CHARACTERIZATION OF BEAKED WHALE (ZIPHIIDAE) AND SPERM WHALE (PHYSETER MACROCEPHALUS) SUMMER HABITAT IN SHELF-EDGE AND DEEPER WATERS OFF THE NORTHEAST U. S.

Sperm whales ( Physeter macrocephalus ) and beaked whales ( Mesoplodon spp. and Ziphius cavirostris ) are deep-diving cetaceans that frequent shelf-edge and Gulf Stream waters off the northeast U. S. coast. Sighting data collected during seven summer (1990, 1991, 1993, and 1995–1998) shipboard surveys were analyzed using a geographic information system to determine habitat use based on bathymetric and oceanographic features. Although sighting rates were lower for beaked whales, both taxa occupied similar habitats. Beaked whales were concentrated at the colder shelf edge, whereas sperm whales were associated with warmer off-shelf water. Mean sighting rates for both taxa were higher in canyon features, but only beaked whale sighting rates were significantly different between canyon and non-canyon habitat (Wilcoxon signed rank test P = 0.007). Within the shared habitat, the two taxa were separated at fine-scale based on oceanographic features.     

Singing through winter nights: Seasonal and diel occurrence of humpback whale (Megaptera novaeangliae) calls in and around the Gully MPA,offshore eastern Canada

Humpback whale use of areas off eastern Canada is poorly understood, a knowledge gap that could impact future conservation efforts. We describe the acoustic occurrence of humpback whales in and around the Gully Marine Protected Area (MPA), an eastern Scotian Shelf submarine canyon. Near‐continuous acoustic recordings sampling at 16 kHz were collected from the MPA and nearby slope areas from October 2012 to September 2014 using near‐bottom recorders. In an offshore region where humpbacks were thought to be rare, we observed calls from October to June with a peak in song and nonsong calls in December and January. This suggests that some individuals occur in Canadian waters in winter and the Gully region may be a North Atlantic humpback whale migratory corridor. Calls were predominantly songs indicating potential mating activities. Song and nonsong calls occurred more at sunset and during hours of darkness than during daylight. This study improves our understanding of the seasonal occurrence of humpback whales on the Scotian Slope and, more specifically, their use of an offshore protected area.     

KILLER WHALE PREDATION ON SPERM WHALES: OBSERVATIONS AND IMPLICATIONS

In October 1997 we observed a herd of approximately 35 killer whales ( Orcinus orca ) attack a pod of nine sperm whales ( Physeter macrocephalus ) 130 km off the coast of central California. During the four hours we watched, adult female killer whales, including some with calves, attacked in waves of four to five animals in what was apparently a "wound and withdraw" strategy. Adult male killer whales stood by until the very end when one charged in and quickly killed a seriously wounded sperm whale that had been separated from the group. The sperm whales appeared largely helpless: their main defensive behavior was the formation of a rosette ("marguerite"-heads together, tails out). When the killer whales were successful in pulling an individual out of the rosette, one or two sperm whales exposed themselves to increased attack by leaving the rosette, flanking the isolated individual, and leading it back into the formation. Despite these efforts, one sperm whale was killed and eaten and the rest were seriously, perhaps mortally, wounded. We also present details of two other encounters between sperm whales and killer whales that we observed. Although sperm whales, because of various behavioral and morphological adaptations, were previously thought to be immune to predation, our observations clearly establish their vulnerability to killer whales. We suggest that killer whale predation has potentially been an important, and underrated, selective factor in the evolution of sperm whale ecology, influencing perhaps the development of their complex social behavior and at-sea distribution patterns.     

