Swimming performance and endurance of the Norway lobster Nephrops norvegicus

The swimming behaviour of the Norway lobster, Nephrops norvegicus (L.), was studied in the laboratory. The lobsters were stimulated to swim repeatedly until they failed to respond to a single tactile stimulus. The position of the initiating stimulus on the body determined the height of the escape path above the bottom of the tank. A stimulus to the rostrum resulted in a low swimming trajectory, rarely exceeding a height of 0.1 m. A similar tactile stimulus to the abdomen produced higher mean swimming trajectories up to 0.5 m. Several parameters of swimming performance and endurance were measured for each swimming sequence, including distance, duration, mean and maximum velocities and the frequency and number of tail beats. Results showed no significant differences in the average swimming performance and endurance of males and females, but there was a sex difference in the relationship between swimming performance and N. norvegicus size.     

Some direct observations on the ecology and behaviour of the Norway lobster Nephrops norvegicus

A detailed investigation of a small area of sea bed occupied by the Norway lobster Nephrops norvegicus (L.) was carried out by diving and television observations at depths of 30 m in Loch Torridon, Scotland. The density of burrows was 1/2 m², but only a proportion of these were occupied by N. norvegicus. Although about 70% of the larger burrows were occupied by N. norvegicus, giving a density of 1 lobster/8 m², very few juveniles (carapace length less than 30 mm) were found in the area. Many of the small burrows were occupied by the gobiid fish Lesueurigobius friesii (Collett). There was evidence that N. norvegicus frequently change their burrows, and fighting for burrows was observed. N. norvegicus remain within their burrows during the day, emerge around sunset to forage for food during the night, and then return to their burrows at dawn. This, and other aspects of their burrowing behaviour, have a marked effect on the commercial trawl catches of N. norvegicus which show large seasonal and diurnal variations in size and sex composition.     

Behavioural and physiological responses of the Norway lobster,<Emphasis Type="Italic"> Nephrops norvegicus</Emphasis> (Crustacea: Decapoda), to sulphide exposure

The Norway lobster, Nephrops norvegicus, is moderately tolerant of sulphide [the lethal time to 50% mortality (LT₅₀) was 22.5 h when exposed to 500 M sulphide] but, whenever possible, it attempts to avoid the presence of sulphide in its immediate vicinity. Any sulphide entering the animal is oxidized to thiosulphate, which accumulates in the haemolymph and in the tissues. During exposure to low concentrations of sulphide, the rate of oxygen consumption is maintained or even enhanced even though the lobsters become quiescent. The apparent increase in oxygen consumption is probably due to the oxidation of sulphide to thiosulphate. At higher concentrations, oxygen consumption decreases, perhaps because of the inhibitory effect of sulphide on electron transport, and N. norvegicus resorts to anaerobic metabolism as indicated by the accumulation of lactate in the haemolymph and in the tissues.Communicated by L. Hagerman, Helsingør     

Effect of aerial exposure on concentrations of selected metabolites in blood of the Norwegian lobsterNephrops norvegicus (Crustacea: Nephropidae)

The Norwegian lobsterNephrops norvegicus (L.) collected from Firth of Clyde, Scotland between December 1987 and March 1988, was unable to survive longer than 18 h experimental emersion at 10°C. During this time the partial pressure of oxygen (P O₂) in the venous blood decreased rapidly and the lobster supplemented cellular energy requirements by anaerobic metabolism. This was indicated by the rapid accumulation ofL-lactate in the blood. Although the survival rate increased (to ca 36 to 48 h) if lobsters were kept on ice, the accumulation ofL-lactate in the blood was not significantly different from lobsters at 10°C, despite the temperature difference. There was no indication thatN. norvegicus was able to further metabolize circulatingL-lactate during emersion. On emersion there was also a marked hyperglycemia in the blood due to the stress of handling and asphyxiation. There was fairly good agreement between results obtained during laboratory studies and simulated fishing activity in the Firth of Clyde. Both sets of results are discussed in the context of adaptation to air breathing within the Crustacea and an assesment of post-harvest treatment of lobsters.     

Observations on the burrows and burrowing behaviour of two mud-dwelling decapod crustaceans,Nephrops norvegicus and Goneplax rhomboides

The burrows of the Norway lobster Nephrops norvegicus (L.) and of the crab Goneplax rhomboides (L.) were studied in Loch Torridon, Scotland. Polyester resin casts of burrows in the sea were made by divers to reveal their subsurface form. Tunnels made by N. norvegicus were usually simple, with two or more openings on the mud surface, and penetrated to a depth of about 30 cm. G. rhomboides burrows did not descend more than about 15 cm beneath the surface, but were usually more complex than the lobster burrows and had several openings. The methods of burrow construction used by the two crustaceans are described from aquarium observations. Neither N. norvegicus nor G. rhomboides show obvious morphological adaptations for burrowing, and it is suggested that the fossorial habit was adopted very early by decapods. The burrows of N. norvegicus do not seem to have assumed any functions in addition to the original one of providing refuge from predators. There is not sufficient known of the biology of the crab to indicate whether the same is true in its case.     

