AGGREGATIONS OF MATURE MALE SPERM WHALES ON THE GALÁPAGOS ISLANDS BREEDING GROUND

The distribution and behavior of mature (12.3–16.3 m) male sperm whales ( Physeter macrocephalus ) were studied on the Galápagos Islands breeding ground from April to June 1995. In contrast to previous research seasons when males were observed only in close spatial and temporal proximity to mixed schools of females and immature animals, in 1995 males were sighted in loose aggregations, separated by hours to days from our vessel's encounters with mixed schools. Only one of ten identified males was observed in spatial proximity to a mixed school.
Aggregations consisted of two to four (minimum estimates) mature males travelling within a range of a few kilometers and were characterized by consistency of heading among individuals. Aggregations moved over time. During encounters, one to three males were observed at the surface at the same time, with interindividual distances of less than 1,000 m. Synchrony of heading was apparent between spatial associates, and its extent appeared to be related to interindividual distance. Clustering (two or more individuals within 100 m) was observed on only two occasions. No agonistic behaviors were seen.
Functions of mature male aggregation on a breeding ground remain unclear. Possible explanations for our observations are local prey abundance, or some form of sociality mediated over spatial scales of hundreds to thousands of meters.     

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.     

A MASS STRANDING OF SPERM WHALES IN OREGON: SEX AND AGE COMPOSITION OF THE SCHOOL

A school of 41 sperm whales, Physeter macrocephalus , that stranded near the mouth of the Siuslaw River, Oregon (43°59'N, 124°08'W), on 16 June 1979 consisted of 13 males and 28 females. Their ages were estimated by assuming that each postnatal dentin layer represents one year. The males were all sub-adults, 9.3–11.5 m long and 14–21 yr old. The females were 9.3–11.4 m long, and 11 to about 58 yr old. Ten females were dissected; nine were sexually mature and three were carrying fetuses 2.64–4.62 m long. None of the 28 females was visibly lactating.     

DIETS OF FIN, SEI, AND SPERM WHALES IN BRITISH COLUMBIA: AN ANALYSIS OF COMMERCIAL WHALING RECORDS, 1963–1967

Diets of fin ( Balaenoptera physalus ), sei ( Balaenoptera borealis ), and sperm whales ( Physeter macrocephalus ) were estimated from the stomach contents of individuals killed along the British Columbia coast from 1963 to 1967. The dominant prey types of fin whales were euphausiids, with minor contributions from copepods and fish. Sei whale stomachs contained primarily copepods in three years, whereas euphausiids or a variety of fish dominated the diet in the other two years. Sperm whales consumed primarily North Pacific giant squid ( Moroteuthis robusta ), but secondary prey differed between males and females. Female sperm whales frequently consumed ragfish ( Icosteus spp.) and other fish, whereas the male diet also contained rockfish ( Sebastes spp.). The high abundance of euphausiids along the British Columbia coast likely contributed to the presence of a summer resident population of fin whales. The high abundance of large copepods farther north probably influenced the migration of sei whales through the offshore waters of British Columbia. Sperm whale stomach contents differed by sex reflecting location and possibly breeding behaviors.     

AN ASSESSMENT OF CHANGES WITH TIME IN THE MARKING PATTERNS USED FOR PHOTOIDENTIFICATION OF INDIVIDUAL SPERM WHALES, PHYSETER MACROCEPHALUS

We observed changes with time in the patterns of characteristic fluke markings used to identify sperm whales. Changes were categorized as minor, moderate, or major based on their severity. These change types were found to occur at rates of 0.9%, 11.8%, and 1.3% per individual per year, respectively. Gain and loss rates for each of seven different mark types were also calculated. The highest estimated rate was the gain of small nicks at 0.08 per individual per year. Most individuals identified by us possess at least a few characteristic marks and, therefore, changes of the type observed in this study are unlikely to severely affect their recognizability. For all but one mark type, gain rates were higher than loss rates, indicating that individuals may be accumulating marks with age. Over long periods this could eventually make individuals unrecognizable, with the result that population sizes calculated from these data may be overestimated. As long as photoidentification studies are conducted sufficiently often, and these changes are as gradual as they appear to be, this problem should be minimal.     