IMPACTS OF COMMERCIAL WHALE WATCHING ON MALE SPERM WHALES AT KAIKOURA, NEW ZEALAND

Abstract Male sperm whales are the basis for a commercially important whale-watching industry at Kaikoura, New Zealand. We examined the influence of whale-watching boats and aircraft over three years using observations from an independent research boat and from shore. We employed an information-theoretic approach to determine which factors were necessary to explain variation in blow interval, time at surface, and time to first click. In almost all analyses, models required the inclusion of the presence of the research boat or whale-watching boats or airplanes. The only exception was the model explaining variation in blow intervals observed from shore, which required only season. We also analyzed spatial behavior at the surface. Resident whales changed direction significantly more in the presence of whale-watching boats compared to encounters with only the research boat present. No such difference was observed for encounters with aircraft. Our results thus indicate that sperm whales off Kaikoura respond to whale-watching activities, although these changes are small and most likely not of biological importance. However, resident whales responded less to these activities compared to transient whales, possibly indicating habituation and, more importantly, the need to monitor continued activities closely.     

Spatial-temporal patterns in intra-annual gray whale foraging: Characterizing interactions between predators and prey in Clayquot Sound, British Columbia, Canada

In Clayquot Sound, British Columbia, gray whales (Eschrichtius robustus) forage primarily on mysids (Family Mysideae) and also on crab larvae (Family Porcellanidae) that are constrained to specific habitat, which relate to bathymetric depths. In this paper we characterize the interactions of gray whales and their prey by analyzing fine scale spatial‐temporal patterns in foraging gray whale distribution within a season. Kernel density estimators are applied to two seasons (1998 and 2002) of high‐resolution data on foraging by gray whales. By partitioning data from each foraging season into several time periods (12 in 1998 and 11 in 2002), using a temporal autocorrelation function, and generating kernel density estimated surfaces for each time period, it is possible to identify discrete areas of increasing and declining foraging effort. Our results indicate that gray whales forage on mysids throughout a season and opportunistically forage on crab larvae. The episodic crab larvae feeding may reduce, but not eliminate, pressure to mysid populations enabling mysids to reassemble swarms and continue to support gray whale foraging in the latter part of the season. Results suggest that when managing marine environments, gray whale populations require multiple and connected habitats for summer foraging.     

Sperm whale depredation of sablefish longline gear in the northeast Pacific Ocean

Interactions between marine mammals and fisheries include competition for prey (catch), marine mammal entanglement in fishing gear, and catch removal off fishing gear (depredation). We estimated the magnitude of sperm whale depredation on a major North Pacific longline fishery (sablefish) using data collected during annual longline surveys. Sperm whale depredation occurs while the longline gear is off‐bottom during retrieval. Sperm whales were observed on 16% of longline survey sampling days, mostly (95% of sightings) over the continental slope. Sightings were most common in the central and eastern Gulf of Alaska (98% of sightings), occasional in the western Gulf of Alaska and Aleutian Islands, and absent in the Bering Sea. Longline survey catches were commonly preyed upon when sperm whales were present (65% of sightings), as evidenced by damaged fish. Neither sperm whale presence (P = 0.71) nor depredation rate (P = 0.78) increased significantly from 1998 to 2004. Longline survey catch rates were about 2% less at locations where depredation was observed, but the effect was not significant (P = 0.34). Estimated sperm whale depredation was <1% of the annual sablefish longline fishery catch off Alaska during 1998 to 2004.     

An assessment of sperm whale occurrence and social structure off São Miguel Island,Azores using fluke and dorsal identification photographs

Female and immature sperm whales form stable social units in tropical to subtropical waters. One such area is the Azores archipelago, where details of their year-round occurrence and social organization are not well known. We used year-round sightings data collected from whale watching vessels to assess sperm whale occurrence and social structure off São Miguel Island, Azores from 2010 to 2017. Individuals were photo-identified by their flukes (n = 393) and dorsal pigmentation marks were examined to explore their use in assisting with reidentifications. Of all cataloged whales, 78.8% were sufficiently distinctive to be reidentified from dorsal pigmentation patterns. Associations between individuals were analyzed to determine social structure and delineate social units. We identified 12 units comprising 2–13 members that had stable, and perhaps preferred associations for periods up to the eight years of the study. Preferences between some pairs of units may exist, but more research is required to better understand dynamics within and between units. This local scale study is an important contributor to our knowledge of geographic variation on a global scale. We recommend making use of all available information from the entire archipelago, to further increase our understanding of sperm social organization in the Azores.     

