Body size of male Atlantic walruses (Odobenus rosmarus rosmarus) from Svalbard

Morphometric data are given for 41 adult male Atlantic walruses ( Odobenus rosmarus rosmarus) immobilized at Svalbard. Total body weight ranged from 960 to 1450 kg. Standard body length ranged from 258 to 380 cm, while curvilinear tusk length ranged from 17 to 65 cm. Immobilized animals were selected for their large size. The sample is therefore biased towards larger animals. Based on regression equations developed for walruses caught in Greenland, the weights of Svalbard walruses were estimated from standard body length and axillary girth. Estimated weights ranged from 632 to 1883 kg. The present data suggest that Atlantic walruses at Svalbard can attain the same body size as Pacific ( O. r. divergens ) and Laptev walruses (O. r. laptevi).     

A morbillivirus antibody survey of Atlantic walrus, narwhal and beluga in Canada

A longitudinal serologic survey was conducted for morbillivirus antibodies in Atlantic walruses (Odobenus rosmarus rosmarus), narwhal (Monodon monoceros), and beluga (Delphinapterus leucas) from the Northwest Territories, Nunavut and the St. Lawrence estuary (Canada). Sixty-five of 131 (50%) walruses sampled between 1984 and 1993 had detectable morbillivirus neutralizing antibodies. Positive walrus were identified from four of five Arctic sampling sites, to as far back as 1984. Prevalence of morbillivirus neutralizing antibodies in walruses from Foxe Basin ranged from a high of 76% (n = 21) in 1993 to a low of 22% (n = 28) in 1984. Limitations in sample acquisition may have produced underestimates for the 1984 data. There are no reports of clinical morbillivirus infection in walruses. Our results are consistent with the hypothesis that a morbillivirus similar or identical to phocine distemper virus (PDV) has circulated among walrus populations of the eastern Canadian Arctic, at least since the early 1980s. No narwhal (n = 79) or beluga (n = 445) from Arctic waters were identified as having antibodies to dolphin morbilivirus (DMV) above the threshold serum dilution of log2 4. Also, none of the beach-cast cetacean carcasses (n = 28) from the Gulf of St. Lawrence and the St. Lawrence estuary were positive for antibodies to DMV. This indicates that Gulf of St. Lawrence, St. Lawrence estuary, and Arctic cetaceans either have not been exposed to DMV or an antigenically related morbillivirus, or are not susceptible to infection.     

REPRODUCTION IN MALE ATLANTIC WALRUSES (ODOBENUS ROSMARUS ROSMARUS) FROM THE NORTH WATER (N BAFFIN BAY)

Age at sexual maturity and timing of the mating season were determined in male Atlantic walruses ( Odobenus rosmarus rosmarus , L.) from the "North Water" subpopulation in northern Baffin Bay. Testes and epididymides of 174 male walruses (between 0 and 30 yr old) from NW Greenland (1987–1990) were studied macroscopically and a subset of 57 specimens was analyzed microscopically. In physically mature bulls ( i.e. , ≥12 yr old), sperm or apparently ripe spermatids were found between 9 November and 12 July. In younger walruses these signs of fertility were found in a few specimens (7–11 yr old) collected between 9 January and 28 May. The mating season seems to peak in January—April. The youngest sexually mature individual was 7 yr old and the oldest apparently immature individual was 13 yr old. Average age of sexual maturity was 10.9 yr (95% C.I.: 9–6–12.2 yr) and all were sexually mature by the time they were 14 yr old. The non-spermiogenetic testes and epididymides showed accelerated growth between about the 5–6th and about the 12–15th year of life, indicating that sexual maturation occurs during these years. The length of the baculum increased gradually until about 12–15 yr of age, when physical maturity was reached.     

