Population dynamics of sperm whales (Physeter macrocephalus) in Guadeloupe,French Caribbean: A mark-recapture study from 2001 to 2013

Sperm whales are present along the Caribbean islands in family groups of mature females, juveniles, and newborns. Their abundance and demographic rates remain poorly known. Using photo-identification data, we estimated the abundance and annual survival of sperm whales in Guadeloupe from January to April 2001–2013. A total of 1,492 photographs of tail flukes were scored (ranging from 1 to 5) in terms of quality (Q) and marking (M) level. A total of 789 photographs were selected with Q ≥ 3 and M ≥ 3, corresponding to 109 individually identified adult females or immatures of both sexes. Capture histories were built and analyzed using the Pollock's robust design capture-mark-recapture model. The study consisted of 13 primary periods sequenced in 5–8 secondary periods. Mean annual survival rate of residents was 0.945, 95% CI [0.864, 0.979]. Abundance varied between years from 75 to 35 individuals. The geometric growth rate over the period 2001–2013 was 0.938, 95% CI [0.878, 0.997], suggesting a population decline of −6.2%/year. Sperm whales are moving between Guadeloupe and Dominica, with a high temporary emigration rate (0.163) and 50% of the individuals of Guadeloupe resighted in Dominica. These results highlight the fragility of the population and the necessity to consider it as one unit.     

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.     

SPERM WHALE SURFACE ACTIVITY FROM TRACKING BY RADIO AND SATELLITE TAGS1

Three 12-m sperm whales (Physeter catodon) were tagged and tracked west of Dominica in the southeast Caribbean to follow the surfacing patterns and movements of these presumed subadult males. Whale N was tagged in April 1993 with a 30-MHz radio tag and tracked for two days. Whale H was tagged in April 1995 with a 30-MHz radio tag and tracked for 4.6 d. Whale A was tagged in April 1995 with a satellite-monitored tag tracked by ARGOS for 21.5 d, the first four of which were concurrent with the tracking of Whale H, an associate. The tagged whales remained west of Dominica for at least 2, 5, and 13 d, respectively. Whales N and A then moved southward to waters off Martinique. There were no apparent effects on the whales by tagging or the presence of the tags. The whales averaged speeds of 2.6-3.5 km/h. Surfacings, indicated by tag signals, were of two types: short surfacings apparently primarily for respiration, averaging 7-10.5 min between repeated longer dives, occurring day and night; and extended surfacings seemingly for rest and social interactions with conspecifics, occurring mostly in daylight. Whales were near the surface for 20.4%–22.6% of the total time (26.6%–27.1% during the day and 14.9%–17.1% at night). Delayed blowing was observed as Whale N surfaced for 8.3 min between 47- and 45-min dives but delayed the first of its 31 blows for 1.5 min after surfacing.     

Residency and abundance of sperm whales (Physeter macrocephalus) in Nemuro Strait,Hokkaido, Japan

We examined the trend in residence patterns and abundance of male sperm whales in Nemuro Strait, Japan, based on long-term photo-identification (1,513 survey days, total 2,969 photos) between 2006 and 2017. A total of 225 unique individuals were identified during this study, with an average of 36 (SE = 2.55) new individuals identified in each season. The model chosen by maximum likelihood suggests that residence time around Nemuro Strait is 769 (SE = 372.4) days, with individuals staying in the strait about 48 days (SE = 8.36) per year. While the migration patterns of male sperm whales visiting this area are still unclear, these findings along with previous studies suggest that males move from one breeding area to another neighboring area every several weeks, shifting their home ranges gradually over a period of a few years. The abundance of sperm whales in Nemuro Strait varied greatly from year to year; from 28 (95%CI: 24–44) in 2015 and (95%CI: 22–48) in 2016 to 66 (95%CI: 57–84) in 2011. This study provides important knowledge of abundance and residency for Nemuro Strait, information which will contribute to further research on the social structure and movement pattern of male sperm whales.     

