila. 2001. Structure and regional composition of the Gulf of Maine humpback whale (Megaptera novaeangliae) population. International Whaling Commission Document SC/53/NAH12.

Rose, G.A., and R.L. O’Driscoll. 2002. Capelin are good for cod: can the northern stock rebuild without them? ICES Journal of Marine Science 59:1018-1026.

Rosenbaum, H.C., P.J. Clapham, J. Allen, M. Nicole-Jenner, C. Jenner, L. Florez Gonzalez, J. Urban R., P. Ladron G., K. Mori, M. Yamaguchi and C.S. Baker. 1995. Geographic variation in ventral fluke pigmentation of humpback whale Megaptera novaeangliae populations worldwide. Marine Ecology Progress Series 124:1-7.

Salden, D.R. 1989. An observation of apparent feeding by a sub-adult humpback whale off Maui, Hawaii. In Abstracts of the Eighth Biennial Conference on the Biology of Marine mammals, Pacific Grove, CA.

Salden, D.R., L.M. Herman, M. Yamaguchi and F. Sato. 1999. Multiple visits of individual humpback whales (Megaptera novaeangliae) between the Hawaiian and Japanese winter grounds. Canadian Journal of Zoology 77:504-508.

Smith, T.D., J. Allen, P.J. Clapham, P.S. Hammond, S. Katona, F. Larsen, J. Lien, D. Mattila, P.J. Palsboll, J. Sigurjonsson, P.T. Stevick, and N. Oien. 1999. An ocean-basin-wide mark-recapture study of the north Atlantic humpback whale (Megaptera novaeangliae). Marine Mammal Science 15:1-32.

Steiger, G.H., J. Calambokidis, R. Sears, K.C. Balcomb and J.C. Cubbage. 1991. Movements of humpback whales between California and Costa Rica. Marine Mammal Science 7:306-310.

Stevick, P.T. 1999. Age-length relationships in humpback whales: a comparison of strandings in the western North Atlantic with commercial catches. Marine Mammal Science 15:725-737.

Stevick, P.T., N. Oien and D.K. Mattila. 1998. Migration of a humpback whale (Megaptera novaeangliae) between Norway and the West Indies. Marine Mammal Science 14:162-166.

Stevick, P.T., J. Allen, P.J. Clapham, N. Friday, S.K. Katona, F. Larsen, J. Lien, D.K. Mattila, P.J. Palsbøll, J. Sigurjonsson, T.D. Smith, N. Øien, and P.S. Hammond. 2001. Trends in abundance of North Atlantic humpback whales, 1979-1993. International Whaling Commission Document SC/53/NAH2.

Stevick, P.T., B.J. McConnell, and P.S. Hammond. 2002. Patterns of movement. In Marine mammal biology; an evolutionary approach. Edited by A.R. Hoelzel. Blackwell, Oxford, UK.

Stevick, P.T., J. Allen, M. Berube, P.J. Clapham, S.K. Katona, F. Larsen, J. Lien, D.K. Mattila, P.J. Palsboll, J. Robbins, J. Sigurjonsson, T.D. Smith, N. Oien, and P.S. Hammond. 2003. Segregation of migration by feeding ground origin in North Atlantic humpback whales (Megaptera novaeangliae). Journal of Zoology, London 259:231-237.

Stone, G.S., L. Florez-Gonzalez and S. Katona. 1990. Whale migration record. Nature 346:705.

Straley, J.M. 1990. Fall and winter occurrence of humpback whales (Megaptera novaeangliae) in southeastern Alaska. Reports of the International Whaling Commission Special Issue 12:319-323.

Straley, J.M., C.M. Gabriele and C.S. Baker. 1993. Seasonal characteristics of humpback whales (Megaptera novaeangliae) in southeastern Alaska. In Proceedings of the Third Glacier Bay Science Symposium. National Park Service, Anchorage, AK.

Straley, J.M., C.M. Gabriele and C.S. Baker. 1994. Annual reproduction by individually identified humpback whales (Megaptera novaeangliae) in Alaskan waters. Marine Mammal Science 10:87-92.

Swingle, W.M., S.G. Barco, T.D. Pitchford, W.A. McLellan and D.A. Pabst. 1993. Appearance of juvenile humpback whales feeding in the nearshore waters of Virginia. Marine Mammal Science 9:309-315.

