Hematology and serum chemistry of harp (Phoca groenlandica) and hooded seals (Cystophora cristata) during the breeding season, in the Gulf of St. Lawrence, Canada

Standard hematologic and serum chemistry parameters were determined from 28 harp seals (Phoca groenlandica) and 20 hooded seals (Cystophora cristata) sampled from 6 March 2001 to 13 March 2001 during the breeding season. Whole blood was collected immediately postmortem from harp seal mother-pup pairs and from six hooded seal pups, and from live-captured adult hooded seals and three hooded seal pups; blood was analyzed within 24 hr at a local human hospital. A certified veterinary laboratory validated subsamples of whole blood and analyzed all serum chemistry parameters. Significant interlaboratory differences in mean values of packed cell volume (PCV) and mean cell volume (MCV) were found. Significant differences were found between samples from the five seal groups (adult male hooded seals, lactating female hooded seals, unweaned hooded seal pups; lactating female harp seals, and unweaned harp seal pups) for hematology and most serum chemistry parameters. In general, age-class influenced mean values of PCV, hemoglobin (HB), red blood cell (RBC) counts, MCV, mean cell hemoglobin (MCH), mean cell hemoglobin concentration (MCHC), and nucleated red blood cell (NRBC) counts per 100 leucocytes, but most age-related variations were species specific. Harp seal pups had significantly lower mean values of HB, PCV, MCH, and MCHC than did other seal groups, and significantly lower mean RBC counts than did hooded seal pups. Mean NRBC counts per 100 leukocytes were more than three times higher in harp seal pups than in hooded seal pups, but this difference was not statistically significant. Mean MCV were significantly lower in harp and hooded seal pups compared to those of adult harp and hooded seals. Differences in hemograms between pup species were likely because of the precocious development of hooded seal pups, which are weaned within 4 days, compared to 12 days for harp seal pups. Among adult seal groups, male hooded seals had significantly higher mean values of PCV and HB than did female harp and hooded seals, and significantly higher mean RBC counts than did adult female hooded seals. Among adult females, mean values of MCH and MCHC were statistically higher in hooded seals than in harp seals. Adult female harp and hooded seals did not differ significantly in other RBC parameters and mean leukocyte counts. Mean values of glucose, blood urea nitrogen, total bilirubin, alanine aminotransferase (ALT), alkaline phosphatase (ALP), total protein, and albumin showed species-specific variations between adults and pups. Except for ALP, few significant differences in mean enzyme activities of aspartate aminotransferase (AST), ALT, creatine kinase and gamma-glutamyltransferase were found between seal groups. Mean concentrations of electrolytes (calcium, phosphorus, sodium, potassium, chloride, magnesium, and total carbon dioxide) varied with age class, but variations in potassium and magnesium were species specific. Harp seal pups had significantly higher mean phosphorus and potassium levels compared to other seal groups.     

Multi-decadal trends in biomarkers in harp seal teeth from the North Atlantic reveal the influence of prey availability on seal trophic position

Arctic food webs are being impacted by borealisation and environmental change. To quantify the impact of these multiple forcings, it is crucial to accurately determine the temporal change in key ecosystem metrics, such as trophic position of top predators. Here, we measured stable nitrogen isotopes (δ¹⁵N) in amino acids in harp seal teeth from across the North Atlantic spanning a period of 60 years to robustly assess multi-decadal trends in harp seal trophic position, accounting for changes in δ¹⁵N at the base of the food web. We reveal long-term variations in trophic position of harp seals which are likely to reflect fluctuations in prey availability, specifically fish- or invertebrate-dominated diets. We show that the temporal trends in harp seal trophic position differ between the Northwest Atlantic, Greenland Sea and Barents Sea, suggesting divergent changes in each local ecosystem. Our results provide invaluable data for population dynamic and ecotoxicology studies.     

Zooarchaeological analysis of prehistoric vertebrate exploitation at the Grand Bay Site,Carriacou, West Indies

This paper discusses zooarchaeological analysis of vertebrate faunal specimens from Grand Bay, a Ceramic Age (ca.ad 400–1300) site on the island of Carriacou in the Grenadines, West Indies. Using faunal data to assess subsistence patterns of vertebrate exploitation during late site occupation, we can begin to better understand Grand Bay procurement strategies and coral reef exploitation. Preliminary zooarchaeological results imply that Grand Bay vertebrate exploitation emphasized marine resources over terrestrial resources, with particular emphasis on coral reef habitats and fish. The faunal data are discussed in relation to common patterns of prehistoric vertebrate exploitation in the Caribbean and fishing strategies. The Grand Bay faunal sample also provides a foundation from which to formulate future research foci and question zooarchaeological approaches to understanding prehistoric coral reef exploitation in the Caribbean.     