DIRECT ESTIMATION OF WITHIN-GROUP HETEROGENEITY IN PHOTO-IDENTIFICATION OF SPERM WHALES

Heterogeneity in photo-identification rates among individuals is a potentially serious problem in many studies of cetacean biology, especially the analysis of populations. However, this heterogeneity is usually difficult to identify or measure. Two instances in which closed groups of female and immature sperm whales ( Physeter macrocepbalus ) were tracked and identified using fluke photographs over periods of days off the Galápagos Islands allowed direct examination of heterogeneity in identification rates. A group of nine animals followed in 1999 provided almost no evidence for heterogeneity (permutation test for heterogeneity, P = 0.48), with an estimated coefficient of variation in identification rates of 0.03 (95% CI from 1,000 bootstrap replications: 0.00–0.10). In contrast, the identification rates of a group of 22 animals followed in 1995 seemed to show potentially important differences ( P = 0.058, CV = 0.20, 95% CI = 0.07–0.28). These differences were not related to the internal social structure of the group or to differences in numbers of markings on the flukes, but smaller whales had lower identification rates. Thus, young sperm whales may be underrepresented in photo-identification studies, but adults within groups seem to have similar identification rates. Situations in which animals are photo-identified from closed populations of known size are particularly useful for examining heterogeneity. They should be vigorously exploited by those who use photo-identification to examine population or behavioral biology.     

GRAY WHALES (ESCHRICHTIUS ROBUSTUS) OFF SAKHALIN ISLAND,RUSSIA: SEASONAL AND ANNUAL PATTERNS OF OCCURRENCE1

The annual return, seasonal occurrence, and site fidelity of Korean-Okhotsk or western gray whales on their feeding grounds off northeastern Sakhalin Island, Russia, were assessed by boat-based photo-identification studies in 1994-1998. A total of 262 pods were observed, ranging in size from 1 to 9 whales with an overall mean of 2.0. Sixty-nine whales were individually identified, and a majority of all whales (71.0%) were observed in multiple years. Annual sighting frequencies ranged from 1 to 18 d, with a mean of 5. 4 d. The percentage of whales reidentified from previous years showed a continuous annual increase, reaching 87.0% by the end of the study. Time between first and last sighting of identified individuals within a given year was 1-85 d, with an overall mean of 40.6 d. Annual calf proportions ranged from 4.3% (1997) to 13.2% (1998), and mother-calf separations generally occurred between July and September. The seasonal site fidelity and annual return of whales to this part of the Okhotsk Sea emphasize its importance as a primary feeding ground for this endangered population.     

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.     

First demographic insights on historically harvested and poorly known male sperm whale populations off the Crozet and Kerguelen Islands (Southern Ocean)

Age and sex dependent spatial segregation has resulted in limited knowledge of the ecology and demography of sperm whale adult males feeding seasonally in high latitudes. This study focused on adult males interacting with the Patagonian toothfish (Dissostichus eleginoides) fishery operating off the Kerguelen and Crozet Archipelagos. Demographic parameters were estimated using a 10‐yr‐long photo‐identification data set paired with multistate closed robust design capture‐mark‐recapture models. The examination of a set of 29,078 photographs taken from fishing vessels during sperm whale depredation events resulted in identification of 295 individuals with nine visiting both study areas. Dispersal between both study regions was estimated to be 1% per year. The mean annual number of interacting sperm whales was estimated to n = 82 (95% CI 58–141) in Crozet and n = 106 (95% CI 76–174) in Kerguelen. Transient proportions were 13% in Crozet and 26% in Kerguelen. Corrected for transience, apparent survival estimates were 0.953 (95% CI 0.890–0.993) in Crozet, and 0.911 (95% CI 0.804–0.986) in Kerguelen. These survival and population size estimates are the first for depredating adult males in high latitudes, and can be used in evaluating the current conservation status of this historically harvested stock and to investigate depredation trends in 35 both Crozet and Kerguelen Islands.     