A SIMPLE PHOTOGRAMMETRIC TECHNIQUE TO MEASURE SPERM WHALES AT SEA

Knowledge of whale length is important to ecological studies. However, photographic techniques to measure sperm whales traditionally require high vantage points or a complicated stereo system. Furthermore, these traditional techniques require an alongside approach that often prevents individual identification. For simple and fast size measurements at sea, I used a laser range finder alongside a digital camera to obtain distance to the fluke at the same time as photo-identification. The camera/lens and laser range finder were calibrated on objects of known lengths. The coefficient of variation (CV) for test objects was low (CV = 0.21%). Forty-seven individually identified sperm whales were measured repetitively on up to 12 different occasions, and the CV was lower (CV = 1.3%) than for other photogrammetric techniques (CV = 4.4%–5.1%). A regression of log fluke span to log total length from whaling and stranding data yielded an r  ² of 0.87 (CV of residuals = 6.7%). Thirty-eight female/immature sperm whales were measured in the Gulf of Mexico (median = 9.3 m, range = 7.1–12.3 m), 167 in the Gulf of California (median = 10.7 m, range = 8.4–13.1 m) and 13 bachelor males off Kaikoura, New Zealand (median = 14.2, range = 11.7–15.8 m). The results were within known sperm whale size and suggested that the population in the Gulf of Mexico was made up of smaller animals than that of the Gulf of California. This technique is easy to implement and allows the measurement of identified individuals.     

Temporal variation in dwarf sperm whale (Kogia sima) habitat use and group size off Great Abaco Island, Bahamas

Dwarf sperm whales (Kogia sima) are among the most commonly stranded yet least known pelagic cetaceans. Few studies have occurred at sea, and none have quantified temporal and spatial variation in dwarf sperm whale abundance and group size. We assessed seasonal and spatial variation in dwarf sperm whale group size and abundance off Great Abaco Island, Bahamas using surveys within a 126 km² study area with depths between 2 and 1,600 m. After correcting for survey effort and variation in sighting efficiency among sea states, we found that dwarf sperm whale group size and habitat use varied seasonally. In summer, dwarf sperm whale groups were small (median = 2.5, range = 1–8) and were found only in the two deep habitats within the study area (slope 400–900 m, deep 900–1,600 m). In winter, group sizes increased (median = 4, range = 1–12) and sightings were almost six times higher in the slope habitat, where vertical relief is highest, than other habitats. Our results suggest that studies of pelagic cetaceans and conservation plans must explicitly account for seasonal variation in group size and habitat use.     

A citizen science approach to long-term monitoring of humpback whales (Megaptera novaeangliae) off Sydney,Australia

The Cape Solander Whale Migration Study is a citizen science project that annually counts northward migrating humpback whales (Megaptera novaeangliae) off Cape Solander, Sydney, Australia. Dedicated observers have compiled a 20-year data set (1997–2017) of shore-based observations from Cape Solander's high vantage point. Using this long-term data set collected by citizen scientists, we sought to estimate the humpback whale population trend as it continues to recover postexploitation. We estimated an exponential growth rate of 0.099 (95% CI = 0.079–0.119) using a generalized linear model, based on observer effort (number of observation days) and number of whales observed, equating to 10% per annum growth in sightings since 1997. We found that favorable weather conditions for spotting whales off Cape Solander consisted of winds <30 km/hr from a southerly through a north westerly direction. Incidental observations of other cetacean species included the endangered blue whale (Balaenoptera musculus) and data deficient species such as killer whales (Orcinus orca) and false killer whales (Pseudorca crassidens). Citizen science-based studies can provide a cost-effective approach to monitoring wildlife over the time necessary to detect change in a population. Information obtained from citizen science projects like this may help inform policy makers responsible for State and Federal protection of cetaceans in Australian waters and beyond.     

