Mechanism of an epibiont burden: Crepidula fornicata increases byssus thread production by Mytilus edulis

Epibionts of mussels can have detrimental effects on their basibionts,such as reduced growth rates, lower fecundity, increased mortalityand an enhanced risk of dislodgement of the overgrown bivalvesdue to stronger hydrodynamic forces. In blue mussels Mytilusedulis, the epibiotic American slipper limpet Crepidula fornicatareduces growth and survival. In a field experiment we testedthe hypothesis that an enhanced byssus thread production withhigh energetic costs for the mussels due slipper limpet epibiontsis the underlying mechanism for the epibiont burden. Byssusthread production in overgrown mussels was twice as high asin unfouled M. edulis (11 ± 0.9 and 5.4 ± 0.6byssus threads/mussel/day, respectively). A control experimentrevealed intermediate byssus thread production (8.4 ±0.8 byssus threads/mussel/day) in mussels cleaned of C. fornicataat the beginning of the experiments, indicating that C. fornicatais responsible for the effects observed. We conclude that increasedbyssus production in fouled M. edulis is a functional responseto higher drag caused by epibionts and that it is associatedwith increased energy expenditure that reduces allocation ofresources for other processes such as growth, reproduction andsurvival. Such indirect effects of epibionts, mediated by anenhanced byssus production, may be widespread in byssus-producingbivalves, with important implications for their population dynamics. (Received 12 January 2006; accepted 21 November 2006)     

Demography and reproductive parameters of a free-ranging group of Japanese macaques (Macaca fuscata) at Katsuyama

Demographic and reproductive data were analyzed for a period of 28 years in the females of a free-ranging group of Japanese macaques at Katsuyama, Okayama Prefecture, Japan. The overall mean, age-specific fecundity rates were 5.43% for 4-year-olds and 41.86% for 5-year-olds, increasing to a peak of 66.67% for 13-year-olds. Fecundity remained relatively high (52.31–54.24%) in 16–19-year-olds, but decreased sharply (45.45–17.86%) in 20–23-year-olds, and became very low in 24–26-year-olds. Females aged 27 years or more did not produce infants. The average age at first birth was 5.41 years. Births peaked in mid-May. The timing of the first births each year remained essentially unchanged during the study period, whereas the timing of the median and last births shifted towards the later part of the season. The mean interbirth interval for all females was 1.56 years. The value was 1.54 years for multiparous females and 1.29 years for females following infant loss. These intervals were significantly shorter than those for primiparous females, and females with surviving infants. The overall mean infant mortality within the first year of life was 10.2%. The value was 8.6% for 10–14-year-olds, and 7.5% for 15–19-year-olds. The timing of birth differed among the four female matrilineal dominance rank-classes. The female fecundity rates increased as a function of matrilineal dominance rank. It is suggested that all demographic and reproductive data should be analyzed in detail with respect to the group's history.     

Modelled population growth based on reproduction differs from life tables based on age determination in Danish raccoon dogs ( Nyctereutes procyonoides )

The population of raccoon dogs (Nyctereutes procyonoides) in Denmark has increased rapidly from 1995 when the first was recorded until today where 3291 raccoon dogs are trapped, shot by hunters or road killed. The aims of this study are to present the first data on reproduction and life tables of raccoon dogs in Denmark and to compare mortality from modelled life tables with game bag records and sampled raccoon dogs in different age groups. In this study, the uteri of 89 adult females (> 10 months) were examined for placental scars (PSC), and 561 individuals (289 males, 272 females) were aged using pulp cavity width and dental lines in canine teeth. The litter size of raccoon dogs in Denmark is to date the largest litter size recorded in the wild (mean ± SE) 10.8 ± 0.4, range 1–16 pubs and fecundity 8.4 ± 0.6 pubs. The percent-reproducing females are 78–83%, based on dark and all PSC, respectively. A significant difference was found between the proportion of individuals composing the different age groups based on age determination of individuals collected (Ntage) and the modelled number of individuals in age groups based on fecundity and different mortality rate (Ntmodel), X2 = 8, p < 0.05. The discrepancy between the relatively high reproduction and lifetables may be due to older and more experienced animals that avoid culling. A low population density in a newly founded Danish population of raccoon dogs, together with a milder climate where raccoon dogs can forage during the winter, may cause an exceptionally high reproduction in Danish raccoon dogs.     