Behavioural responses of competent lobster postlarvae to odor plumes

We determined the ability of competent laboratory-reared Homarus americanus postlarvae to use odor plumes as a potential mechanism of orientation during the habitat selection process at settlement. In a Y-maze apparatus, both swimming direction (upstream or downstream) and arm selection (control or experimental) were strongly influenced by the nature of the chemical stimulus tested. In control conditions (unconditioned water in both proximal arms of the experimental device), postlarvae swam predominantly upstream (61:39) but displayed no arm preference (48:52). When submitted to a metabolite solution of adult conspecifics or macroalgae, postlarvae swam consistently upstream (73%) and showed a significant preference for the arm providing these stimuli (69%). In contrast, postlarvae submitted to cunner (Tautogolabrus adspersus) metabolites swam predominantly downstream (68:32), and those that did swim upstream significantly avoided the experimental arm (30:70). In repeated trials with the same individuals, postlarvae behaved similarly in 80% of the assays, suggesting a low intra-individual variability in behavioural responses. It is concluded that distance chemoreception may play a role in habitat selection by lobster postlarvae at settlement.     

Investigations of carotenoids in some fauna of the Adriatic Sea. III. Leander (Palaemon) serratus and Nephrops norvegicus (Crustacea: Decapoda)

Investigations have been carried out on the carotenoids in Leander (Palaemon) serratus (Pennant) and Nephrops norvegicus (L.) (Crustacea: Decapoda) from the Adriatic Sea. The presence of the carotenoids was determined by means of columnar and thin-layer chromatography. The following carotenoids were found: Leander serratus: -carotene, echinenone, isocryptoxanthin, cryptoxanthin, canthaxanthin, lutein, isozeaxanthin, zeaxanthin, astaxanthin, and astacene; Nephrops norvegicus: -carotene, lutein, zeaxanthin, astaxanthin, and astacene.     

Effect of the neurotransmitters dopamine,serotonin and norepinephrine on the ciliary activity of mussel (Mytilus edulis) larvae

The acute and long-term effects of neurotransmitters dopamine (DA), serotonin (SE) and norepinephrine (NE) on the feeding rates of Mytilus edulis veliger larvae were investigated through concentration-response curves. Increasing DA concentrations increasingly inhibited food uptake. Acute exposure to high levels of DA caused long-term inhibitory effects on feeding rates (10^–5 MDA) and growth rates (3x10^–4 MDA). Feeding activity was also inversely related to NE concentration. SE concentrations between 10^–8–3x10^–7 M supported enhanced feeding rates. Neither NE nor SE showed long-term inhibitory effects on feeding at concentrations <10^–4 M. These results were consistent with the observed effects of the different neurotransmitters on the swimming pattern of the larvae. The experimental evidence supports the model of ciliary control in adult mussels, involving dual innervation of the ciliated cells of the velum, with excitatory serotonergic and inhibitory dopaminergic fibers.     

Natural variation in the concentrations of haemocyanin from three decapod crustaceans, Nephrops norvegicus, Liocarcinus depurator and Hyas aranaeus

Natural variation in the concentrations of haemocyanin ([Hc]) is examined for three decapod species collected from two different locations (Gullmarsfjord and Kattegat) off the west coast of Sweden (August to September 1998). Only one of the frequency distributions for [Hc] is normally distributed, and median values differ both between and within species. Differences in [Hc] cannot be attributed to sex, reproductive condition or moulting condition (over the limited range of moult stages examined) for any of the species. While body size did not scale with [Hc] for the Norway lobster Nephrops norvegicus or the swimming crab Liocarcinus depurator, it did for the spider crab Hyas araneus. To our knowledge this is the first time a negative relationship between body size and [Hc] has been reported for any crustacean. Thus the right-skewed frequency distribution of spider crab [Hc] could be accounted for by the right-skewed body size distribution recorded. The shape of the frequency distributions for [Hc] from the other two species could not be accounted for through differences in the factors examined here, although it is suggested that the amount of food consumed (or not) may be important. Received: 19 July 1999 / Accepted: 4 October 1999     

Highly directional swimming by scalloped hammerhead sharks,Sphyrna lewini,and subsurface irradiance,temperature, bathymetry,and geomagnetic field