FEMALE REPRODUCTIVE PATTERNS AND FETAL GROWTH OF ATLANTIC WALRUSES (ODOBENUS ROSMARUS ROSMARUS) IN FOXE BASIN,NORTHWEST TERRITORIES,CANADA

We examined the reproductive tracts of 79 female Arlantic walruses (Odohenus rosmarus rosmarus) caught in July and August by Inuit hunters in northern Foxe Basin, NWT, Canada. We found no evidence of past or present ovulation in animals less than 5 yr old, but the poor representation of juvenile and adolescent animals precluded statistical analysis of the average age of maturity. The ovulation frequency of 64 mature females (0.52) indicated biennial ovulation, while the pregnancy rate (0.33) and birth rate (0.30) suggested that females gave birth once every three years. The sex ratio of 17 fetuses was not significantly different from 1:1. Fetuses increased 2.9 mm d^-1 in crown-rump length and 0.12 g^1/3 d^-1 in mass over the range of collection dates. Based on the inverse regressions of fetal growth, implantation occurred in late June early July.     

Body growth in Atlantic walruses (Odobenus rosmarus rosmarus) from Greenland

Morphometric data were collected from 105 Atlantic walruses ( Odobenus rosmarus rosmarus ) in northwestern Greenland in the periods 1977–78 and 1989–91. Of these 21 walruses were subjected to a detailed study on body composition.
The asymptotic maximum standard body length of Atlantic walruses in NW Greenland was 269 cm for females and 314 cm for males. This is similar to Pacific walruses, but significantly longer than Atlantic walruses from Hudson Bay. Despite this, walruses from NW Greenland apparently do not attain the same total body mass as Pacific walruses ( O.r.divergens ).
The asymptotic maximum body weights for walruses in NW Greenland were estimated to be 720 kg for females and 1114 kg for males.
Total body surface area was proportional to the two-thirds power of total body weight.
The percentage proportion of blubber and viscera were both negatively correlated to body mass, while skin and muscles constituted a nearly fixed proportion. On average, blubber constituted 19% of total body mass of adult females, 15% of adult males and 24% of subadults of both genders. The average walrus consisted of 18% blubber, 12% skin, 12% viscera and 58% blood, muscle and skeleton. Muscles were estimated to constitute 44% of total body weight.
Allometric functions for weight of internal organs relative to body mass are presented.     

Population structure and gene flow of the Atlantic walrus (Odobenus rosmarus rosmarus) in the eastern Atlantic Arctic based on mitochondrial DNA and microsatellite variation

The population structure of the Atlantic walrus, Odobenus rosmarus rosmarus , was studied using 11 polymorphic microsatellites and restriction fragment length polymorphism detected in the NADH-dehydrogenase ND1, ND2 and ND3/4 segments in mtDNA. A total of 105 walrus samples were analysed from northwest (NW) Greenland, east (E) Greenland, Svalbard and Franz Joseph Land. Two of the 10 haplotypes detected in the four samples were diagnostic for the NW Greenland sample, which implied that the group of walruses in this area is evolutionary distinct from walruses in the other three areas. One individual sampled in E Greenland exhibited a Pacific haplotype, which proved a connection between the Pacific walrus and walruses in eastern Greenland. The Franz Joseph Land, Svalbard and E Greenland samples shared the most common haplotype, indicating very little differentiation at the mtDNA level. Gene flow ( Nm ) estimates among the four areas indicated a very restricted exchange of female genes between NW Greenland and the more eastern Atlantic Arctic samples, and a closer relationship between the three samples composing the eastern Atlantic Arctic. The genetic variation at 11 polymorphic microsatellite loci grouped individuals into three populations, NW Greenland, E Greenland and a common Franz Joseph Land–Svalbard population, which were connected by moderate gene flow.     

Proportion of higher trophic-level prey in the diet of Pacific walruses (Odobenus rosmarus divergens)

During nutritionally stressful situations, Pacific walruses (Odobenus rosmarus divergens) may switch from preying on benthic invertebrates to higher trophic-level prey (HTLP) (e.g., pinnipeds and/or seabirds). We applied a Bayesian mixing model to stable isotope (C and N) data from analyses of various tissues (tongue and lumbar muscle, skin, and liver) to quantify the proportional contribution of HTLP to walruses (n = 293 individuals). The mode contribution of HTLP to walrus diet was ~22 % (±10 %) based on muscle mixing models, which is consistent with results from contaminant studies of Atlantic walruses (Odobenus rosmarus rosmarus), but higher than estimates based on historical stomach content analyses of Pacific walruses. A broader range in the proportion of HTLP (0–60 %) shown by mixing models using stable isotope data from liver and skin of walruses indicated they pursue an opportunistic foraging strategy. Data from the HTLP-consuming walruses were comparable with our stable isotope data of a known “seal-eating” walrus. No significant difference was evident between the estimated contributions of HTLP to the diet of male versus female walruses (P > 0.01). This finding suggests that changes in diet base for walruses are not influenced by the sex of the predator.     