The effects of spiromesifen on life history traits and population growth of two-spotted spider mite (Acari: Tetranychidae)

Laboratory bioassays were conducted to evaluate the effects of spiromesifen on gross fecundity, gross fertility, net fertility and population growth of two-spotted spider mite (Tetranychus urticae Koch) after treatments with four acaricide concentrations: 180 mg/l, i.e. maximum recommended concentration for use in glasshouses against spider mites, 18, 1.8, and 0.18 mg/l, i.e. concentration discriminative for eggs and immatures in preliminary studies which produced 100% mortality of these stages. Quiescent female deutonymphs were treated in the first assay, and young pre-ovipositing females in the second and third, in which exposure lasted 6 h and 20 h, respectively. In the first assay, the 180, 18, and 1.8 mg/l concentrations significantly reduced gross fecundity (61–85%), gross fertility (64–87%) and net fertility (85–94%) of the surviving females. In the second one, only the highest concentration achieved a significant statistical reduction in gross fecundity (52%), gross fertility (67%) and net fertility (84%). In the third assay, fecundity and fertility reduction under the two highest concentrations was 98–99% and 93–98%, whereas it was 50–74% under the 1.8 mg/l concentration, and statistically different from control values. In all three trials, treatments with 180, 18, and 1.8 mg/l concentrations significantly reduced the instantaneous rate of increase. In the third assay, treatments with the two highest concentrations caused population decline. Sublethal activity of the 0.18 mg/l concentration was not found in any assay to be statistically significant. Sublethal effects of spiromesifen and its impact on T. urticae management are discussed.     

Taxonomic Status and Reproduction Dynamics of the Non-Indigenous Cercopagis in the Gulf of Riga (Baltic Sea)

Overwintering population of Cercopagis is represented by resting eggs which hatch generally in May. Although representatives of the first parthenogenetic generation (the spring form individuals) differ morphologically from individuals of the subsequent generations (the summer form individuals) and could be keyed to a separate species – Cercopagis (Apagis) ossiani – our analysis confirm that there is only one Cercopagis species in the Gulf of Riga: Cercopagis pengoi. Notable seasonal dynamics was observed for the gamogenetic mode of reproduction, being strongly associated with the total population density. Gamogenetic females develop since the end of June and reach the maximum absolute abundance in July whereas their proportion is generally the highest in fall. The gamogenetic absolute fecundity drops to the lowest level late July but increased afterwards until the late August. One resting egg was, an average, found in 45.4%, two in 53.4% and three in 1.2% of females. Parthenogenetic fecundity was significantly higher in May–June compared to other months studied. Brood pouch of parthenogenetic females was found to contain 11.6 ± 1.0 and 10.2 ± 0.3 embryos in the spring and summer form individuals, respectively. Variation in the two modes of reproduction and fecundity is probably solely not controlled by temperature, but also by food availability and population density.     

Size composition and reproductive biology of the ornate jobfish <Emphasis Type="Italic">Pristipomoides argyrogrammicus</Emphasis> (Lutjanidae) off Ishigaki Island,Okinawa

The ornate jobfish Pristipomoides argyrogrammicus is an important lutjanid species for fisheries in the Okinawan region of Japan. The present study estimated the size composition and reproduction of this species in the waters around Ishigaki Island, Okinawa Prefecture. The length–frequency distribution indicated that males grow larger than females. The fork length (FL; mm)–whole body weight (BW; g) relationship and FL–total length (TL) relationship were as follows: BW = 1.048 × 10⁻⁵ FL^3.121 and TL = 1.101 × FL + 2.196, respectively. The main spawning season was estimated as between April and August, since higher gonadosomatic index (GSI) values were found for both sexes, and matured oocytes were observed in females during these months. The developmental stage of ovaries correlated with the GSI of females. Fecundity ranged from 9,530 to 98,260 oocytes in fish of 177.0 to 278.0 mm FL, and the FL–fecundity relationship was as follows: fecundity = 9.525 × 10⁻⁸ FL^4.903.     

