Variability In Foraging By Humpback Whales Megaptera Novaenangliae On The Kodiak Alaska Feeding Ground

Humpback whales (Megaptera novaeangliae, Borowski 1781) in the North Pacific migrate from mid- to high- latitude summer feeding grounds along the Pacific Rim, including areas off the coasts of the U.S., Canada, Russia and eastern Asia, to tropical breeding grounds each winter along Pacific coasts of Mexico and Central America as well as the offshore islands of Mexico, Hawaii, and Japan. Humpback whales in the North Pacific and elsewhere were reduced to very low numbers during a period of intense commercial exploitation that ended in 1965. As the population recovers in abundance, the range of cultural and genetic diversity that survived the exploitation-driven bottleneck is able to adapt, endure and evolve. My work uses genetic tools and photo identification data to investigate the population dynamics, mitochondrial (mt) DNA control region evolution and potential drivers of a specialized feeding behavior in a recovering subpopulation of humpback whales in the Glacier Bay and Icy Strait (GBIS) sub-region of the southeastern Alaska (SEAK) feeding ground. I first collated and reconciled available DNA profiles (mtDNA control region, 10 microsatellite loci and sex) from 556 individuals using tissue samples collected from 1987 to 2012. Photo identification records associated with 692 of 1,026 total genetic samples collected in SEAK (now archived within the SEAK DNA Register and Tissue Database) corresponded to extensive life-history information, extending back to the early 1970s, as archived within the SEAK Regional Database, curated by the National Park Service (NPS) and University of Alaska, Southeast (UAS). Changes in population structure in GBIS over 32 years (1973-2005) were investigated in order to determine whether the increase in local abundance was attributable to local fidelity and recruitment or immigration from outside of SEAK. Two temporal strata were defined: 'Founder' individuals identified between 1973-1985 (n = 74, n = 46 with DNA profiles) and 'Contemporary' individuals identified between 2004-2005 (n = 171, n = 114 with DNA profiles). There was no significant genetic differentiation between the strata, indicating that it is unlikely that the population increase within GBIS was due largely to immigration of whales from elsewhere in the North Pacific. However, two additional haplotypes were documented in the Contemporary stratum at low frequency, one of which was previously unreported in the North Pacific (haplotype A8, see below). This relative stability in haplotype frequencies over time argues for strong regional fidelity of the maternal lineages represented in GBIS between 1973 and 1985. After excluding the 42 Contemporary whales with no photo ID record of a mother or genotype available for maternity inference, at least 73.6% (n = 95) of the Contemporary stratum was either a returning Founder or a recruited descendant of a Founder female. Of all genetically confirmed females with genotypes in the Founder stratum, 96% (n = 24) were either represented in the Contemporary stratum, had at least one confirmed descendant in the Contemporary stratum, or both. This high proportion, in addition to the large proportion of the verifiable Contemporary stratum that were either returning Founders or a descendant of a Founder female, provides clear evidence for local fidelity and recruitment to GBIS. The discovery of the A8 haplotype, which differs by one base pair from a common haplotype referred to as A-, represents an increase in mtDNA diversity for the North Pacific humpback whale from 28 to 29 haplotypes. To investigate the origin of this new haplotype, we re-evaluated n = 1089 electropherograms of n = 710 individuals with A- haplotypes from both the SEAK DNA Register and Tissue Database and the ocean-wide program, SPLASH (Baker et al. 2013). From this review, we identified two individuals with the A8 haplotype (a cow and calf, both sampled in GBIS) and n = 20 individuals with clear heteroplasmy for haplotypes A-/A8. The majority of A-/A8 individuals (n = 15) were sampled in SEAK. Genotype exclusion and likelihood were used to identify one of the heteroplasmic females, #196 (first sighted in SEAK in 1982), as the likely mother of the A8 cow and grandmother of the A8 calf, establishing the inheritance and germ-line fixation of the new haplotype from the parental heteroplasmy. Based on life history records and estimates of pairwise relatedness from microsatellite genotypes, it appears likely that the A-/A8 and the A8 individuals are descendants from a common maternal ancestor one or more generations prior to the three generations documented here. Humpback whale sociality takes a distinct form in Icy Strait, where whales form large, coordinated groups with repeated membership across several decades. Twenty-one years of group association records (1985-2005, n = 2,204 groups) were used to investigate the hypothesis that kin selection influences membership in large, stable groups. Of the 2204 groups recorded, 113 consisted of 6 or more individuals; a size considered unexpectedly large assuming a Poisson distribution of group size with a mean of 1.7. A total of n = 71 individuals (n = 48 with DNA profiles) were encountered in a large group in at least one year, n = 38 individuals (n = 34 with DNA profiles) were encountered in a large group in at least two years, n = 29 individuals (n = 27 with DNA profiles) were encountered in a large group in at least three years, decreasing to n = 2 individuals (n = 2 with DNA profiles) that were encountered in a large group in at least 20 years. There were no significant differences in mtDNA frequencies between large group feeders and the Founder and Contemporary strata or when compared to whales never encountered in large groups in Icy Strait, indicating that group membership is not predominately passed through one maternal lineage. Sex ratios did not deviate significantly from 1:1 for those feeding in large groups over an increasing number of years, as would be expected if females were actively recruiting offspring into large groups. The average pairwise relatedness for large group feeders was not significantly greater than expected by chance and did not increase for those feeding in large groups over an increasing number of years. Of the 179 known offspring of females encountered in a large group, only 6% were also encountered in a large group in Icy Strait as an adult and only 2.2% in the same large group as their mother. These results indicate that kin selection is not the primary driver of membership in large, stable groups and pose an interesting dynamic in local habitat use: individuals are recruited to GBIS through local maternal fidelity but do not usually associate closely with direct maternal kin. The extensive collection of DNA profiles now archived with the individual-based data within the SEAK Regional Database allowed us to integrate genetics and photo ID to answer ecologically relevant questions regarding the whales in GBIS. Together, these results demonstrate that GBIS provide habitat for a distinct collection of individuals that exhibit strong fidelity and local recruitment, some of which engage in a highly specialized feeding behavior. Further, GBIS is a local feeding habitat for two individuals with a newly arising North Pacific mtDNA haplotype. These findings reveal local genotypic and cultural variation and highlight the importance of habitat protection for species with fine-scale habitat use and strong fidelity to local migratory destinations.