Differences in size and growth rates of male and female migrating Japanese eels in Pearl River, China

The Sr/Ca ratios in otoliths of silver Japanese eels Anguilla japonica , in Pearl River, China, indicated that both sexes did not stay in brackish water and grew in fresh water from the glass eel stage until spawning migration. This did not support the hypothesis that females tended to distribute upstream and males might be restricted to estuaries. The back-calculated total length of males at glass eel stage was not significantly different from that of females, indicating that the hypothesis that small glass eels became males and larger ones became females may not be true. The mean (±S.D.) age and total length of males at migration were 6·4±1·6 years and 48·3±4·5 cm, which were significantly smaller than for females, 8·3±1·6 years and 61·4±4·1 cm. The age of migration was related inversely to growth rate for both sexes. Growth parameters of the von Bertalanffy growth equation were K =0·21 cm year°1, L _∞=55·7 cm and t _o=-0·55 year for males and K =0·14 cm year⁻¹, L _∞=77·5 cm and t _o=-0·60 year for females. The difference in asymptotic length ( L _∞) between males and females may be because females postpone migration to achieve larger size for maximizing reproductive success.     

WALRUS LIFE-HISTORY MOVEMENTS RECONSTRUCTED FROM LEAD ISOTOPES IN ANNUAL LAYERS OF TEETH

Putative life history movements of male Atlantic walrus ( Odobenus rosmarus nsmarus ) harvested at Sanirajak in northern Foxe Basin, Canada, were reconstructed by comparing Pb isotopes in tooth cementum growth layer groups (GLGs) to those in whole-cementum samples from other walrus of Foxe Basin and nearby areas. The analyses yielded three life history patterns including one, "prodigal sons," not previously recognized in large mammals. Isotope ratios in GLGs of "local" Sanirajak walrus were relatively constant over time and similar to the whole-cementum data for most walrus caught in northern Foxe Basin. An "immigrant" walrus acquired the local signal only in young adulthood. In two "prodigal sons," GLG Pb isotope ratios changed significantly from the local signature after weaning but returned to it in late maturity, shortly before being harvested. Possible recruitment and dispersal areas for the two types of outliers included regions in northern and eastern Hudson Bay, some 300–600 km distant. The return of mature males to their natal group after associating with other groups for several years, as seen in some of the walrus, may be more common among mammals than previously thought. However, without a reconsttuctive technique such as employed here, it could only be detected through long-term studies of known individuals and all the relevant social groups.     

Haul-out and diving activity of male Atlantic walruses (Odobenus rosmarus rosmarus) in NE Greenland

During August-September 1989 and 1990, movements, haul out and dive activity of male Atlantic walruses ( Odobenus rosmarus rosmarus L.) were studied at a terrestrial haul-out site situated in an inshore foraging area in NE Greenland at 76± 30' N. Data were collected from direct observations of a group of about 50 males during August, including walruses that could be individually identified from natural markings, and from tracking of 8 adults equipped with satellite-linked radio transmitters during August-September. In both years, instrumented walruses hauled out for a total of 29.3% of the sampling time. In 1989, when ice floes were available for hauling out, the walruses spent 11% of the time on ice, whereas in 1990, when ice was absent from the study area, they only hauled out on land. Duration of haul-out periods, which did not differ between months or years, averaged 11 h (0.46 d) on ice (S.D. = 5.9, range: 1–29 h, n = 19 periods), and 38 h (1.6 d) on land (S.D. = 11.7, range: 13–64 h, n = 43). The walruses mainly hauled out during the afternoon and evening. Numbers hauling out on land during August were negatively correlated with wind direction, precipitation (rain) and wind-chill. In 1989, the duration of periods of absence from the terrestrial haul-out site (i.e. presumed foraging trips) averaged 206 h or 8.5 d (S.D. = 106.9, range: 48–412 h, n = 13), whereas, in 1990, such trips averaged only 81 h or 3.4 d (S.D. = 37.9, range: 24-156 h, n = 24), reflecting that walruses used the haul-out site more frequently when ice was absent. Direct observations of foraging walruses showed that they were submerged about 81% of the time.     