Todd, S., P. Stevick, J. Lien, F. Marques and D. Ketten. 1996. Behavioural effects of exposure to underwater explosions in humpback whales (Megaptera novaeangliae). Canadian Journal of Zoology 74:1661-1672.

Tomilin, A.G. 1967. Mammals of the USSR and adjacent countries. Cetacea, Volume 9. Translated from Russian by the Isreal Program for Scientific Translations, Jerusalem.

Tonnessen, J.N., and A.O. Johnsen. 1982. The history of modern whaling. University of California Press, Berkeley and Los Angeles, CA.

Urbán R., J., A. Jaramillo L., A Aguayo L., P. Ladrón de Guevara P., M. Salinas Z., C. Alvarez F., L. Medrano G., J.K. Jacobsen, K.C. Balcomb, D.E. Claridge, J. Calambokidis, G.H. Steiger, J.M. Straley, O. von Ziegesar, J.M. Waite, S. Mizroch, M.E. Dahlheim, J.D. Darling and C.S. Baker. 2000. Migratory destinations of humpback whales wintering in the Mexican Pacific. Journal of Cetacean Research and Management 2:101-110.

Volgenau, L., S.D. Kraus and J. Lien. 1995. The impact of entanglements on two substocks of the western North Atlantic humpback whale, Megaptera novaeangliae. Canadian Journal of Zoology 73:1689-1698.

Waite, J.M., M.E. Dahlheim, R.C. Hobbs, S.A. Mizroch, O. von Ziegesar-Matkin, J.M. Straley, L.M. Herman and J. Jacobsen. 1999. Evidence of a feeding aggregation of humpback whales (Megaptera novaeangliae) around Kodiak Island, Alaska. Marine Mammal Science 15:210-220.

Waring, G.T., J.M. Quintal and S.L. Swartz. 2000. U.S. Atlantic and Gulf of Mexico marine mammal stock assessments – 2000. NOAA Technical Memorandum NMFS-NE-162.

Webb, R.L. 1988. On the Northwest: Commercial whaling in the Pacific Northwest 1790-1967. UBC Press, Vancouver, BC.

Weinrich, M.T., J. Bove and N. Miller. 1993. Return and survival of humpback whale (Megaptera novaeangliae) calves born to a single female in three consecutive years. Marine Mammal Science 9:325-328.

Whitehead, H. 1982. Populations of humpback whales in the Northwest Atlantic. Report of the International Whaling Commission 32:345-353.

Whitehead, H. 1985. Humpback whale songs from the North Indian Ocean. Investigations on Cetacea 17:157-162.

Whitehead, H. 1987. Updated status of the humpback whale, Megaptera novaeangliae, in Canada. Canadian Field-Naturalist 101:284-294.

Whitehead, H., R.R. Reeves and P.L. Tyack. 2000. Science and the conservation, protection and management of wild cetaceans. In Cetacean societies: fields studies of dolphins and whales. Edited by J. Mann, R. Connor, P. Tyack and H. Whitehead. University of Chicago Press, Chicago, IL.

Wiley, D.N., R.A. Asmutis, T.D. Pitchford and D.P. Gannon. 1995. Stranding and mortality of humpback whales, Megaptera novaeangliae, in the mid-Atlantic and southeast United States, 1985-1992. Fishery Bulletin, U.S. 93:196-205.

Winn, H.E., R.K. Edel and A.G. Taruski. 1975. Population estimate of the humpback whale (Megaptera novaeangliae) in the West Indies by visual and acoustic techniques. Journal of the Fisheries Research Board of Canada 32:499-506.

Yablokov, A.V. 1994. Validity of whaling data. Nature 367:108.

The Author

Robin W. Baird received his Ph.D. in Biology from Simon Fraser University in 1994, with his thesis focusing on the foraging behaviour and ecology of killer whales in southern British Columbia. He has been undertaking research on marine mammals since 1986, including three seasons of work on whales off Nova Scotia. He has studied the diving behaviour of humpback whales in Hawaiian waters for two years. Dr. Baird is currently a Post-doctoral Fellow at Dalhousie University, and works for the U.S. National Marine Fisheries Service in Beaufort, North Carolina.

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