BILIARY BILE ACID COMPOSITION OF THE PHYSETERIDAE (SPERM WHALES)

Abstract: The bile acid composition of bile obtained from the hepatopancreatic ducts of three species of sperm whales (Cetacea: Physeteridae) was investigated. Bile acids were isolated by adsorption chromatography and analyzed by sequential HPLC, SIMS, and GLC-MS. In each species the dominant bile acids were deoxycholic acid (a secondary bile acid formed by bacterial 7α-dehydroxylation of cholic acid), and chenodeoxycholic acid (a primary bile acid) which together composed more than 86% of biliary bile acids in all three species. In Physeter catodon (sperm whale) deoxycholic acid constituted 79%, and in Kogia breviceps (pygmy sperm whale) it was 61% of biliary bile acids. The sperm whale, which differs from other whales in having a remnant of a large intestine, is the second mammal identified to date in which deoxycholic acid is the predominant bile acid. The high proportion of deoxycholic acid indicates that in the Physeteridae, anaerobic fermentation occurs in its cecum, and that bile acids undergo enterohepatic cycling. Also found were minor proportions of cholic acid, as well as bacterial derivatives of chenodeoxycholic acid (ursodeoxycholic acid, lithocholic acid, and the 12β-epimer of allo-deoxycholic acid). Bile acids were conjugated with taurine in all species; however, in the sperm whale ( Physeter ) glycine conjugates were present in trace proportions. The bile acid hydroxylation pattern (12α- but not 6α-hydroxylation), lack of primary 5α- (allo) bile acids, and presence of glycine conjugated bile acids suggests the possibility that sperm whales originated from ancient artiodactyls.     

PYGMY SPERM WHALES KOGIA BREVICEPS IN THE NORTHEAST ATLANTIC: NEW INFORMATION ON STOMACH CONTENTS AND STRANDINGS

Little is known about the feeding ecology of pygmy sperm whales ( Kogia breviceps ) in the Northeast Atlantic. Results are presented on the stomach contents of five whales stranded on the Galician coast (NW Spain) between 1995 and 2002 and seven whales stranded on the French Atlantic coast between 1984 and 2001. These results are compared with those obtained from the stomach contents of two pygmy sperm whales (a pregnant female and her calf ) stranded on the Scottish (UK) coast in 1999, the first records of the species in Scotland. In 13 out of 14 cases, food remains consisted almost entirely of cephalopod beaks, although some crustacean and fish remains were also present. In all the Spanish specimens, the identified prey were oceanic species: the cephalopods Histioteuthis reversa , H. bonnellii , Todarodes sagittatus , the viperfish Chauliodus sloani , and the giant mysid Gnatophausia sp. The same cephalopod species were found in the stomachs of the whales stranded in Scotland, although both whales had also consumed neritic cephalopod species such as Rossia macrosoma and other sepiolids. In the French specimens, almost all prey identified were oceanic cephalopods ( H. reversa , Brachioteuthis riseii , T. sagittatus , Taonius pavo , etc. ), but remains of crustaceans and a neritic squid ( Loligo forbesi ) were also found. One whale from France had eaten mainly Henslow's swimming crab ( Polybius henslowi ). Results from the present study are consistent with those found by other authors in the Azores and the Canary Islands in that pygmy sperm whales appear to be mainly teuthophagous, with histioteuthid squids forming an important part of the diet. Strandings records suggest that occurrence of pygmy sperm whales in the NE Atlantic may be seasonal, with most strandings occurring in autumn and winter.     

THE TAPETUM FIBROSUM IN THE EYES OF TWO SMALL WHALES

The pygmy sperm whale ( Kogia breviceps ) and the bottlenose dolphin ( Tursiops truncatus ) are equipped with a tapetum fibrosum and, with other Cetacea, are the only carnivores known to possess this typically ungulate tapetal type. Tapeta from two regions of a retina, each with a different spectral reflectance (blue and green), were found to have significantly different fibrillar diameters and inter-fibrillar spacing. When the measured values are applied to a dielectric reflector model, the predicted wavelengths agree with the observed reflectance of the flat-mounted tapeta. The spatial properties of the fibrils change progressively from the tapetal origin in the fibroblast layer to the pigment epithelium, suggesting that different wavelengths may be reflected systematically with tapetal depth. The very large number of reflecting layers characterizing these tapeta, relative to those of other carnivores, may provide for increased spectral purity and efficiency.     