"A feeding aggregation of humpback whales (Megaptera novaeangliae) in the Kodiak Island region has received little previous study. A mark-recapture experiment was conducted in 2001 and 2002 to estimate its abundance. Historical abundance was back-calculated from this estimate, whaling records, and suspected survival and productivity values within a population model. The current population was estimated at 157 whales and the pre-whaling population at 343 whales. Prey consumption by humpback whales was modeled using three methods for two hypothetical diets based on prey availability surveys conducted within the study area and stomach contents of commercially caught whales. By assuming current consumption is proportional to prey availability, the current population removes an estimated 9,600 tons of prey annually. Historical populations may have removed over 19,000 tons of prey annually"--Leaf iii.

Relative to other baleen whale populations, the humpback whales Megaptera novaeangliae inhabiting Southeast Alaska are noteworthy in that they form large, enduring groups when foraging on schools of Pacific herring Clupea harengus pallasi. These groups use a variety of unusual feeding tactics when attacking prey, including the production of loud feeding calls, the release of bubbles, and the waving of their large pectoral flippers. Past observations of these groups have been largely anecdotal so little is known about their social behavior or the function of their feeding tactics. In particular, it is unclear if these pods are based on cooperative interactions or if they represent groups of individuals who are competing for prey that occurs in dense aggregations. In addition, little is known about the underlying social structure of these pods and whether they are composed of kin. The objective of this study was to employ field and laboratory techniques to gain insight into the function of the unusual feeding tactics, and to describe the basic social structure of these pods. In order to test the prediction that bubbles, feeding calls, and flipper movements represent prey manipulation tactics, herring schools were brought into the laboratory and subjected to various simulated humpback whale feeding behaviors. I found that these simulated behaviors produced strong avoidance responses from the herring schools, suggesting that humpback whales use these tactics to manipulate the behavior of their prey. Seven years of field observations revealed that humpbacks with enduring social bonds (i.e., high coefficients of association) specialized on herring and represented a small proportion of the entire whale population. Calves born to these "core members" were never observed to return and feed with their mothers in subsequent years, suggesting that these groups are not composed of close kin. This was verified by analysis of haplotype variation in the mitochondrial genome and microsatellite variation in the nuclear genome. Individuals within these pods appear to invest in by-product benefits, with the enduring bonds between whales in Chatham Strait (cf. krill feeders in Frederick Sound) possibly representing combinations of individuals performing compatible tasks (Le., bubble blower, herder, vocalizer).