TRENDS IN AGE STRUCTURE AND PRODUCTIVITY OF PACIFIC WALRUSES HARVESTED IN THE BERING STRAIT REGION OF ALASKA, 1952–2002

Pacific walruses ( Odobenus rosmarus divergens ) are harvested by subsistence hunters in Alaska as they migrate north through the Bering Strait in the spring. Harvest records and biological specimens have been collected from the Bering Strait communities of Little Diomede, Gambell, and Savoonga since the 1950s. Harvest levels in the Bering Strait region peaked in the late 1980s and declined thereafter; however, there was considerable variation in the size and composition of the harvests among communities and over time. The relationships among characteristics of the community harvests and the presence of temporal trends were investigated using generalized linear models. The proportion of females in the catch increased over time in all three communities, while the proportion pregnant among harvested females declined over the range of sample years. The ages of harvested walruses increased over time in all three communities through the 1980s, after which trends in age stabilized or began to decline. The age of first reproduction was significantly older for females sampled in 1975–1985 than for females sampled between 1952 and 1962 or 1992 and 1998. Factors thought to have influenced the size and composition of the catch over the past 50 yr include hunter preferences, harvest management regimes, environmental conditions, and changes in the population itself.     

Early growth and juvenile population structure of lemon sharks Negaprion brevirostris in the Atol das Rocas Biological Reserve, off north‐east Brazil

Lemon sharks Negaprion brevirostris were sampled in the Atol das Rocas, a nursery area, on nine occasions from March 1999 to October 2003, during which 157 individuals were tagged and 35 were recaptured. The male : female sex ratio of captured individuals was 1 : 1·12. Mean ±  s . d . growth rates were 24·7 ± 3·4 cm year⁻¹ in total length ( L _T), 20·7 ± 3·2 cm year⁻¹ in fork length, and 19·5 ± 2·7 cm year⁻¹ in precaudal length. There was no significant difference in growth rates between males and females. Mean ±  s . d . increase in mass was 2565 ± 762 g year⁻¹. The von Bertalanffy growth parameters estimated by the Fabens method based on L _T were: k  = 0·077, L _∞ = 399·9 cm and t ₀ = −2·16. Despite the large variation of environmental conditions, particularly of tidal range and currents, and the lack of protective mangrove cover in the nursery area at Atol das Rocas, juvenile lemon sharks grew relatively faster than at other nurseries. Such rapid growth could be a response to abundant food availability or high risk of predation by adults that enter the nursery area.     

Sexual and geographical variation in life history parameters of the shorthorn sculpin

A total of 293 shorthorn sculpins Myoxocephalus scorpius from Tromsø, northern Norway, were sampled between November 1998 and April 1999 to determine sex, total length, age, growth, maturity and mortality. Females grew to larger sizes ( L _∞=26·9 v. 18·5 cm), matured later (2 v. 1 year of age) at larger size (maturation length=16 v. 14 cm L _T), and had lower instantaneous mortality rates (0·93 v. 1·20 year⁻¹) than males. The life history parameters of shorthorn sculpins in northern Norway were more similar to the parameters of short-lived central European populations than to the parameters of the long-lived population of Newfoundland. This study confirms that northern Norwegian shorthorn sculpins exhibit sexual dimorphism as in other shorthorn sculpin populations. The relationships between growth pattern, age at maturity and mortality rates observed in the Tromsø population and in other shorthorn sculpin populations, correspond well with the predictions from a published life history model.     