Metabolic cost of first reproduction in young female European roe deerCapreolus capreolus

In female roe deerCapreolus capreolus (Linnaeus, 1758), like in several deer species, first reproduction occurs before they have reached their full body size. This study quantifies, in young females, the energetic costs induced by the contemporaneously occurring events: growth and first reproduction. Resting metabolic rate and body mass were measured in young primiparous females from first mating to 4 months after parturition, and compared to values measured in fully-grown adult multiparous females. Throughout the 10-month period from mating to fawning, body mass increased in yearlings. Prior to the blastocyst implantation (month −5 before parturition) young females were lighter than adult females whereas after parturition they had the same body mass. Our results suggest that body growth was reinitiated during pregnancy in primiparous females. From mating to fawning, except in the first part of pregnancy, mass-specific metabolism was higher in primiparous females than in multiparous individuals indicating the occurrence of an additional cost due to growth in young females. The depressed level observed at the beginning of gestation could allow the resumption of growth at lower cost. Thus, the allocation of resources to both reproduction and growth was not detrimental to first reproduction in young female roe deer under experimental conditions withad lib feeding.     

On reproduction of longfin thornyhead <Emphasis Type="Italic">Sebastolobus macrochir</Emphasis> (Scorpaeniformes: Sebastidae) in waters off the eastern coast of the Kurils

According to data obtained off the eastern coast of the Northern Kurils in October–December 1998, the body length of spawning females of the longfin thornyhead Sebastolobus macrochir is 26–41 cm (on average, 31.5 ± 0.27). Mass spawning is recorded in females 28 cm in length and longer. The individual absolute fecundity of females is 69355-411710 (on average, 189086 ± 13662) eggs. The reproductive potential of females of the longfin thornyhead is estimated as 189 million eggs. It is supposed that all females participate in spawning.     

On the biology of the Japanese swimming crab <Emphasis Type="Italic">Charybdis japonica</Emphasis> (Portunidae) in waters of primorye at the northern boundary of their range

Data on the biology of the Japanese swimming crab, such as sexual dimorphism, size and weight structure, sex ratio, allometric growth, and fecundity were obtained in Sukhodol Bay (Ussuri Bay). The maximum carapace width was 116 mm and the largest weight was 340 g in males and 107 mm and 210 g, respectively, in females. The female to male ratio was 1.0: 2.4. The mean number of eggs in a clutch was 571300 (94000–1786000). Molting lasted from August through October, and a mass molt was recorded in August. As compared to crabs of the central part of the range, off the Korean Peninsula, Charybdis japonica in Ussuri Bay had larger size and higher fecundity. The breeding period in Ussuri Bay was slightly shifted from June–August to late June-September. The mass hatching of larvae occurred in July.     

Sublethal effects of iridovirus disease in a mosquito

Recognition of the importance of debilitating effects of insect virus diseases is currently growing. Commonly observed effects of sublethal infection at the individual level include extended development times, reduced pupal and adult weights, and lowered fecundity. However, for the most part, sublethal infections are assumed to be present in survivors of an inoculum challenge, rather than demonstrated to be present by microscopy or molecular techniques. Invertebrate iridescent viruses are dsDNA viruses capable of causing disease with symptoms obvious to the naked eye, a “patent” infection, that is lethal. Furthermore, inapparent “covert” infections may occur that are non-lethal and which can only be detected using bioassay or molecular techniques. In this study, replication of Invertebrate iridescent virus 6 in Aedes aegypti larvae was demonstrated in the absence of patent disease. A sensitive insect bioassay (using Galleria mellonella) allowed the detection of covert infections, which were more common than patent infections. A concentration-response relationship was detected for the incidence of patent infections. Covert infections were up to 2 orders of magnitude commoner than patent infections, but the prevalence of covert infections did not appear to be related to virus inoculum concentration. Exposure of larvae to virus inoculum resulted in extended juvenile development times. A reduction in the mean and an increase in the variability of fecundity and adult progeny production was observed in females exposed to an inoculum challenge, although formal analysis was not possible. Males appeared capable of passing virus to uninfected females during the mating process. Covertly infected females were smaller and had shorter lifespans than control or virus-challenged females. A conservative estimate for the reduction in the net reproductive rate (R ₀) of such insects was calculated at slightly more than 20% relative to controls. Received: 5 October 1998 / Accepted: 13 February 1999     

Reproduction and larval ecology of the Arctic fish speciesArtediellus atlanticus (Cottidae)