Le Paléolithique moyen de l'Abric Romaní. Comportements écosociaux des groupes néandertaliens

The Abric Romaní is one of the most important Upper Pleistocene sequence of the Iberian Peninsula, with archaeological levels belonging to the Middle Paleolithic and with one of the earliest Aurignacian samples from Western Europe. Research has been focused on the interpretation of patterns of manage and consume of natural resources and the spatial organization of the activities inside the site, and for that reason, the fieldwork strategy is lead to excavate the whole surface of the site. The results of more than 25 years of research on the Abric Romaní conclude that Neanderthal communities settled in the site forming domestic groups around hearths. Depending on the intensity of occupation, related to the length of occupation and with the number of people of these groups, this spatial domestic structure evolves forming the variability spectrum of settlement impacts in the site. In the same way, the manage of lithic and faunal resources increases its variability following the intensity of occupation, using clears technical patterns of lithic production and for catching and processing faunal preys. This variability from behaviors well established is interpreted like the way for adapting new historical and environmental situations.     

Cenozoic radiolarian paleobiogeography: Implications concerning plate tectonics and climatic cycles

A preliminary study of the paleobiogeographic patterns of radiolarian facies during the Paleogene and subsequent time shows that:(1) Through time radiolarian assemblages display distinct faunal provincialism reminiscent of modern faunal distributions correlated with planetary temperature gradients and surface oceanic conditions. The equatorial—tropical radiolarian fauna extended apparently unrestricted across the Pacific Ocean, the Caribbean Sea and the Atlantic Ocean through Early Miocene time. In the Caribbean Sea and the Atlantic Ocean, radiolarians reached their maximum abundance in the Eocene and Oligocene. Subsequently, they gradually declined to virtual disappearance in these areas in the early Miocene. Their Pacific counterparts remained practically undisturbed, except that post early Miocene assemblages there showed a marked trend toward decreasing test thickness. This trend has since been a worldwide characteristic of Neogene radiolarian assemblages and their modern equivalents. It is postulated that the disappearance of radiolarians in the Carribean Sea and the Atlantic Ocean at the end of the Paleogene is related to the onset of the emergence of the isthmus of Panama which interrupted the preexisting oceanic circulation between the Pacific and Atlantic Oceans.(2) Throughout the Paleogene there have been marked sequential fluctuations in the radiolarian assemblages of the Caribbean Sea which indicate intermittent incursions of higher-latitude fauna in this area. Associated with the faunal fluctuations are cyclic variations in the total carbonate of the sediment with patterns also comparable in duration to Pleistocene carbonate cycles in the equatorial Pacific known to have been induced by climatic changes. Based on similarities with Pleistocene climatic cycles in the equatorial Pacific and elsewhere, it is surmised that the faunal and lithologic fluctuations observed in Paleogene radiolarian sediments were also induced by the biologic and physico-chemical processes associated with worldwide changes in the climatic conditions of that time.     

What's in a beach? Soil micromorphology of sediments from the Lower Paleolithic site of 'Ubeidiya, Israel