GENETIC VARIATION OF KOGIA SPP. WITH PRELIMINARY EVIDENCE FOR TWO SPECIES OF KOGIA SIMA

Concordance between mitochondrial DNA (mtDNA) markers and morphologically based species identifications was examined for the two currently recognized Kogia species. We sequenced 406 base pairs of the control region and 398 base pairs of the cytochrome b gene from 108 Kogia breviceps and 47 K. sima samples. As expecred, the two sister species were reciprocally monophyletic to each other in phylogenetic reconstructions, but within K. sima , we unexpectedly observed another reciprocally monophyletic relationship. The two K. sima clades resolved were phylogeographically concordant with all of the haplotypes in one clade observed solely among specimens sampled from the Atlantic Ocean and with those in the other clade observed solely among specimens sampled from the Indo-Pacific Ocean. These apparently allopatric clades were observed in all phylogenetic reconstructions using the maximum parsimony, maximum likelihood, and neighborjoining algorithms, with the mtDNA gene sequences analyzed separately and combined. The nucleotide diversity for the combined gene sequence haplotypes of the two K. sima clades resolved in our analyses was 0.58% and 1.03% for the Atlantic and Indo-Pacific, respectively, whereas for the two recognized sister species, nucleotide diversity was 1.65% and 4.02% for K. breviceps and K. sima , respectively. The combined gene sequence haplotypes have accumulated 44 fixed base pair differences between the two K. sima clades compared to 20 fixed base pair differences between the two recognized sister species. Although our results are consistent with species-level differences between the two K. sima clades, recognition of a third Kogia species awaits supporting evidence that these two apparently allopatric clades represent reproductively isolated groups of animals.     

ESTIMATING BODY LENGTH OF PYGMY RIGHT WHALES (CAPEREA MARGINATA) FROM MEASUREMENTS OF THE SKELETON AND BALEEN

Various parts of the skeleton and/or the longest baleen plate of 46 specimens of Caperea marginata from Australia and New Zealand were measured and related to body length. Of the 32 skull, postcranial and baleen-plate measurements available, eight were analysed and seven found to be good predictors of body length, by using a curvilinear model describing their relationship with body length. Greatest skull width, supraoccipital length and mandible length had the smallest prediction limits (± 0.28-0.33 m in small animals, ±0.44-0.58 m in large animals) when compared with postcranial measurements (scapula length, vertebra 7 centrum width). Baleen-plate length was also a useful predictor of body length (±0.32-0.77 m). There was a substantial increase in the arch of the skull as body length increased. Bulla length was not a good predictor of body length, because measurements were highly variable and because the bulla grew little during postnatal life. Physical maturity occurred at body lengths of at least 5.9 m, also the shortest length at which both epiphyses of the humerus and proximal epiphyses of the radius and ulna were fused. Weaning appears to occur at about 3-3.5 m. The following approximate relative age/length classes were erected: dependent calves, <3.6 m; subadults, 3.6-5.5 m; adults, >5.5 m. Females were significantly longer than males in the sample of 22 animals greater than 5.9 m, length of the smallest recorded physically mature animal.     

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.     

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.     