DURATION OF STEREOTYPED UNDERWATER VOCAL DISPLAYS BY MALE ATLANTIC WALRUSES IN RELATION TO AEROBIC DIVE LIMIT

During the breeding season from January to mid-April, adult male Atlantic walruses (Odobenus rosmarus rosmarus) dive repeatedly for an average duration of 4–6 min and give stereotyped underwater vocal displays. Between dives, they surface for 1–2 min, take 4–6 breaths, and give stereotyped vocalizations between breaths. Male walruses vocalize in the presence of groups of females and calves, young adult males, or by themselves as lone singers. This pattern is repeated throughout the breeding season and can be maintained for extended periods, sometimes exceeding 48 h. The prolonged underwater vocal displays of male walruses seem possible because the animals do not exceed the aerobic dive limit (ADL), estimated to be 9.8 min for a 1,100-kg animal, nor do they exceed the behavioral ADL of 7.9 min, determined from the histogram of dives for males singing alone. The number of breaths taken after dives and the postdive surface times remained fairly constant despite dive duration, suggesting that the walruses remained within their aerobic dive limits. The duration of most dives made by displaying males vocalizing alone during the breeding season, and dive duration of walruses feeding for protracted periods outside the breeding season, are both roughly half the estimated ADL.     

Towards a Better Understanding of the Effects of UV on Atlantic Walruses,Odobenus rosmarus rosmarus: A Study Combining Histological Data with Local Ecological Knowledge

Walruses, Odobenus rosmarus, play a key role in the Arctic ecosystem, including northern Indigenous communities, which are reliant upon walruses for aspects of their diet and culture. However, walruses face varied environmental threats including rising sea-water temperatures and decreasing ice cover. An underappreciated threat may be the large amount of solar ultraviolet radiation (UV) that continues to reach the Arctic as a result of ozone loss. UV has been shown to negatively affect whales. Like whales, walrus skin is unprotected by fur, but in contrast, walruses spend long periods of time hauled-out on land. In this study, we combined the results of histological analyses of skin sections from five Atlantic walruses, Odobenus rosmarus rosmarus, collected in Nunavik (Northern Quebec, Canada) with qualitative data obtained through the interviews of 33 local walrus hunters and Inuit Elders. Histological analyses allowed us to explore UV-induced cellular lesions and interviews with experienced walrus hunters and Elders helped us to study the incidences and temporal changes of UV-induced gross lesions in walruses. At the microscopic scale, we detected a range of skin abnormalities consistent with UV damage. However, currently such UV effects do not seem to be widely observed at the whole-animal level (i.e., absence of skin blistering, erythema, eye cataract) by individuals interviewed. Although walruses may experience skin damage under normal everyday UV exposure, the long-term data from local walrus hunters and Inuit Elders did not report a relation between the increased sun radiation secondary to ozone loss and walrus health.     

Age, growth and mortality of picarel, Spicara smaris L. (Pisces: Centracanthidae), from the eastern Adriatic (Croatian coast)

Age, growth and mortality of picarel Spicara smaris L. collected in the eastern middle Adriatic during April and May in 1999 were studied. The total lengths of individuals ranged from 6.3 to 19.8 cm, and the weight from 2.2 to 78.3 g. The slopes ( b values) of the total length–weight regressions indicated negative allometric growth for females ( b =2.8± 0.03) and isometric growth for males ( b =3.08±0.04). The oldest female and male were 4 and 6 years, respectively. The von Bertalanffy growth parameters for the sexes combined were ( L _∞=22.76 cm, K =0.277, t ₀=−0.793). The overall sex ratio was 1 : 4.1 in favour of females. Individuals larger than 17.9 cm were all males.     

Age estimation, growth, maturity and distribution of the roundnose grenadier from the Rockall trough

This paper summarizes the history of commercial exploitation of roundnose grenadier Coryphaenoides rupestris in the North Atlantic. Length frequencies of C. rupestris in 1993, from 400 to 1200 m on the slopes of the Rockall trough indicate a reduction since the 1970s in the modal length of fish found at 700–1000 m. Ages ranged from 2 to 50 years for males and 2 to 60 years for females, with most between 10–38 years. Females attained a greater asymptotic pre-anus length ( L _∞=19.5 cm) than males ( L _∞=15.5 cm) and had a greater weight for a given age (male W _∞=761g, female W _∞=1132g). This species may have a protracted spawning period. Using pre-anus lengths, 50% of male fish were mature at 10 cm (ages 8–10) while 50% of female fish were mature at 12 cm (ages 9–11). At the greatest depths sampled the length frequency of fish was bimodal with a hiatus between 9 and 11 cm (ages 8–12). Highest catch rates occurred on the Donegal slope in September at a depth of 800–1000 m.     