A micromorphological study of archaeological sediments from the early Pleistocene site of 'Ubeidiya (Jordan Valley, Israel) was conducted to provide microenvironmental detail for the hominin occupation contexts and investigate site formation issues. Previous research shows that the hominin groups occupied the marshes and pebbly beaches at the shores of a lake during a regressive period, but given that some portions of the lithic and faunal assemblages are abraded and others fresh, there remains a question of whether the archaeological assemblages are in situ or reworked, and if reworked, by what mechanisms and from where. The rates of sedimentation within the regressive cycle, by which we can learn about the frequency and duration of exposed surfaces amenable for hominin occupation is also unknown. Finally, the artificial nature of some of the pebbly layers has been questioned. The micromorphological analysis yielded the identification of twelve microfacies; the majority of these represent fluvially derived floodplain soils or distal mudflow deposits, and a minor number are sediments of lacustrine origin: mudflats and shallow subaqueous sediments. These represent the natural habitats of the 'Ubeidiya hominins and might serve as a reference to similar contexts of other early hominin sites. The sedimentary model proposed here entails the rapid deposition of fluvially derived low-energy sediments at and around the shoreline, followed by prolonged periods of exposure, during which surfaces stabilized within a relatively wet, marshy environment. This interpretation suggests that the abraded portions of the archaeological assemblages are a result of prolonged surface exposure rather than high-energy transport from a distant source or to wave reworking at the shoreline, and supports the consideration of these assemblages as archaeological palimpsests, with locally reworked fresh and abraded elements. No micromorphological evidence supporting anthropogenic agency in the formation of the pebbly layers was found. The entire regressive cycle entailed unvarying climatic conditions with seasonal fluctuations and episodic lacustrine incursions, and with a trend towards arid conditions in the end.     

Monoclonal antibody to alkaline phosphatase from the intestinal mucosa of the harp seal, Phoca groenlandica.

1. Hybridoma secreting a monoclonal antibody APP.1 to the harp seal alkaline phosphatase (A1Ph) was obtained by fusing murine myeloma Sp 2/0 cells with the splenocytes of BALB/c mice immunized with purified isozyme K. 2. The antibody has no effect on the enzyme activity and shows a high affinity for harp seal A1Ph (KD = 8.5 x 10(-10) M). The antibody has similar affinities for the AlPh of harp seal, fur seal, common seal and deer. 3. The antibody APP.1 was coupled to Sepharose and employed in chromatographic purification of the harp seal intestinal AlPh. Alkaline phosphatase isolated on this immunosorbent has a spec. act. of 20,800 units per mg of protein. 4. The antibody-enzyme complex gives an excellent immunocytochemical labeling of tissue sections, cell cultures and smears.     

UK intertidal foraminiferal distributions: implications for sea-level studies

Foraminiferal assemblages have been collected from ten intertidal study areas situated on the east, south and west coasts of the UK. The assemblages display a vertical zonation which indicates that the distribution of foraminifera in these intertidal environments is usually the direct function of altitude with the duration and frequency of intertidal exposure the most important factors. Multivariate analyses separate foraminiferal assemblages into two faunal zones: a high- and middle marsh zone consisting of differing abundances of Jadammina macrescens, Trochammina inflata and Miliammina fusca; and a low-marsh and tidal flat zone dominated by calcareous foraminiferal species, notably Elphidium williamsoni, Haynesina germanica and Quinqueloculina spp. These faunal zones are similar to those in other mid-latitude, cool temperate intertidal environments although there are spatial and temporal variations between areas. The altitudinal ranges of the faunal zones are employed to identify the vertical relationship of the local environment in which the assemblage accumulated to a reference tide level.     

A PREHISTORIC BREEDING POPULATION OF HARP SEALS (PHOCA GROENLANDICA) IN THE BALTIC SEA

The pelagic and gregarious, low Arctic harp seal ( Phoca groenlandica ) is the most common seal species in most refuse faunas from coastal hunter-gatherer sites dating from the late Atlantic to the early Subboreal period ( ca. 4000-2000 cal B. C.) in the Baltic Sea. Our main objective was to examine the migration contra breeding population hypotheses regarding the Baltic harp seals. Analyses of epiphyseal fusion data and osteometry of archeological harp seal remains from 25 dwelling-sites suggest that a local breeding population established itself in the early Subboreal period. In the Middle Neolithic the rookery possibly was situated in the Baltic proper, south of Aland and west of Gotland. The mean adult size of the Baltic harp seals decreased, suggesting minimal genetic exchange with the north Atlantic Ocean population. Genetic drift, interspecific competition, and over-hunting by humans are all factors likely to have contributed to the eventual extinction of harp seals in the Baltic Sea.     

