Hunting during the last 200 years reduced many populations of mysticete whales to near extinction. To evaluate potential genetic bottlenecks in these exploited populations, we examined mitochondrial DNA control region sequences from 90 individual humpback whales (Megaptera novaeangliae) representing six subpopulations in three ocean basins. Comparisons of relative nucleotide and nucleotype diversity reveal an abundance of genetic variation in all but one of the oceanic subpopulations. Phylogenetic reconstruction of nucleotypes and analysis of maternal gene flow show that current genetic variation is not due to postexploitation migration between oceans but is a relic of past population variability. Calibration of the rate of control region evolution across three families of whales suggests that existing humpback whale lineages are of ancient origin. Preservation of preexploitation variation in humpback whales may be attributed to their long life-span and overlapping generations and to an effective, though perhaps not timely, international prohibition against hunting.Humpback whales (Megaptera novaeangliae) once numbered >125,000 individuals distributed into three oceanic populations: the North Pacific, the North Atlantic, and the southern oceans. Within each population, observations of migratory movement by marked individuals suggest that humpback whales form relatively discrete subpopulations that are not separated by obvious geographic barriers (1). Before protection by international agreement in 1966, the world-wide population of humpback whales had been reduced by hunting to <5000, with some regional subpopulations reduced to <200 individuals (Table 1).To evaluate the possibility that commercial hunting reduced genetic variation in baleen whales, we examined nucleotide sequence variation in the mitochondrial (mt) DNA from 90 humpback whales collected from the three major oceanic basins. We chose humpback whales for this evaluation because their well-described subpopulation divisions and well-documented history of exploitation provide a historical framework within which to evaluate genetic data (Table 1). We chose mtDNA as a genetic marker because of its power in describing the genetic structure of maternal lineages within populations and its sensitivity to demographic changes in populations (20). To allow the use of small skin samples collected by biopsy darting, we applied the polymerase chain reaction (PCR) and direct "solid-phase" sequencing methodology (21) to the mtDNA control region or "D-loop," a noncoding region that is highly variable in most vertebrates (22).The publication costs of this article were defrayed in part by page charge payment. This article must therefore be hereby marked "advertisement" in accordance with 18 U.S.C. §1734 solely to indicate this fact.We first verified that oceanic populations of humpback whales are independent demographic units by estimating mtDNA gene flow with a cladistic analysis of the control region sequences. We then evaluated mtDNA diversity within each o...

Photographs of individually identified humpback whales Megaptera novaeangliae were collected in regions throughout the central and eastern North Pacific during the years 1977 to 1983. A comparison of these photographs revealed extensive movement between seasonal habitats. Whales found wintering near Hawaii traveled to summer feeding regions throughout the coastal waters of Alaska. Whales wintering near Mexico were found in Alaskan feeding regions and near the Farallon Islands off central California. I-lttle exchange was found between the 2 wintering grounds or among the 5 summering grounds studied. Fidelity to a given feeding region was demonstrated by a high proportion of migratory return. Evidence of fidelity to a given wintering ground was less conclusive. The coloration of humpback whale flukes showed a longitudinal cline across the 5 feeding regions. Flukes of whales from the easternmost feeding regions were, on average, darker than those from the westernmost feeding regions. Whales in Hawaii and Mexico were similar in fluke coloration and the average coloration on both wintering grounds was intermediate between the extremes of the feeding regions. We propose that humpback whales in the eastern and central North Pacific form a single 'structured stock' consisting of several geographically-isolated 'feeding herds' which intermingle on 1 or more wintering grounds. Mark-recapture analyses of resightlng data indicate that the Hawaiian wintering congregation is 4 to 6 times larger than the southeastern Alaska feeding herd. Within a structured stock, sets of whales interact with different probabilities in each seasonal habitat. This, in turn, has important implications for the social organization and management of these whales.

The population structure of variation in a nuclear actin intron and the control region of mitochondrial DNA is described for humpback whales from eight regions in the North Pacific Ocean: central California, Baja Peninsula, nearshore Mexico (Bahia Banderas), offshore Mexico (Socorro Island), southeastern Alaska, central Alaska (Prince Williams Sound), Hawaii and Japan (Ogasawara Islands). Primary mtDNA haplotypes and intron alleles were identified using selected restriction fragment length polymorphisms of target sequences amplified by the polymerase chain reaction (PCR-RFLP). There was little evidence of heterogeneity in the frequencies of mtDNA haplotypes or actin intron alleles due to the year or sex composition of the sample. However, frequencies of four mtDNA haplotypes showed marked regional differences in their distributions (phi ST = 0.277; P < 0.001; n = 205 individuals) while the two alleles showed significant, but less marked, regional differences (phi ST = 0.033; P < 0.013; n = 400 chromosomes). An hierarchical analysis of variance in frequencies of haplotypes and alleles supported the grouping of six regions into a central and eastern stock with further partitioning of variance among regions within stocks for haplotypes but not for alleles. Based on available genetic and demographic evidence, the southeastern Alaska and central California feeding grounds were selected for additional analyses of nuclear differentiation using allelic variation at four microsatellite loci. All four loci showed significant differences in allele frequencies (overall FST = 0.043; P < 0.001; average n = 139 chromosomes per locus), indicating at least partial reproductive isolation between the two regions as well as the segregation of mtDNA lineages. Although the two feeding grounds were not panmictic for nuclear or mitochondrial loci, estimates of long-term migration rates suggested that male-mediated gene flow was several-fold greater than female gene flow. These results include and extend the range and sample size of previously published work, providing additional evidence for the significance of genetic management units within oceanic populations of humpback whales.

Reproductive histories of indnldually identified, female humpback whales were documented on both the summering and wintering grounds of an endangered but currently unexploited population. Interbirth or 'calving' intervals of mature females were on average longer and more variable than previously reported, ranging from 1 to at least 5 yr. In Hawaii, multiple sightings of 18 females provided an estimated calving rate (calves [mature female]-' yr-') of 0.58. In southeastern Alaska, multiple sightings of 4 1 females provided an estimated calving rate of 0.37 The survival of an individual through at least its first year of life was documented in 5 cases. Three of these, first identified as calves in southeastern Alaska, continued to return to this feeding region as juveniles. The possible weaning of a year-old whale was observed in Hawaii, and the apparent death of a calf was documented in southeastern Alaska. We suggest that the estimated calving rate from sightings of females in Hawaii is inflated by sighting biases and that the lower estimate from southeastern Alaska is a better measure of current reproductive rates. A comparison of this estimate with historical estimates of pregnancy rates from whaling records provides no evidence of a marked density-dependent increase in the reproductive rate of humpback whales.

A total of 326 humpback whales (Megaptera novaeangliae) were individually identified in southeastern Alaska during five summer seasons (July to September) and four late seasons (November to February) spanning the years 1979 to 1983. Peak numbers of whales were found late in August or early in September. Whales arrived 1–2 wk later in 1982 than in 1981. Whales sighted in both the summer and late seasons of 1981 and 1982 remained about 3.7 mo and one whale remained for at least 4.9 mo. Humpback whales from southeastern Alaska wintered in Hawaiian or Mexican waters, but generally did not travel to other feeding regions. The most rapid migratory transit between Hawaii and southeastern Alaska was 79 d. Based on mark‐recapture analyses of the photographic data, we estimate a population of 270–372 whales in the southeastern Alaska feeding herd.