The Laptev Sea walrus Odobenus rosmarus laptevi: an enigma revisited

The walrus ( Odobenus rosmarus ) is in some current systematic schemes divided into three subspecies: O. r. rosmarus in the North Atlantic, O.   r. divergens in the North Pacific and O.   r. laptevi in the Laptev Sea. These three subspecies have been described as differing in body size, but the taxonomic status of O.   r. laptevi is disputed. The current study applies molecular and morphometric methods to assess the taxonomic status of O.   r. laptevi and to analyse the systematic and phylogeographic relationships between the three purported walrus subspecies. Tusk length and tusk circumference were measured from the few skulls available of O.   r. laptevi , and the obtained values were within the ranges reported for Pacific walruses. Thus, morphologically, subspecies status for O.   r. laptevi is not supported according to the Amadon–Mayr '75% rule'. Phylogenetic analyses and haplotype networks based on mitochondrial nucleotide sequence data of NADH dehydrogenase 1, 16S rRNA, cytochrome oxidase I and the d -loop of the control region of the historic O.   r. laptevi bone material and contemporary O.   r. rosmarus and O.   r. divergens showed that the Laptev Sea walrus groups with individuals from the North Pacific. Thus, the mitochondrial sequence data do not support the recognition of three walrus subspecies as reciprocally monophyletic evolutionary units with independent evolutionary histories. Only O.   r. rosmarus and O.   r. divergens meet this criterion with the present sampling. Accordingly, we recommend that Odobenus r. laptevi be abandoned and the Laptev walrus instead be recognized as the westernmost population of the Pacific walrus, Odobenus r. divergens. However, further research is recommended to assess whether the Laptev walrus could be considered as a significant unit in terms of conservation and management, since it is unique in several ecological parameters.     

The growth and mortality of larval herring, Clupea harengus L., in the River Blackwater Estuary, 1978–1980

Herring larvae were sampled in the Outer Thames Estuary and the River Blackwater Estuary in the springs of 1978, 1979 and 1980. Data were collected on larval stage, yolk sac and post yolk sac, larval length and total larval numbers. Newly hatched larvae were 6.8±0.5 mm long and the growth rate of yolk sac larvae was estimated at 0.18 mm d⁻¹ ( L = 6.8±0.186 t ). The growth rate of post yolk sac larvae increased to 0.43mm d⁻¹ ( L = 11±48.0±43 t ). Mortality estimates, derived from total numbers in the summed estuary segments, varied between the years and the cohorts within the year. In 1979 the mortality rates were 0.061 d⁻¹ and 0.074 d⁻¹ for the two cohorts. The mean size of the larval population was estimated at 2.48×10⁹ (1.63–3.77 × 10⁹) which agreed well with population size estimates from egg laying and from catch in numbers at age together with estimated fishing mortality rates.     

Accumulation of PSP toxins in Atlantic mackerel: seasonal and ontogenetic variations

Atlantic mackerel Scomber scombrus are known to be lethal vectors of paralytic shellfish poisoning (PSP) toxins to predators. To elucidate dynamics of PSP toxin accumulation in this species, mackerel were sampled in the Gulf of St Lawrence from May to October 1993. Mackerel appear to retain toxins (saxitoxin, gonyautoxins 2 and 3) year-round. The toxin content of the liver, as determined by high performance liquid chromatography, increased significantly with fish age ( r² =0.40) and length ( r² =0.52), suggesting that mackerel progressively accumulate PSP toxins throughout their life. The toxin content of the liver also increased significantly during the summer feeding sojourn in the Gulf of St Lawrence. Comparison of profiles of saxitoxin derivatives indicated that zooplankton were the likely source of PSP toxins found in mackerel. The mean ± S.D toxin content was 17.4 ± 10.6 nmol liver⁻¹ and the mean ± S.D. PSP toxicity was 112.4 ± 67.0 μg saxitoxin equivalents 100 g⁻¹ liver wet weight ( n =247).