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Social information is predicted to enhance the quality of animals' migratory decisions in dynamic ecosystems, but the relative benefits of social information in the long-range movements of marine megafauna are unknown. In particular, whether and how migrants use nonlocal information gained through social communication at the large spatial scale of oceanic ecosystems remains unclear. Here we test hypotheses about the cues underlying timing of blue whales' breeding migration in the Northeast Pacific via individual-based models parameterized by empirical behavioral data. Comparing emergent patterns from individual-based models to individual and population-level empirical metrics of migration timing, we find that individual whales likely rely on both personal and social sources of information about forage availability in deciding when to depart from their vast and dynamic foraging habitat and initiate breeding migration. Empirical patterns of migratory phenology can only be reproduced by models in which individuals use long-distance social information about conspecifics' behavioral state, which is known to be encoded in the patterning of their widely propagating songs. Further, social communication improves pre-migration seasonal foraging performance by over 60% relative to asocial movement mechanisms. Our results suggest that long-range communication enhances the perceptual ranges of migrating whales beyond that of any individual, resulting in increased foraging performance and more collective migration timing. These findings indicate the value of nonlocal social information in an oceanic migrant and suggest the importance of long-distance acoustic communication in the collective migration of wide-ranging marine megafauna.

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Migratory marine megafauna generally move vast distances between productive foraging grounds and environmentally stable breeding grounds, but characterizing how they use these habitats to maintain homeostasis and reproduce is difficult. We used isotope analysis of blue whale skin strata (n = 621) and potential prey (n = 300) to examine their migratory and foraging strategies in the eastern Pacific Ocean. Our results suggest that most whales in the northeast Pacific use a mixed income and capital breeding strategy, and use the California Current Ecosystem as their primary summer-fall foraging ground. A subset of individuals exhibited migratory plasticity and spend most of the year in the Gulf of California or Costa Rica Dome, two regions believed to be their primary winter-spring breeding grounds. Isotope data also revealed that whales in the southern Eastern Tropical Pacific generally do not forage in the northeast Pacific, which suggests a north-south population structure with a boundary near the equator.

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Linking individual and population scales is fundamental to many concepts in ecology [1], including migration [2, 3]. This behavior is a critical [4] yet increasingly threatened [5] part of the life history of diverse organisms. Research on migratory behavior is constrained by observational scale [2], limiting ecological understanding and precise management of migratory populations in expansive, inaccessible marine ecosystems [6]. This knowledge gap is magnified for dispersed oceanic predators such as endangered blue whales (Balaenoptera musculus). As capital breeders, blue whales migrate vast distances annually between foraging and breeding grounds, and their population fitness depends on synchrony of migration with phenology of prey populations [7, 8]. Despite previous studies of individual-level blue whale vocal behavior via bio-logging [9, 10] and population-level acoustic presence via passive acoustic monitoring [11], detection of the life history transition from foraging to migration remains challenging. Here, we integrate direct high-resolution measures of individual behavior and continuous broad-scale acoustic monitoring of regional song production (Figure 1A) to identify an acoustic signature of the transition from foraging to migration in the Northeast Pacific population. We find that foraging blue whales sing primarily at night, whereas migratory whales sing primarily during the day. The ability to acoustically detect population-level transitions in behavior provides a tool to more comprehensively study the life history, fitness, and plasticity of population behavior in a dispersed, capital breeding population. Real-time detection of this behavioral signal can also inform dynamic management efforts [12] to mitigate anthropogenic threats to this endangered population [13, 14]).

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Knowing the migratory movements and behaviour of baleen whales is fundamental to understanding their ecology. We compared δ15N and δ13C values in the skin of blue (Balaenoptera musculus), fin (Balaenoptera physalus) and sei (Balaenoptera borealis) whales sighted in the Azores in spring with the values of potential prey from different regions within the North Atlantic using Bayesian mixing models to investigate their trophic ecology and migration patterns. Fin whale δ15N values were higher than those recorded in blue and sei whales, reflecting feeding at higher trophic levels. Whales' skin δ15N and δ13C values did not reflect prey from high-latitude summer foraging grounds; instead mixing models identified tropical or subtropical regions as the most likely feeding areas for all species during winter and spring. Yet, differences in δ13C values among whale species suggest use of different regions within this range. Blue and sei whales primarily used resources from the Northwest African upwelling and pelagic tropical/subtropical regions, while fin whales fed off Iberia. However, determining feeding habitats from stable isotope values remains difficult. In conclusion, winter feeding appears common among North Atlantic blue, fin and sei whales, and may play a crucial role in determining their winter distribution. A better understanding of winter feeding behaviour is therefore fundamental for the effective conservation of these species.