 State of maturity, egg numbers, as well as gonadosomatic index (GSI), and relative fecundity were determined in 209 females of the cottid species Artediellus atlanticus. Specimens were caught by Agassiz and bottom trawls during two expeditions of RV “Polarstern” to the shelf off northeast Greenland (ARK VII/2; July/ August 1990) as well as to the Barents Sea and the shelf areas surrounding Svalbard (ARK VIII/2 – Arctic EPOS; June/July 1991). Analysis of the state of maturity showed that A. atlanticus spawns from late summer to autumn. Individuals from northern populations spawn later (November) than those from more southern locations (July to September). The total number of eggs (40–216) was small due to their large diameter and the small size of the fish. The mean GSI of females from the central Barents Sea (36.9±6.5%; n=24) was significantly higher (P<0.001) than the GSI of females caught off eastern Svalbard (18.7±4.0%; n=14), compared at stage IV of maturity. The mean relative fecundity was significantly higher (P<0.001) for females of the central Barents Sea (17.9±2.3; n=19) than for females caught on more northern stations east of Svalbard (6.9±1.8; n=17) and north of Svalbard (6.4±0.5; n=7). Individuals of A. atlanticus that were maintained at 0°C spawned in the aquaria, allowing observation of the development of their eggs and larvae for 1 year. The eggs needed more than 200 days to develop. The larvae hatched at a very advanced stage and were similar to the adults in appearance and behaviour. The larvae grew at a rate of 0.02 mm per day and showed a benthic mode of life. Received: 7 April 1995/Accepted: 30 July 1995     

Phytoplankton blooms and fish recruitment rate: Effects of spatial distribution

We consider the spatio-temporal dynamics of a spatially-structured generalization of the phytoplankton-zooplankton-fish larvae model system proposed earlier (Biktashev et al., 2003, J. Plankton Res. 5, 21–33; James et al., 2003, Ecol. Model. 160, 77–90). In contrast to Pitchford and Brindley (2001, Bull. Math. Biol. 63, 527–546), who were concerned with small scale patchiness (i.e., 1–10m), on which the (stochastic) raptorial behaviour of individual larvae is important, we address here the much larger scale ‘patchy’ problems (i.e., 10–100 km), on which both larvae and plankton may be regarded as passive tracers of the fluid motion, dispersed and mixed by the turbulent diffusion processes. In particular, we study the dependence of the fish recruitment on carrying capacities of the plankton subsystem and on spatio-temporal evolution of that subsystem with respect to the larvae hatching site(s). It is shown that the main features found both in the nonstructured and age-structured spatially uniform models are observed in the spatially structured case, but that spatial effects can significantly modify the overall quantitative outcome. Spatial patterns in the metamorphosed fish distribution are a consequence of quasi-local interaction of larvae with plankton, in which the dispersion of larvae by large scale turbulent eddies plays little part due to the relatively short timescale of the larvae development. As a result, in a strong phyto/zooplankton subsystem, with fast reproduction rate and large carrying capacity of phytoplankton and high conversion ratio of zooplankton, recruitment success depends only on the localization and timing of the hatching with respect to the plankton patches. In a weak phyto/zooplankton system, with slow reproduction rate and small carrying capacity of phytoplankton and low conversion ratio of zooplankton, the larvae may significantly influence the evolution of the plankton patches, which may lead to nontrivial cooperative effects between different patches of larvae. However, in this case, recruitment is very low.     

A laboratory study of the effect of non-phytoplankton diets on the reproduction of Calanus helgolandicus

The effect of five non-phytoplankton and one phytoplankton dietson reproduction is described for the copepod Calanus helgolandicusand compared to field fecundity estimates. A fecundity increasefollowed ingestion of larvae of the sea urchin Sphaerechinusgranularis (LSU) and the oyster Crassostrea gigas (OYS) by copepodfemales, whereas a decrease followed ingestion of eggs of thecopepods Acartia and Temora spp. (COP), as a function of proteinconcentration in diets. With larvae of the sea urchin Paracentrotuslividus (SSU) and the dinoflagellate Prorocentrum minimum (PM;standard phytoplankton diet), fecundity was identical. Althoughprotein concentrations differed greatly from one food treatmentto the other, mean fecundity values were rather low. For unknownreasons, and despite their very high protein concentrationsin comparison to in situ, none of these diets could improvefemale fecundity or reach maximum egg production rates reportedin C.helgolandicus. Hatching rates were >80%, stable andsimilar with both PM and COP diets. With SSU, LSU and OYS diets,egg viability was lower than in situ and with PM, showing astrong and mild decrease with time with LSU and SSU, respectively.Ingestion of zygotes of the brown alga Fucus spiralis (ZYG)led to the death of females. These results suggest that denseblooms of zygotes of macroalgae and invertebrate embryos andlarvae, occurring in the continental shelf as part of copepoddiets, could have different effects on the demographic responseof C.helgolandicus.     