Foraging ecology and niche overlap in pygmy (Kogia breviceps) and dwarf (Kogia sima) sperm whales from waters of the U.S. mid‐Atlantic coast

A complementary approach of stomach content and stable isotope analyses was used to characterize the foraging ecology and evaluate niche overlap between pygmy (Kogia breviceps) and dwarf (K. sima) sperm whales stranded on the U.S. mid‐Atlantic coast between 1998 and 2011. Food habits analysis demonstrated both species were primarily teuthophagous, with 35 species of cephalopods, and 2 species of mesopelagic fishes represented in their overall diets. Pianka's Index of niche overlap suggested high overlap between whale diets (O_n = 0.92), with squids from the families Histioteuthidae, Cranchidae, and Ommastrephidae serving as primary prey. Pygmy sperm whales consumed slightly larger prey sizes (mean mantle length [ML] = 10.8 cm) than dwarf sperm whales (mean ML = 7.8 cm). Mean prey sizes consumed by pygmy sperm whales increased with growth, but showed no trend in dwarf sperm whales. Significant differences were not detected in δ¹⁵N and δ¹³C values of muscle tissues from pygmy (10.8‰ ± 0.5‰, ?17.1‰ ± 0.6‰), and dwarf sperm whales (10.7‰ ± 0.5‰, ?17.0‰ ± 0.4‰), respectively. Isotopic niche widths also did not differ significantly and dietary overlap was high between the two species. Results suggest the feeding ecologies of the pygmy and dwarf sperm whales are similar and both species occupy equivalent trophic niches in the region.     

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.     

ATOC AND OTHER FACTORS AFFECTING THE DISTRIBUTION AND ABUNDANCE OF HUMPBACK WHALES (MEGAPTERA NOVAEANGLIAE) OFF THE NORTH SHORE OF KAUAI

Humpback whale diving behavior changes subtly when exposed to signals transmitted from the Acoustic Thermometry of Ocean Climate (ATOC) sound projector located 14 km offshore the island of Kauai. This study considered whether such responses would lead to changes in distribution and abundance. A land-based shore station measured humpback whale locations (scan samples) for both nearshore (<5 km) and offshore (5–10 km) areas. Control observations were made in 1994 and 1998. In 1998 multipleday blocks with ATOC transmissions were interspersed with multiple-day control blocks without transmissions. Sighting rates were higher in 1998 (ATOC) than in 1994 (control year), probably due to better sighting conditions, but may reflect increased population size. Sighting rates did not differ between control and ATOC conditions in 1998. A seasonal sighting peak was observed in both years. No vessel effect on sighting rate was detected in 1998.
There was no effect of ATOC on the distance from the shore station to whales, or the depth of water where pods were located. However, the distribution of whales shifted slightly eastward during the ATOC blocks and the mean distance between the ATOC source and pods was greater during transmissions. Nonetheless, more whales were found close to the source when it was on, suggesting a more variable response rather than simple avoidance, with whales found both closer to, and farther away from, the source during transmissions.     

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.     

A DWARF FORM OF THE SPINNER DOLPHIN (STENELLA LONGIROSTRIS) FROM THAILAND

A very small form of the spinner dolphin has been found to inhabit the Gulf of Thailand. Ten specimens taken incidentally in a local shrimp fishery differ from specimens of this species collected elsewhere in body size and shape, skull size and shape, number of teeth and numbers of thoracic and lumbar vertebrae. Four cranially adult males were 129–137 cm long, well below the ranges for the Indian Ocean, western Pacific, central Pacific, eastern Pacific and Atlantic. The skull is also very small. Tooth counts and vertebral counts average lower than in other series. The color pattern is not significantly different from that of spinner dolphins in the central and western Pacific, Atlantic and Indian Oceans but differs from that of the small eastern spinner of the eastern Pacific. The Gulf of Thailand specimens are morphologically separable from all other specimens, but it is to be expected that when larger samples are available there will be some overlap. The dwarf form may overlap in body and skull size with small spinner dolphins taken incidentally in a gillnet fishery for sharks off northern Australia. The geographic range of the dwarf spinner may be restricted. The size and status of the population and the impact of the shrimp fishery are unknown and should be investigated. The dwarf spinner may have an ecology different from that of other spinners, feeding mainly on reef-associated and benthic organisms rather than mesopelagic animals.     