Submarine canyons: hotspots of benthic biomass and productivity in the deep sea

Submarine canyons are dramatic and widespread topographic features crossing continental and island margins in all oceans. Canyons can be sites of enhanced organic-matter flux and deposition through entrainment of coastal detrital export, dense shelf-water cascade, channelling of resuspended particulate material and focusing of sediment deposition. Despite their unusual ecological characteristics and global distribution along oceanic continental margins, only scattered information is available about the influence of submarine canyons on deep-sea ecosystem structure and productivity. Here, we show that deep-sea canyons such as the Kaikoura Canyon on the eastern New Zealand margin (42°01′ S, 173°03′ E) can sustain enormous biomasses of infaunal megabenthic invertebrates over large areas. Our reported biomass values are 100-fold higher than those previously reported for deep-sea (non-chemosynthetic) habitats below 500 m in the ocean. We also present evidence from deep-sea-towed camera images that areas in the canyon that have the extraordinary benthic biomass also harbour high abundances of macrourid (rattail) fishes likely to be feeding on the macro- and megabenthos. Bottom-trawl catch data also indicate that the Kaikoura Canyon has dramatically higher abundances of benthic-feeding fishes than adjacent slopes. Our results demonstrate that the Kaikoura Canyon is one of the most productive habitats described so far in the deep sea. A new global inventory suggests there are at least 660 submarine canyons worldwide, approximately 100 of which could be biomass hotspots similar to the Kaikoura Canyon. The importance of such deep-sea canyons as potential hotspots of production and commercial fisheries yields merits substantial further study.     

Cooperative hunting behavior,prey selectivity and prey handling by pack ice killer whales (Orcinus orca), type B,in Antarctic Peninsula waters

Currently, there are three recognized ecotypes (or species) of killer whales (Orcinus orca) in Antarctic waters, including type B, a putative prey specialist on seals, which we refer to as “pack ice killer whale” (PI killer whale). During January 2009, we spent a total of 75.4 h observing three different groups of PI killer whales hunting off the western Antarctic Peninsula. Observed prey taken included 16 seals and 1 Antarctic minke whale (Balaenoptera bonaerensis). Weddell seals (Leptonychotes weddellii) were taken almost exclusively (14/15 identified seal kills), despite the fact that they represented only 15% of 365 seals identified on ice floes; the whales entirely avoided taking crabeater seals (Lobodon carcinophaga; 82% relative abundance) and leopard seals (Hydrurga leptonyx; 3%). Of the seals killed, the whales took 12/14 (86%) off ice floes using a cooperative wave‐washing behavior; they produced 120 waves during 22 separate attacks and successfully took 12/16 (75%) of the Weddell seals attacked. The mean number of waves produced per successful attack was 4.1 (range 1–10) and the mean attack duration was 30.4 min (range 15–62). Seal remains that we examined from one of the kills provided evidence of meticulous postmortem prey processing perhaps best termed “butchering.”     

Nuclear and mitochondrial markers reveal distinctiveness of a small population of bottlenose whales (Hyperoodon ampullatus) in the western North Atlantic

Small populations at the edge of a species' distribution can represent evolutionary relics left behind after range contractions due to climate change or human exploitation. The distinctiveness and genetic diversity of a small population of bottlenose whales in the Gully, a submarine canyon off Nova Scotia, was quantified by comparison to other North Atlantic populations using 10 microsatellites and mitrochondrial DNA (mtDNA) control region sequences (434 bp). Both markers confirmed the distinctiveness of the Gully (n = 34) from the next nearest population, off Labrador (n = 127; microsatellites -F(ST)= 0.0243, P < 0.0001; mtDNA -Phi(ST) = 0.0456, P < 0.05). Maximum likelihood microsatellite estimates suggest that less than two individuals per generation move between these areas, refuting the hypothesis of population links through seasonal migration. Both males and females appear to be philopatric, based on significant differentiation at both genomes and similar levels of structuring among the sexes for microsatellites. mtDNA diversity was very low in all populations (h = 0.51, pi = 0.14%), a pattern which may be due to selective sweeps associated with this species' extreme deep-diving ecology. Whaling had a substantial impact on bottlenose whale abundance, with over 65 000 animals killed before the hunt ceased in the early 1970s. Genetic diversity was similar among all populations, however, and no signal for bottlenecks was detected, suggesting that the Gully is not a relic of a historically wider distribution. Instead, this unique ecosystem appears to have long provided a stable year-round habitat for a distinct population of bottlenose whales.     