New data on the reproductive biology of shortspine thornyhead <Emphasis Type="Italic">Sebastolobus alascanus</Emphasis> (Sebastidae) in the northwestern Pacific

New data on fecundity, sex ratio in different size groups, and the rate of sexual maturation of shortspine thornyhead Sebastolobus alascanus in the northwestern part of the Pacific Ocean are provided. Individual absolute fecundity of the studied females varies in the range of 175 000–821000 (on average, 392500) eggs. In Pacific waters off the Kuril Islands and Kamchatka, among individuals with a length to 41 cm, males dominate; in size groups of 42–59 cm, sex ratio is almost equal; among larger fish, females prevail. In the western part of the Bering Sea among analyzed fish with a length to 44 cm, sex ratio is almost equal; among larger individuals, females dominate. Half of males and females of shortspine thornyhead off the eastern coast of the northern Kuril Islands and southeastern Kamchatka mature at a length of approximately 41 cm that can be recommended as a commercial size for the given species in this area.     

Sex-specific response to nutrient limitation and its effects on female mating success in a gift-giving butterfly

Animals with complex life cycles respond to early food limitation by altering the way resources are allocated in the adult stage. Response to food limitation should differ between males and females, especially in organisms whose mating systems include nutritional nuptial gifts. In these organisms, males are predicted to keep their allocation to reproduction (sperm and nuptial gift production) constant, while females are predicted to sacrifice allocation to reproduction (egg production) since they can compensate by acquiring nuptial gifts when mating. In this study, we investigated how dietary nitrogen limitation during the larval stage affects sex-specific resource allocation in Pieris rapae butterflies. Also, we tested whether nutrient-limited females increased nuptial gift acquisition as a way to compensate for low allocation to reproduction. We found that as predicted females, but not males, sacrifice allocation to reproduction when larval dietary nitrogen is limited. However, females were unable to compensate for this low reproductive allocation by increasing their mating rate to acquire additional gifts. Females reared on low nitrogen diets also reduced wing coloration, a potential signal of female fecundity status. We suggest that female mating frequency is constrained by male mate choice based on females’ wing coloration. This study provides new insights into how larval dietary nitrogen, a key nutritional resource for all herbivores, alters male and female allocation to reproduction as well as to ornamentation.     

DIRECT SELECTION ON LIFE SPAN IN DROSOPHILA MELANOGASTER

An important issue in the study of the evolution of aging in Drosophila melanogaster is whether decreased early fecundity is inextricably coupled with increased life span in selection experiments on age at reproduction. Here, this problem has been tackled using an experimental design in which selection is applied directly to longevity. Selection appeared successful for short and long life, in females as well as males. Progeny production of females selected for long life was lower than for short-lived females throughout their whole life. No increase of late-life reproduction in long-lived females occurred, as has been found in selection experiments on age at reproduction. This discrepancy is explained in terms of the inadequacy of the latter design to separate selection on life span from selection on late-life fecundity. Moreover, starvation resistance and fat content were lower for adults selected for short life. In general, the data support the negative-pleiotropy–disposable-soma theory of aging, and it is hypothesized that the pleiotropic allocation of resources to maintenance versus to reproduction as implicated in the theory might involve lipid metabolism. It is argued that further research on this suggestion is urgent and should certainly comprise observations on male reproduction because these are for the greater part still lacking. In conclusion, the longevity of D. melanogaster can be genetically altered in a direct-selection design, and such an increase is accompanied by a decreased general reproduction and thus early reproduction.     

The effects of habitat complexity on aggression and fecundity in zebrafish (<Emphasis Type="Italic">Danio rerio</Emphasis>)

Female zebrafish housed in aquaria with spatial complexity (plastic plants) over a 13–16-week period showed reduced levels of aggressive behavior compared to females in bare tanks. In tanks with plants, there was no relationship between levels of aggression and fecundity but, in bare tanks, females experiencing the highest levels of aggression showed reduced fecundity. Our results suggest that it may be beneficial, when maintaining zebrafish at moderate to high densities or working with especially aggressive strains, to house them in spatially complex conditions.     