THREE MASS STRANDINGS OF SPERM WHALES (PHYSETER MACROCEPHALUS) IN SOUTHERN AUSTRALIAN WATERS

One hundred and fifteen sperm whales (97 female, 15 male, 3 unknown sex) were involved in three mass stranding events during the month of February 1998 along the west and northwest coastlines of Tasmania, Australia. Sixty-six of these whales stranded at Ocean Beach, Strahan; 35 at Greens Beach, Marrawah; and 11 at Black River Beach, Stanley. The remaining whales stranded singly along the coastline. Three mass strandings of this species in such close temporal proximity have not been reported in this area before, and this is the first time that data have been comprehensively collected from complete or near-complete groups of sperm whales from Tasmanian waters. Adult females dominated the three stranding groups. Total lengths ranged from 417 to 1,200 cm and ages ranged from 0.75 to 64 yr. Four females were lactating and four fetuses were found amongst the groups. Stomach contents were dominated by pelagic cephalopods.     

MIGRATION AND POPULATION STRUCTURE OF NORTHEASTERN PACIFIC WHALES OFF COASTAL BRITISH COLUMBIA: AN ANALYSIS OF COMMERCIAL WHALING RECORDS FROM 1908-1967

Data recorded from 24,862 whales killed by British Columbia coastal whaling stations between 1908 and 1967 revealed trends in the abundance, sex ratios, age structure, and distribution of sperm ( Physeter macrocephalus ), fin ( Balaenoptera physalus ), sei ( Balaenoptera borealis ), humpback ( Megaptera novaeangliae ), and blue ( Balaenoptera musculus ) whales. The catch data were analyzed using annual and monthly mean values. Monthly and annual variation in whaling effort was deduced from accounts of the history of British Columbia coastal whaling, and biases arising from changes in effort were considered in the interpretation of the results. During the later years of whaling (1948-1967), the mean lengths of captured whales declined significantly and pregnancy rates dropped to near zero in fin, sei, and blue whales. Monthly patterns in numbers killed revealed a summer migration of sei and blue whales past Vancouver Island, and confirms anecdotal suggestions that local populations of fin and humpback whales once spent extended periods in the coastal waters of British Columbia. Furthermore, the data strongly suggest that sperm whales mated (April-May) and calved (July-August) in British Columbia's offshore waters. The historic whaling records reveal much about the migratory behavior and distribution of the large whales species as they once were, and may continue to be, in the northeastern Pacific. Verifying the persistence of these trends in the remnant populations is a necessary and logical next step.     

VARIATION IN THE VISUALLY OBSERVABLE BEHAVIOR OF GROUPS OF GALÁPAGOS SPERM WHALES1

The behavior of groups of female and immature sperm whales (Physeter macrocephalus) was measured on 117 d within an 11-yr period off the Galápagos Islands, Ecuador. On each day, up to 18 measures of visually observable behavior were calculated. These concerned speeds, headings, movement patterns, diving synchrony, foraging formations, time spent socializing, and aerial behavior. The measured behavior of the sperm whales was considerably more variable when they were socializing than when foraging. None of the measures showed much correlation with sea-surface temperature, and only measures of consistency of movement were significantly correlated with defecation rate, an indicator of feeding success. However, month-long time periods accounted for over 50% of the variance in eight of eighteen measures, and, in the cases of surface speed and dive synchrony, the effects were statistically significant. In contrast, there was no autocorrelation with lag of one day in the residuals of any of the measures. Thus, behavior may be tracking substantial temporal variation in the whales' environment over scales of about several months. Groups of whales had significantly different travel patterns, but there was little other evidence for group-specific behavior, perhaps because tests of group-specific effects were not of adequate statistical power. Variation in sperm whale behavior, especially over time scales of a few months or longer and spatial scales of a few hundred kilometers or larger, should be considered when estimating densities from sighting surveys.     