FUNCTIONAL AND EVOLUTIONARY ASPECTS OF THE POSTCRANIAL ANATOMY OF DICYNODONTS (SYNAPSIDA, THERAPSIDA)

Abstract:  Restoration of the major skeletal muscles and functional morphological analysis of the postcranium were carried out on two Triassic dicynodont genera, Wadiasaurus and Lystrosaurus . A phylogenetic analysis of 12 selected Permian and Triassic dicynodont taxa was conducted and the postcranial character states were then mapped onto the most parsimonious tree. The analysis revealed changes in pectoral girdle and forelimb morphology, which included reduction of the coracoid plate, increasing robustness of the deltopectoral crest, change in humeral orientation from lateral to caudolateral, increasing prominence of the humeral head, and increasing robustness of the radius. Such changes can be associated with a functional tendency to reduce the lateral component of the propulsive force while still in an abducted mode. On the other hand, changes associated with the pelvic girdle included expansion of the preacetabular iliac process, reduction of the postacetabular iliac process, craniocaudal expansion of the iliac blade, change in the shape of the pubis from flat and plate-like to small and rod-like with a cranial process, and change in acetabular orientation from lateral to caudolateral. The femoral head, starting from a cranioproximal position, progressively became dorsally pronounced and offset from the body. Other features/changes associated with the femur included increasing robustness of the trochanter major, and increasing flattening of the femoral midshaft. Changes in the axial skeleton included increasing stiffening of the trunk to reduce lateral undulations, increasing dorsoventral flexion, and increasing sacral vertebral count, which can be correlated with the preacetabular iliac expansion. These findings suggest that the dicynodont postcranial skeleton evolved towards more upright hindlimb morphology with the body held well off the ground.     

RESPONSE OF NORTH ATLANTIC RIGHT WHALES (EUBALAENA GLACIALIS) TO PLAYBACK OF CALLS RECORDED FROM SURFACE ACTIVE GROUPS IN BOTH THE NORTH AND SOUTH ATLANTIC

The surface active group (SAG) is the most obvious social interaction of the North Atlantic right whale ( Eubalaena glacialis ). SAGs are typically composed of an adult female with two or more males engaged in social behavior near the surface. Distinct calls, believed to be produced by the female, are associated with these groups. Calls recorded from three North Atlantic right whale SAGs and three South Atlantic right whale ( Eubalaena australis ) SAGs were played back to North Atlantic right whales to determine if these sounds are sufficient to attract males to the groups. Playbacks of gunshot sounds produced by North Atlantic right whales were used as a control stimulus. Thirty-six trials were carried out from 1999 to 2001 in the Bay of Fundy, Canada. Whales approached 27 of 31 SAG playbacks and 0 of 5 gunshot playbacks. Where sex was determined ( n = 28), all approaches to North Atlantic SAG recordings were by males. Individuals ( n = 22) of all age and sex classes approached South Atlantic SAG playbacks. These trials indicate that SAG calls from both populations are sufficient to attract right whales to SAGs and that males and females respond differently to stimuli from the North Atlantic. The difference in response to North and South Atlantic SAG stimuli was unexpected. Novelty, species differences in calls, and different seasonal or behavioral context for the recorded stimuli may be responsible for the differences in response.     

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.     

白鲟肝脏和胰脏的组织学与形态学研究

白鲟肝脏较大,可分为左右两大叶及小的中叶,胆囊位于右叶的凹缺内。肝板多由双层细胞构成,肝小叶不明显。肝内毛细胆小管由4个肝实质细胞围成。胰脏有3支,被厚的浆膜。胰岛明显。胰管与胆管汇合后共同开口于小肠最前部背面。对肝、胰实质细胞的显微或亚显微结构进行了描述。     

STOMACH CONTENTS OF FALSE KILLER WHALES (PSEUDORCA CRASSIDENS) STRANDED ON THE COASTS OF THE STRAIT OF MAGELLAN,TIERRA DEL FUEGO