DIGESTIBILITY OF CRUSTACEANS and CAPELIN IN HARP SEALS (PHOCA GROENLANDICA)

Gross energy intake (GEI), apparent digestible efficiency (%DE), and body-mass change were studied both in recently weaned and in subadult harp seals fed capelin ( Mallotus villosus ) and crustaceans ( Parathemisto libellula and Thysanoessa sp.) ad lib. The median %DE of Parathemisto libellula (with a wet-weight energy density (ED) of 3.9 kJ/g) and Thysanoessa sp. (ED: 2.3 kJ/g) was 81% and 83%, respectively, while the median %DE of capelin (ED: 8.6 kJ/g) was 94% and 93% for subadult harp seals and one harp seal pup, respectively. No correlation between age and %DE was obtained (P > 0.05). Despite ad lib. access to the food, the harp seal pups were unable to maintain body mass while on a crustacean diet, and it is therefore suggested that harp seal pups most likely must supplement their crustacean diets with fish to maintain growth.     

Dung beetle assemblage structure across the aridity and trophic resource gradient of the Botswana Kalahari: patterns and drivers at regional and local scales

Understanding pattern and process at both regional and local scales is important for conservation planning although such knowledge of insects is frequently lacking. To assess patterns along a regional gradient of increasing aridity and diminishing food resources in the Botswana Kalahari, Scarabaeine dung beetles were sampled quantitatively using four dung types at three local sites in six regional areas. At regional scale, factor analysis of species abundance extracted a maximum of six factors, each dominated by a single area. Therefore, the statistical significance of regional spatial variation far outweighed that of dung type association. At local scale, six factor analyses of species abundance extracted from four to six factors. The importance of local dung type associations was relatively high but diminished with increasing local spatial heterogeneity. At regional scale, hierarchical analysis of oblique factors divided assemblages into unique local and shared regional components. Primary extended factors accounted for 40–50 % of unique local faunal composition in five out of six areas. Two secondary extended factors showed either high shared proportional contribution to regional assemblage structure in the northeast with a steep decline to the southwest, or an opposite trend. Their point of intersection was consistent with a boundary zone between mesic northeast and arid southwest faunal components in the central Kalahari. Despite some inconsistencies in rank position between regression methods, rainfall, temperature, and mammal density/diversity were the strongest influences on regional patterns defined by secondary factors. Patterns are discussed according to conservation and changes in land usage around reserves.     

From carcass to cave: large mammal exploitation during the Aurignacian at Vogelherd, Germany

Recent results from the zooarchaeological analysis of faunal remains from Vogelherd Cave, southwestern Germany, provide new insight into the subsistence behavior of early modern human groups during the Aurignacian. The results presented here represent the first comprehensive study of the archaeofauna from this site. Several episodes of occupation are inferred at this site, taking place primarily between 31 and 32 ka. Although a wide spectrum of Pleistocene mammals is represented in the Aurignacian at Vogelherd, reindeer (Rangifer tarandus) and horse (Equus ferus) were the primary prey taxa, and they are the most appropriate data sets with which to understand human subsistence on an intrasite level. Hunting of both taxa took place during the late summer and fall, coinciding with reindeer migrations and local abundance of horses. Complete or nearly complete prey carcasses were then transported from the kill locations to the cave for processing. This study shows that Vogelherd was a preferred locale of Aurignacian groups for a broad range of activities, including the time- and labor-intensive exploitation of ungulate prey for meat, marrow, and fat resources, as well as the production and maintenance of artifacts such as figurative artwork, personal ornaments, bone and ivory armatures, and lithic tools. With its rich faunal and artifact assemblages, the Aurignacian deposit at Vogelherd provides a wealth of information on this critical period of the early Upper Paleolithic, when cultural innovations were flourishing.     

DIET OF WEANED PUPS AND SEASONAL VARIATIONS IN BODY CONDITION OF JUVENILE BARENTS SEA HARP SEALS PHOCA GROENLANDICA

Data on blubber depth, body condition, and diet were collected from young of year and juvenile harp seals ( Phoca groenlandica ) in the Barents Sea in the periods February-April and June-October 1990–1997. Harp seal pups feed independently shortly after weaning. In the southern parts of the Barents Sea the diet of harp seal pups consisted mainly of euphausiids ("krill") Thysanoessa sp. and amphipods of the genus Parathemisto . The change in body condition of harp seal pups between weaning (mid-March) and June indicated that considerable amounts of blubber energy, deposited during suckling, were mobilized. This suggests that the early food intake of weaned pups was insufficient to meet the energy requirements. The pattern of low spring and earlysummer body condition, followed by a subsequent rapid improvement in condition during late summer and autumn appeared to occur not only among pups-of-the-year, but also in one- and two-year old harp seals.     