Behavior and social structure of the sperm whales of Dominica,West Indies

There is substantial geographic variation in the behavior and social structure of sperm whales worldwide. The population in the Eastern Caribbean is thought to be isolated from other areas in the North Atlantic. We describe the behavior and social structure of the sperm whales identified off Dominica during an eight year study (2005–2012; 92% of photographic identifications) with supplementary data collected from seven other organizations dating as far back as 1981. A total of 419 individuals were identified. Resighting rates (42% of individuals between years) and encounter rates with sperm whale groups (mean = 80.4% of days at sea) among this population were both comparatively high. Group sizes were small (7–9 individuals) and were comprised of just one social unit (mean = 6.76 individuals, SD = 2.80). We described 17 units which have been reidentified off Dominica across 2–27 yr. Mature males are seen regularly off Dominica, but residency in the area lasts only a few days to a few weeks. Males were reidentified across years spanning up to a decade. Management of this population within the multinational Wider Caribbean Region will require governments to work towards international agreements governing sperm whales as a cross‐border species of concern.     

Hybridization of Southern Hemisphere blue whale subspecies and a sympatric area off Antarctica: impacts of whaling or climate change?

Understanding the degree of genetic exchange between subspecies and populations is vital for the appropriate management of endangered species. Blue whales (Balaenoptera musculus) have two recognized Southern Hemisphere subspecies that show differences in geographic distribution, morphology, vocalizations and genetics. During the austral summer feeding season, the Antarctic blue whale (B. m. intermedia) is found in polar waters and the pygmy blue whale (B. m. brevicauda) in temperate waters. Here, we genetically analyzed samples collected during the feeding season to report on several cases of hybridization between the two recognized blue whale Southern Hemisphere subspecies in a previously unconfirmed sympatric area off Antarctica. This means the pygmy blue whales using waters off Antarctica may migrate and then breed during the austral winter with the Antarctic subspecies. Alternatively, the subspecies may interbreed off Antarctica outside the expected austral winter breeding season. The genetically estimated recent migration rates from the pygmy to Antarctic subspecies were greater than estimates of evolutionary migration rates and previous estimates based on morphology of whaling catches. This discrepancy may be due to differences in the methods or an increase in the proportion of pygmy blue whales off Antarctica within the last four decades. Potential causes for the latter are whaling, anthropogenic climate change or a combination of these and may have led to hybridization between the subspecies. Our findings challenge the current knowledge about the breeding behaviour of the world's largest animal and provide key information that can be incorporated into management and conservation practices for this endangered species.     

Potential vessel collisions with Southern Hemisphere humpback whales wintering off Pacific Panama

Vessel collision is a threat to many whale species, and the risk has increased with expanding maritime traffic. This compromises international conservation efforts and requires urgent attention from the world's maritime industry. Humpback whales (Megaptera novaeangliae) are at the top of the death toll, and although Central America is a wintering area for populations from both the Northern and Southern Hemispheres, existing efforts to reduce ship‐whale collisions are meager. Herein, we evaluated the potential collisions between vessels and humpback whales wintering off Pacific Panama by following the movements of 15 whales tagged with satellite transmitters and comparing these data with tracks plotted using AIS real‐time latitude‐longitude points from nearly 1,000 commercial vessels. Movements of whales (adults and calves) in the Gulf of Panama coincide with major commercial maritime routes. AIS vessel data analyzed for individual whale satellite tracks showed that 53% (8 whales) of whales had 98 encounters within 200 m with 81 different vessels in just 11 d. We suggest implementing a 65 nmi Traffic Separation Scheme and a 10 kn maximum speed for vessel routing into the Gulf of Panama during the wintering season. In so doing, the area for potential whale‐vessel collisions could be reduced by 93%.     