Some reproductive characteristics of endangered Caspian Lamprey (<Emphasis Type="Italic">Caspiomyzon wagneri</Emphasis> Kessler, 1870) in the Shirud River southern Caspian Sea,Iran

Migration and reproduction of the Caspian Lamprey, Caspiomyzon wagneri, in the Shirud River were investigated during late-March to early-May at water temperatures ranging from 11 to 21.25°C. We examined the effect of water temperature on timing of spawning migrations. There was a significant negative relationship between temperature and intensive migration of Caspian Lamprey (p < 0.05). The most intensive migration of lampreys was at night (21:00–3:00 h) and when the water temperatures averaged 16°C (34.43%). The overall sex ratio (male to female) was 1.07 to 1. The individual absolute fecundity was 31 ‘758–51’ 198 eggs (mean±SD—41,924 ± 5,382). The egg diameter was 0.780–1.151 (0.92 ± 0.081) mm. The individual relative fecundity varies from 80.3 to 148.1 (107.2 ± 15.1) eggs per 1 mm of length and from 260.8 to 677.4 (397.6 ± 93) eggs per 1 g of weight. The gonadosomatic index (GSI) of females was 5.83–31.44 (11.22 ± 4.30).     

Life history,secondary production and population dynamics of <Emphasis Type="Italic">Gammarus fossarum</Emphasis> (Koch, 1836) in a constant temperature stream

Population of the freshwater amphipod Gammarus fossarum was investigated in a calcareous stream with almost constant temperature (7–8°C) in the Chočské Vrchy Mts (West Carpathians, Slovakia). Quantitative samples of G. fossarum taken during 2005 showed population densities varying from 100 m⁻² in August to 585 m⁻² in late November. The population was split into juveniles, mature males, mature females without eggs and females with eggs. The percentage of juveniles (40–64%) was always the highest of any of the categories. Ovigerous females occurred throughout the year. The mean sex ratio was 1: 2.4 (male: female), although its values varied considerably with the time of year. Breeding was continuous, although juvenile recruitment peaked in early spring, summer and early winter. Three discrete cohorts were distinguished from the size frequency distributions. The life span was 6–7 months and the individuals matured approximately in the half of life cycle. The mean fecundity was 9.6 embryos per brood. Variation in fecundity was mostly explained by size of the incubating females. The absolute growth of this species was best described by the Gompertz growth function. Relative growth rates (% body DM day⁻¹) fluctuated in a nonlinear manner with size and age. The highest values of daily growth (2–4% of dry mass per day) were noted approximately in the half of life cycle. Annual production, estimated by the size-frequency method, was 1618.9 g dry mass m⁻² and P/B ratio was 5.15.     

Effects of plant density and nutrient levels on fruit polymorphism in Atriplex sagittata

Atriplex sagittata is a heterocarpic species producing three types of fruits which differ in morphology and ecological properties. This study focused on variation in biomass allocation into particular fruit types under different density and fertilization levels. The reduction in total weight under stressful conditions was accompanied by a reduction of reproductive structures in terms of both total fruit production and mean fruit weight. Allocation of biomass to particular fruit types under different density-fertilization treatments showed considerable variation. The non-dormant, bracteolate fruit type (termed C) contributed 80–90% to the total fecundity and its production was not affected by density-fertilization treatments. The production of this fruit type is higher in the upper part of the maternal plant stem. Production of the very dormant ebracteate fruit type (termed A) increased under favourable conditions and was greater on the lower part of the stem, whereas more of the so-called type B fruit (covered by bracteoles and exhibiting dispersal and dormancy characteristics intermediate between types A and C) was produced under suboptimal conditions and was concentrated in the middle part of the plant body. This pattern contradicts to some extent the theory that deeply dormant and less dispersible fruits will be produced under stressful conditions and may be considered a new model for the behaviour of plants with more than two heterocarpic fruits. The type B fruit, intermediate in both morphology and ecological behaviour, provides a continuum between extreme strategies. Received: 8 February 1998 / Accepted: 19 November 1998