Short‐finned pilot whales (Globicephala macrorhynchus) of the Mariana Archipelago: Individual affiliations,movements, and spatial use

Little is known about short‐finned pilot whales (Globicephala macrorhynchus) in the western North Pacific outside of Japanese coastal waters. To expand understanding of short‐finned pilot whale ecology in the region, we conducted small‐boat surveys in 2010?2016 within the Mariana Archipelago to investigate individual associations, movements, spatial use, and dive behavior of short‐finned pilot whales. We collected genetic, photo‐identification, and satellite‐tag data and identified 191 distinctive individuals. A preliminary social network diagram of photo‐cataloged individuals revealed a main cluster that comprised 82% of individuals, representing all five mitochondrial DNA haplotypes identified within the population. Kernel density estimates for tagged short‐finned pilot whales (n = 11) during summer were used to identify areas with the highest probability of use (10% probability density contour), core area (50%) and home range (95%). The area with highest probability of use by short‐finned pilot whales was off the northwest side of Guam. Satellite tag data also suggest that some individuals are island‐associated year‐round. Data from five location‐dive tags demonstrated that the short‐finned pilot whales dove more often to intermediate depths at twilight and night, suggesting they may target prey that forage on the deep scattering layer as it migrates to and from the surface.     

Consensus movements by groups of sperm whales

When animals live in cohesive groups they need to make consensus decisions about movements. As a very large‐scale example of communal movement, nomadic female sperm whales (Physeter macrocephalus) travel about 50 km per day as coherent groups of 10–50 animals spread over several km of ocean. From 543 h of data during which 3,873 headings of small clusters of whales or individual whales were recorded, I quantified the heading behavior of groups foraging off the Galápagos Islands. The groups made both sudden and gradual turns. Using piecewise regression models, I estimate that sudden and gradual turns in heading both occurred at rates of 0.10/h. The mean change in heading was 69° for sudden turns and 84° for gradual turns. The mean duration of gradual turns was 1.3 h, so turns were often slow. Using the recorded headings within 30 min of each of 1,798 focal headings, a regression of heading on time gave a mean rate of turn of the group and error of each focal heading about the mean heading. Absolute heading errors increased with absolute turn rate (r_S = 0.241; P = 0.0000), so turns were often messy. Thus sperm whales often make slow and messy—likely democratic—consensus decisions when groups change heading.     

Vocal clans in sperm whales (Physeter macrocephalus)

Cultural transmission may be a significant source of variation in the behaviour of whales and dolphins, especially as regards their vocal signals. We studied variation in the vocal output of 'codas' by sperm whale social groups. Codas are patterns of clicks used by female sperm whales in social circumstances. The coda repertoires of all known social units (n = 18, each consisting of about 11 females and immatures with long-term relationships) and 61 out of 64 groups (about two social units moving together for periods of days) that were recorded in the South Pacific and Caribbean between 1985 and 2000 can be reliably allocated into six acoustic 'clans', five in the Pacific and one in the Caribbean. Clans have ranges that span thousands of kilometres, are sympatric, contain many thousands of whales and most probably result from cultural transmission of vocal patterns. Units seem to form groups preferentially with other units of their own clan. We suggest that this is a rare example of sympatric cultural variation on an oceanic scale. Culture may thus be a more important determinant of sperm whale population structure than genes or geography, a finding that has major implications for our understanding of the species' behavioural and population biology.     

Population size and migratory connectivity of humpback whales wintering in Las Perlas Archipelago,Panama

From 2003 to 2009, we surveyed Las Perlas Archipelago off the Pacific coast of Panama 53 times between the months of August and October to estimate abundance of humpback whales and to test for a migratory connection with populations from the southern hemisphere. We identified 295 individuals using photo‐identification of dorsal fins, including 58 calves, and the population estimate for a single season was 100–300 solitary adults plus 25–50 mothers with calves; the estimated population of animals across all seasons using a mark and recapture model was over 1,000. Eight of the 139 fluke identifications were matched to whales in photograph catalogues from the Antarctic Peninsula and a ninth was matched to a whale sighted in Chilean waters; four of these nine individuals have also been sighted in Colombia. We conclude that Panama (Las Perlas Archipelago in particular) is an important calving area for humpback whales in the Southern Hemisphere. These data should provide a foundation for monitoring of population change and to increase awareness in Panama about the need to manage vessel traffic and tourism related to the whales at Las Perlas.     