We examined the stomachs of 25 false killer whales collected from a mass stranding of 181 animals along both coasts of the Strait of Magellan, Chile, in March 1989. The 21 stomachs (nine males and 12 females), with food remains contained 11 prey species (nine cephalopods and two fishes) with a total of 442 individuals. Except for one case, food remains were meager, indicating that the animals had not eaten for some time or through stress had vomited on the beach. Eleven of the 21 animals had mud (often with squid beaks) in the esophagus and first stomach. The prey were identified employing squid beaks, fish otoliths and bones, and their wet weights were estimated using regressions between hard parts and known weights of species. The most important prey were the oceanic and neritic-oceanic squids, Martialia byadesi and Illex argentinus, followed by the neritic fish, Macruronus magellanicus. Of less importance were the oceanic squid, Todaroes fillipovae, the oceanic and epipelagic octopus, Ocytboe sp., and the oceanic squid, Moroteuthis ingens. The rest of the prey were poorly represented and included four oceanic squids and one neritic fish. The prey species of these animals were subantarctic, with two antarctic species, abundant over the Patagonian shelf and adjacent oceanic waters around Tierra del Fuego.     

THE GUNSHOT SOUND PRODUCED BY MALE NORTH ATLANTIC RIGHT WHALES (EUBALAENA GLACIALIS) AND ITS POTENTIAL FUNCTION IN REPRODUCTIVE ADVERTISEMENT

North Atlantic right whales ( Eubalaena glacialis ) make a short, distinctive broadband sound that is produced internally called a Gunshot sound. This sound has been recorded in the Bay of Fundy, Canada from both single whales ( n = 9) and social surface active groups ( n =49). Those single whales producing Gunshot sounds whose sex could be determined ( n = 9) were all mature males. Gunshot sounds were produced as part of a stereotyped behavioral sequence by these individuals, including frequent head-lifts and flipper slapping at the surface. In surface active groups, Gunshot sounds were commonly recorded when males were present in the group. The rate of production of Gunshot sounds was weakly correlated with the total number of males present in the group. Given the behavioral context of Gunshot sound production, and production of the sound only by male whales, Gunshots may function in a reproductive context as an advertisement signal to attract females, an agonistic signal directed toward other males, or a combination of the two functions.     

A REVISION OF THE SYSTEMATICS OF THE NIZYMENIACEAE (GIGARTINALES,RHODOPHYTA) BASED ON POLYSACCHARIDES,ANATOMY, AND NUCLEOTIDE SEQUENCES1

The Australian endemic family Nizymeniaceae, based on Nizymenia australis Sonder, consists of three species in the two genera Nizymenia (1 sp.) and Stenocladia (2 spp.). We have reassessed the generic composition of the family based on evidence from nonfibrillar wall polysaccharides, vegetative anatomy, and the nucleotide sequences of an internal transcribed spacer, ITS 2, of the nuclear ribosomal cistron. Investigation of the polysaccharides by constituent sugar analysis, sulfate content determination, and methylation analysis, combined with gas chromatography-mass spectrometry and infrared analysis, showed that the polysaccharides elaborated by the three species were branched, highly sulfated xylogalactans. These polysaccharides also contained significant amounts of mono-O-methyl galactose (5–8 mol% of total sugars), mainly 4-O-methyl galactose. Although no discrete chemical structures could be assigned to the polysaccharides, the analyses showed that those from Nizymenia australis and Stenocladia australis (Sonder) Silva were more alike than either was to that from S. furcata (Harvey) J. Agardh. This polysaccharide affinity was echoed by a suite of vegetative anatomical features. However, the only likely synapomorphy was the presence of refractive, thick-walled medullary rhizines in both N. australis and S. australis. The ITS 2 sequences were inferred from direct sequencing of the products of polymerase chain reaction amplification. Comparison of the ITS 2 sequences of its three species with those of two outgroups indicated that the family Nizymeniaceae is monophyletic but that interspecific relationships within the family could not be resolved. We conclude that there is insufficient evidence to separate any of the species from the rest at the genus level. Therefore, all three species are consolidated into the genus Nizymenia. This necessitates nomenclatural changes of Stenocladia australis to Nizymenia conferta (Harvey) Chiovitti, Saunders, et Kraft comb. nov.