Abundance and Distribution of Sperm Whales in the Canary Islands: Can Sperm Whales in the Archipelago Sustain the Current Level of Ship-Strike Mortalities?

Sperm whales are present in the Canary Islands year-round, suggesting that the archipelago is an important area for this species in the North Atlantic. However, the area experiences one of the highest reported rates of sperm whale ship-strike in the world. Here we investigate if the number of sperm whales found in the archipelago can sustain the current rate of ship-strike mortality. The results of this study may also have implications for offshore areas where concentrations of sperm whales may coincide with high densities of ship traffic, but where ship-strikes may be undocumented. The absolute abundance of sperm whales in an area of 52933 km², covering the territorial waters of the Canary Islands, was estimated from 2668 km of acoustic line-transect survey using Distance sampling analysis. Data on sperm whale diving and acoustic behaviour, obtained from bio-logging, were used to calculate g(0) = 0.92, this is less than one because of occasional extended periods when whales do not echolocate. This resulted in an absolute abundance estimate of 224 sperm whales (95% log-normal CI 120–418) within the survey area. The recruitment capability of this number of whales, some 2.5 whales per year, is likely to be exceeded by the current ship-strike mortality rate. Furthermore, we found areas of higher whale density within the archipelago, many coincident with those previously described, suggesting that these are important habitats for females and immature animals inhabiting the archipelago. Some of these areas are crossed by active shipping lanes increasing the risk of ship-strikes. Given the philopatry in female sperm whales, replacement of impacted whales might be limited. Therefore, the application of mitigation measures to reduce the ship-strike mortality rate seems essential for the conservation of sperm whales in the Canary Islands.     

Whale killers: Prevalence and ecological implications of killer whale predation on humpback whale calves off Western Australia

Reports of killer whales (Orcinus orca) preying on large whales have been relatively rare, and the ecological significance of these attacks is controversial. Here we report on numerous observations of killer whales preying on neonate humpback whales (Megaptera novaeangliae) off Western Australia (WA) based on reports we compiled and our own observations. Attacking killer whales included at least 19 individuals from three stable social groupings in a highly connected local population; 22 separate attacks with known outcomes resulted in at least 14 (64%) kills of humpback calves. We satellite‐tagged an adult female killer whale and followed her group on the water for 20.3 h over six separate days. During that time, they attacked eight humpback calves, and from the seven known outcomes, at least three calves (43%) were killed. Overall, our observations suggest that humpback calves are a predictable, plentiful, and readily taken prey source for killer whales and scavenging sharks off WA for at least 5 mo/yr. Humpback “escorts” vigorously assisted mothers in protecting their calves from attacking killer whales (and a white shark, Carcharodon carcharias). This expands the purported role of escorts in humpback whale social interactions, although it is not clear how this behavior is adaptive for the escorts.     

A new species of baleen whale (Balaenoptera) from the Gulf of Mexico,with a review of its geographic distribution

Bryde's-like whales are a complex of medium-sized baleen whales that occur in tropical waters of all three major ocean basins. Currently, a single species of Bryde's whale, Balaenoptera edeni Anderson, 1879, is recognized, with two subspecies, Eden's whale, B. edeni edeni and Bryde's whale, B. edeni brydei (Olsen, 1913), although some authors have recognized these as separate species. Recently, a new, evolutionarily divergent lineage of Bryde's-like whale was identified based on genetic data and was found to be restricted primarily to the northern Gulf of Mexico (GOMx). Here, we provide the first morphological examination of a complete skull from these whales and identify diagnostic characters that distinguish it from the other medium-sized baleen whale taxa. In addition, we have increased the number of genetic samples of these Bryde's-like whales in the GOMx from 23 to 36 individuals, all of which matched the GOMx lineage. A review of Bryde's-like whale records in the Caribbean and greater Atlantic supports an isolated distribution for this unique lineage, augmenting the genetic and morphological body of evidence supporting the existence of an undescribed species of Balaenoptera from the Gulf of Mexico.