Seasonal energetics of northern phocid seals

The metabolic rate of harp (Pagophilus groenlandicus), harbor (Phoca vitulina), and ringed seals (Pusa hispida) was measured at various temperatures in air and water to estimate basal metabolic rates (BMRs) in these species. The basal rate and body composition of three harp seals were also measured throughout the year to examine the extent to which they vary seasonally. Marine mammalian carnivores generally have BMRs that are over three times the rates expected from body mass in mammals generally, both as a response to a cold-water distribution and to carnivorous food habits with the basal rates of terrestrial carnivores averaging about 1.8 times the mean of mammals. Phocid seals, however, have basal rates of metabolism that are 30% lower than other marine carnivores. Captive seals undergo profound changes in body mass and food consumption throughout the year, and after accounting for changes in body mass, the lowest rate of food intake occurs in summer. Contrary to earlier observations, harp seals also have lower basal rates during summer than during winter, but the variation in BMR, relative to mass expectations, was not associated with changes in the size of fat deposits. The summer reduction in energy expenditure and food consumption correlated with a reduction in BMR. That is, changes in BMR account for a significant portion of the seasonal variation in energy expenditure in the harp seal. Changes in body mass of harp seals throughout the year were due not only to changes in the size of body fat deposits, but also to changes in lean body mass. These results suggest that bioenergetics models used to predict prey consumption by seals should include time-variant energy requirements.     

Comparaisons taphonomiques de sites magdaléniens du Bassin parisien : les faunes de Ville-Saint-Jacques (Seine-et-Marne) et Étiolles (Essonne)

The preservation of faunal remains in the Magdalenian sites of the Paris Basin only occurs for one third of the cases. This taphonomic bias is very likely in relation with the open-air locations where the majority of regional sites have been discovered. Nevertheless, the Magdalenian settlement shows a certain variability across the landscape: lowland sites, hillside sites, rock shelter sites. This paper aims to highlight the potential interactions between such physical context variability and the taphonomical features of Magdalenian faunal preservations. The issues presented here mainly focus on the comparison of the two faunal remains’ assemblages of Étiolles (a lowland site of the Seine valley) and Ville-Saint-Jacques (hill side site closed to the Seine valley). These faunal assemblages have been studied using the same set of qualitative and quantitative criteria in order to facilitate direct comparisons. This paper must be viewed as the first step of a more systematical survey of Magdalenian faunal assemblages in the Paris Basin. Finally, the main goal of this survey aims to point out the respective zooarchaeological potential of such sites, and by the way, to reach their global socioeconomic significations.     

New data on Late Pleistocene small vertebrates from northern France

Quaternary small vertebrate assemblages from northern France, and more generally from northwestern Europe, are poorly recognized, and studied less than those from more southern, Mediterranean areas. However, important sites of human occupation are often found in these northern areas, with significant issues related to the occupation of these sites by Neanderthals and previous humans, as well as faunal dynamics under climatic pressure. In this paper, we present preliminary results from the study of small vertebrate assemblages from two Late Pleistocene sites of northern France: Mutzig (Alsace) and Le Rozel (Normandy). Both are ancient rock-shelters that have been excavated recently and have yielded evidence of Neanderthal occupation, but in very different contexts: Le Rozel is located in a coastal area of northwestern France contemporaneous with a temperate period, while Mutzig is located at the foot of the Vosges mountains in northeastern France and was occupied during a cold period. Consequently, even if these two sites are chronologically close to each other, differences in faunal composition are observed and discussed in relation to the geographic, climatic and biochronological context, in comparing the results from small vertebrates with those from other disciplines (numerical dating, large mammals).     

The effects of climate change on harp seals (Pagophilus groenlandicus)