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BACKGROUND: Species distribution models have shown that blue whales (Balaenoptera musculus) occur seasonally in high densities in the most biologically productive regions of the California Current Ecosystem (CCE). Satellite telemetry studies have additionally shown that blue whales in the CCE regularly switch between behavioral states consistent with area-restricted searching (ARS) and transiting, indicative of foraging in and moving among prey patches, respectively. However, the relationship between the environmental correlates that serve as a proxy of prey relative to blue whale movement behavior has not been quantitatively assessed. METHODS: We investigated the association between blue whale behavioral state and environmental predictors in the coastal environments of the CCE using a long-term satellite tracking data set (72 tagged whales; summer-fall months 1998-2008), and predicted the likelihood of ARS behavior at tracked locations using nonparametric multiplicative regression models. The models were built using data from years of cool, productive conditions and validated against years of warm, low-productivity conditions. RESULTS: The best model contained four predictors: chlorophyll-a, sea surface temperature, and seafloor aspect and depth. This model estimated highest ARS likelihood (> 0.8) in areas with high chlorophyll-a levels (> 0.65 mg/m3), intermediate sea surface temperatures (11.6-17.5 °C), and shallow depths (< 850 m). Overall, the model correctly predicted behavioral state throughout the coastal environments of the CCE, while the validation indicated an ecosystem-wide reduction in ARS likelihood during warm years, especially in the southern portion. For comparison, a spatial coordinates model (longitude × latitude) performed slightly better than the environmental model during warm years, providing further evidence that blue whales exhibit strong foraging site fidelity, even when conditions are not conducive to successful foraging. CONCLUSIONS: We showed that blue whale behavioral state in the CCE was predictable from environmental correlates and that ARS behavior was most prevalent in regions of known high whale density, likely reflecting where large prey aggregations consistently develop in summer-fall. Our models of whale movement behavior enhanced our understanding of species distribution by further indicating where foraging was more likely, which could be of value in the identification of key regions of importance for endangered species in management considerations. The models also provided evidence that decadal-scale environmental fluctuations can drive shifts in the distribution and foraging success of this blue whale population.

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INTRODUCTION: This is the first worldwide systematic and quantitative study to count and identify helminth parasites from 100 blue and 44 fin whale fecal samples collected in the Gulf of California during winter (1993-2014). RESULTS: Blue and fin whale feces had similar prevalence of adult acanthocephalans (Bolbosoma sp.) in feces (18.2% and 14.6%, respectively), but blue whales had significantly higher helminth egg prevalence in feces (100%) and mean intensity (443 ± 318 eggs/g) compared to fin whales (61%, 252 ± 327 eggs/g). Diphyllobothrium sp. eggs were identified in blue whale feces and Diphyllobothridae, Ogmogaster sp. and Crassicauda sp. eggs were identified in fin whale feces. We tested the hypothesis that egg intensity in blue whale's feces varies as a function of age class, reproductive status, sex, preservation and sampling years using a Generalized Linear Model. This model explained 61% of the variance in the helminth egg intensity, but it was not significant. Eighteen blue whale individuals were resampled over time without significant difference between consecutive samples. CONCLUSIONS: Thus, all individual blue whales that migrate to the Gulf of California during winter are permanently parasitized with helminths, while the resident fin whales showed lower prevalence and intensity. This helminth load difference is likely due to their different diets duringsummer-fall, when blue whales feed on other krill species in the California Current System and fin whales shift to school fish prey types in the Gulf of California.

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Acoustic communication is an important aspect of reproductive, foraging and social behaviours for many marine species. Northeast Pacific blue whales (Balaenoptera musculus) produce three different call types-A, B and D calls. All may be produced as singular calls, but A and B calls also occur in phrases to form songs. To evaluate the behavioural context of singular call and phrase production in blue whales, the acoustic and dive profile data from tags deployed on individuals off southern California were assessed using generalized estimating equations. Only 22% of all deployments contained sounds attributed to the tagged animal. A larger proportion of tagged animals were female (47%) than male (13%), with 40% of unknown sex. Fifty per cent of tags deployed on males contained sounds attributed to the tagged whale, while only a few (5%) deployed on females did. Most calls were produced at shallow depths (less than 30 m). Repetitive phrasing (singing) and production of singular calls were most common during shallow, non-lunging dives, with the latter also common during surface behaviour. Higher sound production rates occurred during autumn than summer and they varied with time-of-day: singular call rates were higher at dawn and dusk, while phrase production rates were highest at dusk and night.