The North Pacific humpback whale (Megaptera novaeangliae) population has been increasing at an average annual rate of ~6% since the early 1990s. In northern Southeast Alaska alone, there are now more whales than estimated for the entire North Pacific several decades ago. An understanding of how this growing population is repopulating traditional foraging grounds will benefit from detailed investigations of their prey preferences and trends in whale abundance and distribution relative to those prey. This dissertation examines these issues from late May until early September 2008 in Frederick Sound and Stephens Passage, a Southeast Alaskan feeding area historically used by humpback whales. The foundation for the study is an analysis of the life histories and abundance patterns of euphausiids, the principal prey of humpbacks in the area, during late spring and summer. Four species, Thysanoessa raschii, T. longipes, T. spinifera, and Euphausia pacifica, were identified in plankton net samples collected at random locations throughout the study site (n = 49) and in locations where a strong scattering layer was observed on a 120 kHz echosounder (n = 48). Both sample types varied in euphausiid species composition. Abundance patterns of immature euphausiids coupled with observations of females carrying spermatophores indicated differences between species in spawning schedules. Thysanoessa spp. began spawning in early April with the spring phytoplankton bloom and continued until late June, whereas E. pacifica began spawning in early June and continued until late August. This protracted recruitment of immature euphausiids was geographically widespread throughout the summer in contrast to adults, which, although present all summer, were found primarily in slope and shallow (

Increases in the humpback whale (Megaptera novaeangliae) population have generated considerable interest in understanding the foraging habits of these large marine predators in the Gulf of Alaska. Globally, humpback whales are classified as generalist predators but are known to exhibit localized differences in diet. Intensified predation pressure is of particular concern to resource managers, who have observed whales feeding at juvenile hatchery salmon release sites in Southeast Alaska. We assessed the diets and behavioral tactics of humpback whales foraging near Hidden Falls Hatchery release sites (in Chatham Strait, 2016 to 2018) to better understand their predatory effects on juvenile hatchery-reared salmon. We used skin biopsies, prey sampling, and stable isotope analysis to estimate whales’ diet composition. Aerial footage and photographic sequences were used to assess the foraging tactics used on this prey source. We observed three individual whales repeatedly feeding on juvenile hatchery-reared salmon, and we were able to sample them multiple times over a period spanning shifts in diet. Overall, the diets of these whales were higher trophically than other humpback whales foraging in the area, even before feeding on juvenile hatchery salmon started. These hatchery-feeding whales may be generally more piscivorous than other whales, which focused on planktivorous prey. Our repeat sampling, in conjunction with scheduled introductions of a novel prey source, provided a semi-controlled feeding experiment that allowed for incorporation and turnover rate estimates from humpback whale tissue in a way that was not previously possible for large, free-ranging cetaceans. Finally, during the course of this study we discovered an undescribed feeding tactic employed by hatchery-associated whales. We observed the use of solo bubble-nets to initially corral prey, followed by calculated movements to establish a secondary boundary with the pectoral fins that further condensed prey and increased foraging efficiency. Our study provided the first empirical evidence for what we describe as “pectoral herding”. This work deepens our knowledge about humpback whale foraging ecology, how this innovative species is able to exploit newly available prey, and to what extent they feed on commercially valuable hatchery salmon.

"Summer distributions of fin (Balaenoptera physalus) and humpback (Megaptera novaeangliae) whales were examined relative to bathymetry, oceanography, and zooplankton composition and density in Marmot and Chiniak Bays (Kodiak Island, AK) during 2002 and 2003. Habitat use and habitat partitioning were assessed using Monte Carlo and randomization tests, logistic regression analyses, and kernel density probability contours of high-use areas. Fin whales associated with deeper, cooler waters near areas of maximum slope and consistently used Marmot Bay. Fin whale spatial-temporal distribution likely coincided with Neocalanus copepod concentrations during early summer and adult euphausiids later in summer. Fin whale associations with Pseudocalanus copepods may relate more to that copepods' prevalence than to relevance as prey. Humpback whale site fidelity and association with shallow waters was evident in 2002, but not in 2003. Variability in humpback whale distribution was likely related to their exploitation of forage fish aggregations and threshold foraging needs. High densities of adult euphausiids may promote spatial overlap and shared resource use by fin and humpback whales. This mesoscale snapshot of a dynamic nearshore marine environment and the whales foraging there is an integral step toward identifying and characterizing important habitats for endangered fin and humpback whales"--Leaf iii.