SEASONAL OCCURRENCE OF SPERM WHALE (PHYSETER MACROCEPHALUS) SOUNDS IN THE GULF OF ALASKA, 1999–2001

An acoustic survey for sperm whales was conducted in the Gulf of Alaska. Six autonomous hydrophones continuously recorded sound signals below 500 Hz from October 1999 to May 2001. After recovery, recordings were processed using an automatic process to detect usual clicks of sperm whales. The detection algorithm equalized background noise, summed the data in a frequency band, and then used autocorrelation to detect the whales' highly regular clicks. Detections were checked manually, revealing that 98% of detections did contain clicks. Results indicate that sperm whales are present in the Gulf of Alaska year-round; this result extends what is known from whaling data, which were gathered principally in summer. Sperm whales were more common in summer than winter by a factor of roughly two, and occurred less often at the westernmost site surveyed (52°N, 157°W) than elsewhere in the Gulf. This is the first study of sperm whales based exclusively on remote acoustic sensing. This methodology is feasible because sperm whale clicks extend to frequencies (∼100 Hz) low enough to be recorded by low-sample-rate instruments that operate continuously, and because the detection algorithm has a low false-detection rate. The methodology may be replicated to facilitate comparisons between different time periods and geographic regions.     

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.     

海峡两岸的宽尾凤蝶

本记述分布于台湾海峡两岸的Agehana属风蝶两种：中国宽尾风蝶Agehana elwesl Leech和台湾宽尾凤蝶A．maraho Shiraki et Sonan，是我国特有的两个珍稀蝶种．前分布于福建武夷山和其他自然保护区山岳地带，以及北京、陕西，安徽．江西，浙江．湖南、贵州等地，后只分布于台湾省．对于A．elwesi的成虫形态特征作了详细描述，两种凤蝶的区别外观。包含雄外生殖器的形态特征，绘图并列表比较如．研究结果认为李传隆(1984)提出的宽尾风蝶台湾亚种Agehana elwesl marahe Shiraki et Sonan的论述不能成立，确认它和大陆的宽尾风蝶仍然应为两个独立的种．     

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.     

ESTIMATES OF SPERM WHALE ABUNDANCE IN THE NORTHEASTERN TEMPERATE PACIFIC FROM A COMBINED ACOUSTIC AND VISUAL SURVEY

We estimate the abundance of sperm whales in a 7.8 million km² study area in the eastern temperate North Pacific using data from a ship-based acoustic and visual line-transect survey in spring 1997. Sperm whales were detected acoustically using a hydrophone array towed at 15 km/h and 100 m depth. The hydrophone array was towed for 14,500 km, and locations were estimated acoustically for 45 distinct sperm whale groups. Whales producing slow clicks (>2-s period) were detected at greater distance (up to 37 km), and the estimation of effective strip widths was stratified based on initial click period. Visual survey effort (using 25° binoculars and naked eyes) covered 8,100 km in Beaufort sea states 0–5 and resulted in only eight sightings. The effective strip width for visual detections was estimated from previous surveys conducted using the same methods and similar vessels in the eastern Pacific. Estimated sperm whale abundance in the study area was not significantly different between acoustic (32,100, CV = 0.36) and visual (26,300, CV = 0.81) detection methods. Acoustic techniques substantially increased the number of sperm whales detected on this line-transect survey by increasing the range of detection and allowing nighttime surveys; however, visual observations were necessary for estimating group size.     

SPERM WHALES IN THE GULF OF CALIFORNIA: RESIDENCY, MOVEMENTS, BEHAVIOR, AND THE POSSIBLE INFLUENCE OF VARIATION IN FOOD SUPPLY

Sperm whale movements, residency, population structure, and behavior were investigated in the Gulf of California in 1998 and 1999. Variations in sperm whale movement patterns and behavior were related to changes in prey abundance (jumbo squid, Dosidicus gigas ) determined by fishery statistics. Photo-identification data revealed that seven female sperm whales moved into the Gulf of California from the Galapagos Islands, traveling up to 3,803 km. These are among the longest documented movements for female sperm whales. There were significant differences in speed and distance traveled during a dive cycle between 1998 and 1999 (low and high squid abundance). In 1999 there were also significant differences in small-scale movements and behavior between the northern and the southern part of the study area (high and low prey abundance). These results suggest that when food resources are low, sperm whales travel in straighter lines, dive for longer periods, travel larger distances during dive cycles, and at higher speed. In 1999 there were significant differences in time spent socializing in areas of high prey abundance versus areas of low abundance. All of these changes in behavior were consistent with increased foraging effort when squid abundance was low. A high proportion of mature males and first-year calves were observed in the Gulf of California, suggesting that it is an important sperm whale breeding ground.     