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BACKGROUND: The most traditional scheme for migration among baleen whales comprises yearly migrations between productive waters at high latitude summer feeding grounds and warmer waters at lower latitudes where whales calve and mate, but rarely feed. Evidence indicates, however, that large departures from this scheme exist among populations and individuals. Furthermore, for some populations there is virtually no information on migratory pathways and destinations. Such is the case of Chilean blue whales throughout the Eastern South Pacific; hence, the goal of this study was to assess its migratory behavior. METHODS: Dedicated marine surveys and satellite tagging efforts were undertaken during the austral summer and early autumn on blue whale feeding grounds off Chilean Northern Patagonia (CNP) during 2013, 2015 and 2016. Positional data derived from satellite tags regarding movement patterns and behavior were analyzed using Bayesian switching first-difference correlated random walk models. RESULTS: We instrumented 10 CNP blue whales with satellite transmitters and documented individual variation in departure time, northbound migratory routes and potential wintering grounds. The onset of migration occurred from mid/late austral autumn to well into the austral winter. Blue whales moved in various directions, but ultimately converged toward a general NW movement direction along a wide corridor exceeding 2,000 km. Area-Restricted Search behavior was exhibited within fjords and channels of CNP and also South of Galapagos Archipelago (GA) and northern Peru, but never during migration. Interestingly, dive profiles for one whale that reached GA showed a sharp and consistent increase in depth north of 5°S and extreme deep dives of up to 330 m. DISCUSSION: Information derived from satellite tagged blue whales in this study is the first of its kind off the Eastern Southern Pacific. Our results provide valuable information on their migratory timing, routes and behavior on their northbound migration, particularly regarding the varied migratory plasticity for this particular population. Our results also highlight the first record of two complete migratory paths between CNP and GA and strengthen the hypothesis that GA waters correspond to a potential wintering destination for CNP blue whales. We further hypothesize that this area might be selected because of its biological productivity, which could provide feeding opportunities during the breeding season. Our results suggest that special efforts should be put forward to identify blue whale critical areas and understand key behavioral aspects in order to provide the basis for their conservation on a regional context (i.e., reducing potential ship strike and promote Marine Protected Area (MPA) implementation in Chile, Ecuador and Peru). Indeed, we suggest joint blue whale conservation efforts at the regional level in order to identify and determine potential threats and impacts and, most importantly, implement prospective management actions.

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Steroid hormone assessment using non-invasive sample collection techniques can reveal the reproductive status of aquatic mammals and the physiological mechanisms by which they respond to changes in their environment. A portion of the eastern North Pacific blue whale (Balaenoptera musculus) population that seasonally visits the Gulf of California, Mexico has been monitored using photo-identified individuals for over 30 years. The whales use the area in winter-early spring for nursing their calves and feeding and it therefore is well suited for fecal sample collection. Using radioimmunoassays in 25 fecal samples collected between 2009 and 2012 to determine reproductive state and stress, we validated three steroid hormones (progesterone, corticosterone and cortisol) in adult female blue whales. Females that were categorized as pregnant had higher mean fecal progesterone metabolite concentrations (1292.6 ±â€¯415.6 ng·g-1) than resting and lactating females (14.0 ±â€¯3.7 ng·g-1; 23.0 ±â€¯5.4 ng·g-1, respectively). Females classified as pregnant also had higher concentrations of corticosterone metabolites (37.5 ±â€¯9.9 ng·g-1) than resting and lactating females (17.4 ±â€¯2.0 ng·g-1; 16.8 ±â€¯2.8 ng·g-1, respectively). In contrast, cortisol metabolite concentrations showed high variability between groups and no significant relationship to reproductive state. We successfully determined preliminary baseline parameters of key steroid hormones by reproductive state in adult female blue whales. The presence of pregnant or with luteal activity and known lactating females confirms that the Gulf of California is an important winter-spring area for the reproductive phase of these blue whales. The baseline corticosterone levels we are developing will be useful for assessing the impact of the increasing coastal development and whale-watching activities on the whales in the Gulf of California.