Does disappearance mean extirpation? The case of right whales off Namibia

Right whales off Namibia were severely depleted by early 19th century whaling, and rarely featured in modern whaling catches in the 1920s. Aerial surveys of the Namibian coastline from 1978 and onwards revealed increasing numbers of right whales, but few cow‐calf pairs. Aerial surveys off South Africa since 2009 showed a major decline in the availability of animals without calves. Twenty individual matches were made between 94 whales photographed off Namibia/Northern Cape in 2003–2012 and 1,677 photographed off South Africa in 1979–2012. Eight were adult females that calved in South African waters, but only one was also seen with a calf off Namibia. Twelve out of 13 individuals off Namibia with distinctive dorsal pigmentation were first seen as calves off South Africa. These results strongly indicate connectivity between the two regions, while the presence off Namibia of three adult females from the South African population in the season in which they are believed to conceive suggests that there is unlikely to be any genetic differentiation between the two areas. We conclude that the reappearance of right whales off Namibia represents range expansion from South Africa rather than the survival of a few remnants of an originally separate stock.     

High site fidelity,strong associations,and long‐term bonds: Short‐finned pilot whales off the island of Hawai‘i

Studies of short‐finned pilot whales suggest they travel in stable mixed‐sex groups composed of strongly associated individuals; however, temporal analyses of social structure are lacking. To examine site fidelity, association patterns, and temporal relationships, we analyzed data from 267 encounters of this species off the island of Hawai‘i from 2003 through 2007, identifying 448 distinctive individuals (68.1% seen more than once). About 72% of the whales were linked by association into a single social network, suggesting the possibility of multiple populations using the area. Sighting histories suggested that only some individuals exhibit high site fidelity. Individuals demonstrated preferential associations and community division was strongly supported by average‐linkage hierarchical cluster analysis of the association data. Nine longitudinally stable social units composed of key individuals and their constant companions were identified. Qualitative assignment of age and sex classes of unit members indicated that some segregation between adult males and female/calf pairs may occur. Temporal analyses of individuals encountered on the same day indicate stable long‐term associations. Differential patterns of residency and site fidelity were unexpected and may be indicative of multiple populations around the main Hawaiian Islands. The presence of a resident population demonstrating strong, long‐term site fidelity and associations off Hawai‘i Island may warrant special management considerations.     

An anthropological genetic perspective on creolization in the anglophone caribbean

Variable socio‐cultural influences developed in the colonial Caribbean as a result of competing European hegemonic rule. In this study, we examine how colonial regulations regarding social hierarchies and mate choice worked to influence the genetic landscape of contemporary African Caribbean populations. To this end, 420 individuals from Dominica, Grenada, St. Kitts, St. Lucia, St. Thomas, St. Vincent, Jamaica, and Trinidad were genotyped for 105 autosomal ancestry informative markers. Based on these data, population substructure and admixture were assessed using an exact test, a model‐based clustering method, and principal components analysis. On average, individual admixture estimates of the pooled African Caribbean sample were 77% (SD ± 18%) West African, 15% (SD ± 15%) European, and 7.7% (SD ± 8%) Native American. In general, ancestry estimates were significantly different between Dominica and all other islands. Genetic structure analyses indicated subdivision into two subpopulations on most islands. Finally, unlike all of the other Caribbean populations that clustered adjacent to African populations, the Dominican population was more intermediate between the three parental groups in the principal components plot. As a result of the significant French influence throughout Dominican history, Dominica did not have the same cultural influences that typified other Anglophone colonies. Consequently, there were different social hierarchies and resulting mate choices on Dominica compared with the other considered islands. This study highlights the complex socio‐cultural history of a broad region of the Caribbean and attests to the interplay between social and biological factors in shaping the genetic diversity present in present‐day communities. Am J Phys Anthropol 151:135–143, 2013. © 2013 Wiley Periodicals, Inc.     