舟山渔场及附近海域虾类群落结构特征分析

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PHYLOGENETIC RELATIONSHIPS OF BRYDE'S WHALES IN THE WESTERN NORTH PACIFIC AND ADJACENT WATERS INFERRED FROM MITOCHONDRIAL DNA SEQUENCES1

To clarify phylogenetic relationships of Bryde's whales, we examined the nucleotide sequence of the mitochondrial control region and cytochrome b gene in 33 animals: 12 from offshore waters of the western North Pacific, five from off the Solomon Islands, and 16 from the East China Sea and coastal waters of Kochi in southwestern Japan. For reference purposes, homologous sequences from four Balaenoptera species including four Bryde's whales collected in the eastern Indian Ocean were added. We found whales from the three sampling areas to be genetically distinct. The control region sequences suggested that the whales from the three areas separate at higher than the populational level from one another. The cytochrome b data indicated that genetic differences between whales off the Solomon Islands and animals in the other two areas are equivalent to values found among recognized Balaenoptera species, although such a relationship was not observed between the other two areas. We conclude that whales in the East China Sea and coastal waters of Kochi separate from Bryde's whales in offshore waters of the western North Pacific at higher than the populational level but lower than the specific level (i. e., at the subspecific level) and that whales off the Solomon Islands do not belong genetically to the Bryde's whale as previously recognized.     

THE NORTHEASTERN ALBORAN SEA, AN IMPORTANT BREEDING AND FEEDING GROUND FOR THE LONG-FINNED PILOT WHALE (GLOBICEPHALA MELAS) IN THE MEDITERRANEAN SEA

Little is known about the long-finned pilot whale's population size, structure, distribution, and dynamics in the Western Mediterranean basin. The research region covered since 1992 in southeast Spain, at the edge of the Alboran Sea, is considered an important oceanographic transition zone between the Mediterranean and the Atlantic Ocean. The research ship Toftevaag carried out surveys covering a total of 10,173 nmi (18,840 km) from April to September each year, 1992-1997, all years pooled. Effort for ten-by-ten-mile quadrants was stratified by depth and sea state to ascertain encounter rates. Tracking was used together with photo-identification of animals to analyze home range of groups. Behavior was recorded ad libitum , and underwater video taping was used to analyze specific behavior patterns. One hundred and nine sightings of pilot whales were made. The average group size was 41.4 ± 58.4, ranging from 1 to 350. The average depth at encounters was 848.7 ± 281.2 m ranging from 300 to 1,800 m. Compatison of results for encounter rate and group size with those for other Mediterranean regions, together with site fidelity shown by photo-identification and observations of reproductive behavior, reflect the importance of the Alboran Sea to the species in the Mediterranean.     

THE MICROSCOPIC AND IMMUNOHISTOLOGIC ANATOMY OF THE ENDOCRINE PANCREAS OF PYGMY AND DWARF SPERM WHALES (KOGIIDAE)

Histologic studies of pancreatic tissues of one pygmy sperm whale, Kiogia breviceps , and one dwarf sperm whale, K. simus , demonstrated rather typical exocrine pancreatic anatomy. Peroxidase-antiperoxidase (PAP) techniques determined that the cell composition of the islets of Langerhans resembled that of other mammals. Within islets, cells secreting insulin (B-cells) and glucagon (A-cells), were clearly demonstrated, but, surprisingly, isolated A- and B-cells were also found among pancreatic acinar cells. PAP techniques demonstrated the presence of neuron-specific enolase within islets, but failed to provide a sufficiently clear reaction to demonstrate the presence of somarostatin-producing D-cells. No positive PAP reaction for serotonin occurred.     

黄,渤海沿岸水域游泳动物群落结构时空格局异质性研究

依１９８０年至１９８５年搜集的系统生态学资料分析,黄渤海沿水域游泳动物群落由１８５种动物（１７７种鱼类和８种头足类）组成,暖温性和冷温性的复合区系,以及偶见种,季节种和习见种的多重生态学成分,以暖温性习见种占群落优势地位,其ＮＥＤ和ＢＥＤ分别占６８．８６％和６５．５９％,暖水性偶见种仅占０．１０％和０．１９％。群落结构的季节变化可归结为依ＢＥＤ聚合成暖季组（５－９月）和依ＮＥＤ聚合为冷季组（１０－