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Vocal behavior of blue whales (Balaenoptera musculus) in the Gulf of Corcovado, Chile, was analysed using both audio and accelerometer data from digital acoustic recording tags (DTAGs). Over the course of three austral summers (2014, 2015 and 2016), seventeen tags were deployed, yielding 124 h of data. We report the occurrence of Southeast Pacific type 2 (SEP2) calls, which exhibit peak frequencies, durations and timing consistent with previous recordings made using towed and moored hydrophones. We also describe tonal downswept (D) calls, which have not been previously described for this population. As being able to accurately assign vocalizations to individual whales is fundamental for studying communication and for estimating population densities from call rates, we further examine the feasibility of using high-resolution DTAG accelerometers to identify low-frequency calls produced by tagged blue whales. We cross-correlated acoustic signals with simultaneous tri-axial accelerometer readings in order to analyse the phase match as well as the amplitude of accelerometer signals associated with low-frequency calls, which provides a quantitative method of determining if a call is associated with a detectable acceleration signal. Our results suggest that vocalizations from nearby individuals are also capable of registering accelerometer signals in the tagged whale's DTAG record. We cross-correlate acceleration vectors between calls to explore the possibility of using signature acceleration patterns associated with sounds produced within the tagged whale as a new method of identifying which accelerometer-detectable calls originate from the tagged animal.

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Many aspects of blue whale biology are poorly understood. Some of the gaps in our knowledge, such as those regarding their basic taxonomy and seasonal movements, directly affect our ability to monitor and manage blue whale populations. As a step towards filling in some of these gaps, microsatellite and mtDNA sequence analyses were conducted on blue whale samples from the Southern Hemisphere, the eastern tropical Pacific (ETP) and the northeast Pacific. The results indicate that the ETP is differentially used by blue whales from the northern and southern eastern Pacific, with the former showing stronger affinity to the region off Central America known as the Costa Rican Dome, and the latter favouring the waters of Peru and Ecuador. Although the pattern of genetic variation throughout the Southern Hemisphere is compatible with the recently proposed subspecies status of Chilean blue whales, some discrepancies remain between catch lengths and lengths from aerial photography, and not all blue whales in Chilean waters can be assumed to be of this type. Also, the range of the proposed Chilean subspecies, which extends to the Galapagos region of the ETP, at least seasonally, perhaps should include the Costa Rican Dome and the eastern North Pacific as well.

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Using bacterial and fungal tag-encoded FLX-Titanium amplicon pyrosequencing, the microbiota of the faecal material of two blue whales living in the wild off the coast of California was investigated. In both samples the most predominant bacterial phylum was the Firmicutes with Clostridium spp. being the most dominant bacteria. The most predominant fungi were members of the phylum Ascomycota with Metschnikowia spp. being the most dominant. In this study, we also preliminarily characterised the culturable anaerobic bacteria from the faecal material, using traditional culture and 16S rRNA gene sequencing approaches. In total, three bacterial species belonging to the phylum Firmicutes were identified.

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A number of parasitic diseases have gained importance as neozoan opportunistic infections in the marine environment. Here, we report on the gastrointestinal endoparasite fauna of three baleen whale species and one toothed whale: blue (Balaenoptera musculus), fin (Balaenoptera physalus), and sei whales (Balaenoptera borealis) and sperm whales (Physeter macrocephalus) from the Azores Islands, Portugal. In total, 17 individual whale fecal samples [n = 10 (B. physalus); n = 4 (P. macrocephalus); n = 2 (B. musculus); n = 1 (B. borealis)] were collected from free-swimming animals as part of ongoing studies on behavioral ecology. Furthermore, skin biopsies were collected from sperm whales (n = 5) using minimally invasive biopsy darting and tested for the presence of Toxoplasma gondii, Neospora caninum, and Besnoitia besnoiti DNA via PCR. Overall, more than ten taxa were detected in whale fecal samples. Within protozoan parasites, Entamoeba spp. occurred most frequently (64.7%), followed by Giardia spp. (17.6%) and Balantidium spp. (5.9%). The most prevalent metazoan parasites were Ascaridida indet. spp. (41.2%), followed by trematodes (17.7%), acanthocephalan spp., strongyles (11.8%), Diphyllobotrium spp. (5.9%), and spirurids (5.9%). Helminths were mainly found in sperm whales, while enteric protozoan parasites were exclusively detected in baleen whales, which might be related to dietary differences. No T. gondii, N. caninum, or B. besnoiti DNA was detected in any skin sample. This is the first record on Giardia and Balantidium infections in large baleen whales.