SPERM WHALES TAGGED WITH TRANSPONDERS AND TRACKED UNDERWATER BY SONAR

Abstract: Two sperm whales tagged with acoustic transponder tags were tracked by sonar during a cruise from 16 to 30 October 1991 in the southeast Caribbean west of Dominica Island. The whales dove to depths of 400–600 m and more, including a dive to 1,185 m and one possibly to 2,000 m. They were tracked for periods of 3–14 h, over distances of 8.5–40 km. The tagged whales were found together four and eight days after tagging, and were tracked simultaneously for 13 h, over 31 km. Whale movements on different days at the surface averaged from 0.68 to 0.82 m/set, with dive descent rates from 0.82 to 1.13 m/set, ascent rates from 0.74 to 1.16 m/set, and horizontal movement during dives from 0.76 to 1.29 m/set. Dives lasted from 18 min to 1 h and 13 min, averaging 33 and 41 min on different days. Every track ended when tag signals became obscured at night by dense biological scatterers concentrated in offshore areas where the whales were diving. Both tagged whales appear to have been males of 15 and 11m, each dominant in different groups; but when together the larger whale was dominant, as evidenced by chases and agonistic vocalizations. The whales did not appear to react to the tags or to the sounds associated with tracking (30, 32, and 36 kHz).     

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.     

Social structure of killer whales (Orcinus orca) in a variable low-latitude environment,the Galápagos Archipelago

Killer whales (Orcinus orca) have strong social matrilineal bonds and form groups and long-lasting associations, but little is known about their population or social structure in an equatorial setting such as the waters around the Galápagos Islands. Using 91 encounters and identification photographs from 1991 to 2017, we identified 64 killer whales of which 18 individuals were locals with high resighting rates. Group size was small, ranging from 1 to 15 animals, with 69% of the groups containing four or fewer animals. Using social network analysis (SOCPROG 2.7) whales grouped into three distinct units and one loose association with frequent exchange between different groups. One male–male unit showed a strong association (association strength = 0.55). Overall, associations lasted over at least 3 years. Our data give first evidence of a loose social organization of Galápagos killer whales, similar to fission-fusion societies.     

Population structure of melon‐headed whales (Peponocephala electra) in the Hawaiian Archipelago: Evidence of multiple populations based on photo identification

Despite the presence of melon‐headed whales in tropical and subtropical waters worldwide, little is known about this species. To assess population structure in Hawai‘i, dedicated field efforts were undertaken from 2000 to 2009. Using only good quality photographs, there were 1,433 unique photo‐identified individuals, of which 1,046 were distinctive. Of these, 31.5% were seen more than once. Resighting data combined with social network analyses showed evidence of two populations—a smaller, resident population, seen exclusively off the northwest region of the island of Hawai‘i, and a larger population, seen throughout all the main Hawaiian Islands (hereafter the “main Hawaiian Islands” population). A Bayesian analysis examining the probability of movements of individuals between populations provided a posterior median dispersal rate of 0.0009/yr (95% CI = 0–0.0041), indicating the populations are likely demographically independent. Depth of encounters with the Hawai‘i Island resident population was significantly shallower (median = 381 m) than those with the main Hawaiian Islands population (median = 1,662 m). Resightings of individuals have occurred up to 22 yr apart for the Hawai‘i Island resident population and up to 13 yr apart for the main Hawaiian Islands population, suggesting long‐term residency to the islands for both populations.     

Migratory movements of individual humpback whales photographed off the eastern coast of Australia

Humpback whales (Megaptera novaeangliae) migrate long distances each year on a return journey from low‐latitude breeding grounds to high‐latitude feeding grounds. Migration is influenced by subtle and complex social behaviors and the assumption that whales transit directly through the migratory corridor off the east coast of Australia requires further investigation. From 2003 to 2005, we followed the movements of 99 individual whales within one migratory cycle from three locations, off Byron Bay during the whales' northern migration and in Hervey Bay and at Ballina during the southern migration. The median sighting interval of whales between Byron Bay and Hervey Bay (n = 26) was 52 d (IQR = 42.5–75.5); between Byron Bay and Ballina (n = 21) was 59 d (IQR = 47.0–70.0); and between Hervey Bay and Ballina (n = 33) was 9 d (8.0–14.0). The overall pattern observed from these resightings suggests that Group E1 humpback whales spend approximately two months in the northern quarter of their range during the austral winter months. Intraseason resightings of whales at Ballina (n = 13, median sighting interval = 7 d) also suggest that some individuals, particularly adult males, may circle back north during their general southward journey along this part of the coast, perhaps in an attempt to increase mating opportunities.