Harp seals (Pagophilus groenlandicus) have evolved life history strategies to exploit seasonal sea ice as a breeding platform. As such, individuals are prepared to deal with fluctuations in the quantity and quality of ice in their breeding areas. It remains unclear, however, how shifts in climate may affect seal populations. The present study assesses the effects of climate change on harp seals through three linked analyses. First, we tested the effects of short-term climate variability on young-of-the year harp seal mortality using a linear regression of sea ice cover in the Gulf of St. Lawrence against stranding rates of dead harp seals in the region during 1992 to 2010. A similar regression of stranding rates and North Atlantic Oscillation (NAO) index values was also conducted. These analyses revealed negative correlations between both ice cover and NAO conditions and seal mortality, indicating that lighter ice cover and lower NAO values result in higher mortality. A retrospective cross-correlation analysis of NAO conditions and sea ice cover from 1978 to 2011 revealed that NAO-related changes in sea ice may have contributed to the depletion of seals on the east coast of Canada during 1950 to 1972, and to their recovery during 1973 to 2000. This historical retrospective also reveals opposite links between neonatal mortality in harp seals in the Northeast Atlantic and NAO phase. Finally, an assessment of the long-term trends in sea ice cover in the breeding regions of harp seals across the entire North Atlantic during 1979 through 2011 using multiple linear regression models and mixed effects linear regression models revealed that sea ice cover in all harp seal breeding regions has been declining by as much as 6 percent per decade over the time series of available satellite data.     

Radiocarbon and stable isotope investigations at the Central Rhineland sites of Gönnersdorf and Andernach-Martinsberg, Germany

The late glacial open-air sites of Gönnersdorf and Andernach-Martinsberg in the German Central Rhineland are well known for their Magdalenian occupation and activities. The latter site also produced evidence for a younger, Final Palaeolithic occupation of the locality by people of the Federmessergruppen. Both sites are particularly well preserved, largely due to their burial beneath volcanic deposits of the late glacial Laacher See eruption. We conducted a program of AMS radiocarbon dating and stable isotope analyses with the aim of improving understanding of the chronological history and ecological setting of the two sites.Previously published radiocarbon dates appeared to indicate that the earliest Magdalenian occupation at Gönnersdorf fell around 12,900 uncalibrated ¹⁴C yr BP, while the earliest occupation at Andernach may have been more than 500 radiocarbon years earlier. The AMS determinations presented here revise this impression and suggest that the onset of occupation at the two sites was in fact simultaneous and prior to the warming of Greenland Interstadial GI 1e. At Gönnersdorf, a chronological hiatus exists between the main Magdalenian faunal assemblage and mega-faunal remains interpreted as collected sub-fossil material. At Andernach-Martinsberg, there is a clear chronological hiatus between the Magdalenian occupation and subsequent Federmessergruppen activities at the site. However, an intermediate radiocarbon date on an atypically preserved horse bone is suggestive of ephemeral human visits to the site between these well demonstrated phases. A date of similar age on an elk bone from Gönnersdorf may indicate broadly contemporaneous human presence at Gönnersdorf too.Stable isotope analysis of faunal remains from Gönnersdorf and Andernach-Martinsberg was conducted with the aim of both reconstructing and comparing local environmental conditions at the two sites, and also potentially identifying subtle variations in the chronological development of the two sites not detectable at the level of precision of current radiocarbon dating techniques. No spatial trends in the faunal isotope signatures were observed within each site. In the case of samples with both radiocarbon and isotope data, no chronological pattern was observed for the isotope results. The Magdalenian faunal isotope signatures at the two sites resembled each other, suggesting comparable climatic and environmental conditions. The faunal δ¹³C signatures at Gönnersdorf and Andernach-Martinsberg were similar to those at contemporary European sites. While the faunal δ¹⁵N values were similar to those at contemporary sites in Germany, the UK, and Belgium, they were lower than those from the South of France. This difference in δ¹⁵N values is thought to relate to regional differences in the timing of changes in soil and plant nitrogen cycling in response to ameliorating climatic conditions.     

Underwater calling rates of harp and Weddell seals as a function of hydrophone location

Stereo recordings of harp seal (Pagophilus groenlandicus) and Weddell seal (Leptonychotes weddellii) underwater vocalizations were made near breeding groups using separations between two hydrophones of 159–339 m. Within a large herd, harp seal call numbers varied slightly between channels. Counts of Weddell seal calls were higher near a small herd on the ice than 159–180 m away. Repeat counts of harp seal calls by a single observer differed significantly. Source levels of Weddell seal calls varied and higher amplitude calls would be detected up to 2 orders of magnitude farther away than quieter calls. Hydrophone location (especially near small groups), observer variability and call source level differences will bias the use of monitoring underwater seal vocalizations to index locations, population size or underwater behaviours. Using hydrophone arrays and multiple observers may mitigate these problems. Accepted: 15 September 2000