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Unusually low genetic diversity can be a warning of an urgent need to mitigate causative anthropogenic activities. However, current low levels of genetic diversity in a population could also be due to natural historical events, including recent evolutionary divergence, or long-term persistence at a small population size. Here, we determine whether the relatively low genetic diversity of pygmy blue whales (Balaenoptera musculus brevicauda) in Australia is due to natural causes or overexploitation. We apply recently developed analytical approaches in the largest genetic dataset ever compiled to study blue whales (297 samples collected after whaling and representing lineages from Australia, Antarctica and Chile). We find that low levels of genetic diversity in Australia are due to a natural founder event from Antarctic blue whales (Balaenoptera musculus intermedia) that occurred around the Last Glacial Maximum, followed by evolutionary divergence. Historical climate change has therefore driven the evolution of blue whales into genetically, phenotypically and behaviourally distinct lineages that will likely be influenced by future climate change.

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We examined recordings from a 15-month (May 2009-July 2010) continuous acoustic data set collected from a bottom-mounted passive acoustic recorder at a sample frequency of 6kHz off Portland, Victoria, Australia (38°33'01″S, 141°15'13″E) off southern Australia. Analysis revealed that calls from both subspecies were recorded at this site, and general additive modeling revealed that the number of calls varied significantly across seasons. Antarctic blue whales were detected more frequently from July to October 2009 and June to July 2010, corresponding to the suspected breeding season, while Australian blue whales were recorded more frequently from March to June 2010, coinciding with the feeding season. In both subspecies, the number of calls varied with time of day; Antarctic blue whale calls were more prevalent in the night to early morning, while Australian blue whale calls were detected more often from midday to early evening. Using passive acoustic monitoring, we show that each subspecies adopts different seasonal and daily call patterns which may be related to the ecological strategies of these subspecies. This study demonstrates the importance of passive acoustics in enabling us to understand and monitor subtle differences in the behavior and ecology of cryptic sympatric marine mammals.

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Blue whales (Balaenoptera musculus) were among the most intensively exploited species of whales in the world. As a consequence of this intense exploitation, blue whale sightings off the coast of Chile were uncommon by the end of the 20th century. In 2004, a feeding and nursing ground was reported in southern Chile (SCh). With the aim to investigate the genetic identity and relationship of these Chilean blue whales to those in other Southern Hemisphere areas, 60 biopsy samples were collected from blue whales in SCh between 2003 and 2009. These samples were genotyped at seven microsatellite loci and the mitochondrial control region was sequenced, allowing us to identify 52 individuals. To investigate the genetic identity of this suspected remnant population, we compared these 52 individuals to blue whales from Antarctica (ANT, n = 96), Northern Chile (NCh, n = 19) and the eastern tropical Pacific (ETP, n = 31). No significant differentiation in haplotype frequencies (mtDNA) or among genotypes (nDNA) was found between SCh, NCh and ETP, while significant differences were found between those three areas and Antarctica for both the mitochondrial and microsatellite analyses. Our results suggest at least two breeding population units or subspecies exist, which is also supported by other lines of evidence such as morphometrics and acoustics. The lack of differences detected between SCh/NCh/ETP areas supports the hypothesis that eastern South Pacific blue whales are using the ETP area as a possible breeding area. Considering the small population sizes previously reported for the SCh area, additional conservation measures and monitoring of this population should be developed and prioritized.

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It is generally assumed that species with low population sizes have lower genetic diversities than larger populations and vice versa. However, this would not be the case for long-lived species with long generation times, and which populations have declined due to anthropogenic effects, such as the blue whale (Balaenoptera musculus). This species was intensively decimated globally to near extinction during the 20th century. Along the Chilean coast, it is estimated that at least 4288 blue whales were hunted from an apparently pre-exploitation population size (k) of a maximum of 6200 individuals (Southeastern Pacific). Thus, here, we describe the mtDNA (control region) and nDNA (microsatellites) diversities of the Chilean blue whale aggregation site in order to verify the expectation of low genetic diversity in small populations. We then compare our findings with other blue whale aggregations in the Southern Hemisphere. Interestingly, although the estimated population size is small compared with the pre-whaling era, there is still considerable genetic diversity, even after the population crash, both in mitochondrial (N = 46) and nuclear (N = 52) markers (Hd = 0.890 and Ho = 0.692, respectively). Our results suggest that this diversity could be a consequence of the long generation times and the relatively short period of time elapsed since the end of whaling, which has been observed in other heavily-exploited whale populations. The genetic variability of blue whales on their southern Chile feeding grounds was similar to that found in other Southern Hemisphere blue whale feeding grounds. Our phylogenetic analysis of mtDNA haplotypes does not show extensive differentiation of populations among Southern Hemisphere blue whale feeding grounds. The present study suggests that although levels of genetic diversity are frequently used as estimators of population health, these parameters depend on the biology of the species and should be taken into account in a monitoring framework study to obtain a more complete picture of the conservation status of a population.