The homing behavior of scalloped hammerhead sharks (Sphyrna lewini) to and fro between Espiritu Santo Seamount and Las Animas Island and the surrounding pelagic environment was studied to reveal their mechanisms of navigation in the oceanic environment. Four sharks were tagged with ultrasonic transmitters and tracked at the former location and one shark at the latter site during July, August, or September between 1981 and 1988. Hammerhead swimming movements were highly oriented: the mean coefficient of concentration (r) for sets of ten consecutive swimming directions recorded during eight homing movements by three hammerhead sharks ranged from 0.885 to 0.996. Drift within a current could not explain this directionality, since highly variable directions were recorded from a transmitter floating at the sea surface after becoming detached from a shark. Forward swimming momentum was an unlikely explanation, since highly directional swimming was maintained for a period of 32 min with only a gradual change in course. To maintain directionality over this period, an environmental property should be necessary for guidance. The hammerheads swam at night, with repeated vertical excursions ranging from 100 to 450 m deep, out of view of either the sea surface or the sea floor. The sharks' vertical diving movements were compared to distributions of spectral irradiance (relative to elasmobranch scotopic and photopic visual sensitivities), temperature, and current-flow directions in the water column. No relationships were evident between these properties and the sharks' oriented swimming movements. Movements of scalloped hammerhead sharks to and from a seamount were compared to topographic features in bathymetry and geomagnetic field leading away from the seamount. Sharks swam repeatedly over fixed geographic paths, and these paths occurred less often along submarine ridges and valleys than maxima and minima in the geomagnetic field. No significant difference existed between the degree of association of points from the sharks' tracks and points from track simulations and 20° changes in the slope of the depth record. On the other hand, significantly more points from the sharks' tracks were associated with slope changes in the magnetic intensity record than points from track simulations. A magnetic intensity gradient of 0.037 nanoteslas/m (nT/m) existed at 175 m depth, where a shark swam directionally, and this gradient was three times steeper than that measured at the sea surface and exceeded that recorded at a depth of 200 m. The hammerheads are hypothesized to find the seamount using geomagnetic topotaxis. The shark could be attracted to and move back and forth along ridges and valleys, features in the relief of magnetic field intensities occurring over a geographical area.     

The eye and some effects of light on locomotor activity in Nephrops norvegicus

The compound eye of Nephrops norvegicus (L.) is of the superposition type, well-adapted to the low levels of light prevailing at the sea bed during the activity periods of the species. Only the proximal retinal shielding pigment responds to light, the distal retinal shielding pigment being in the dark-adapted position at all times. The response of the proximal pigment appears to vary seasonally. Field observations compared light intensity at the sea bed with the numbers of N. norvegicus caught by trawl at various times of day in the Irish Sea in summer and winter. Laboratory experiments were combined with these field data to indicate that light is an important modulator of locomotor activity in this species.     

Gender- and oxygen-related irrigation behaviour of the decapod Nephrops norvegicus

The Norway lobster Nephrops norvegicus (L.) inhabits burrows in muddy clay sediments (e.g. on the Swedish west coast), where an autumnal oxygen deficiency in the bottom water can occur. Our experiments investigated whether the irrigation of the burrows would reflect a behavioural adaptation to hypoxia, and whether any gender differences of such behaviour exist. Irrigation is performed by the pleopods which may compensate for a decreasing oxygen tension. Pleopod activity (total number of strokes per sampling time), associated with oxygen concentration and gender, was studied in N. norvegicus kept in artificial burrows resembling their natural habitat. Male and female lobsters were separately exposed to either normoxia (70% oxygen saturation) or hypoxia (30% oxygen saturation). A sexual difference in behaviour was found, where females irrigated the burrow less than males during normoxia. Females showed a significant increase of pleopod activity in hypoxia compared with normoxic conditions, which was not displayed by the males probably due to the degree of individual variation found. However, when only males were studied during progressive hypoxia (from 60 to 5% oxygen saturation), following any changes of irrigational behaviour, a significant increase of accumulated pleopod activity occurred. A major increase of pleopod activity appeared between 60 and 50% oxygen saturation, below which the activity remained high until a critical point (<10% saturation, 11 °C, 33 psu) where irrigation dropped to a level close to that of normoxic values. Activity sessions during hypoxia were longer and had a higher stroke rate than during normoxia. Received: 22 October 1997 / Accepted: 26 February 1998     

Swimming pattern as an indicator of the roles of copepod sensory systems in the recognition of food

The roles of copepod sensory systems in the recognition of food were investigated using the Bugwatcher, a video-computer system designed to track and describe quantitatively the swimming patterns of aquatic organisms. The swimming behavior of the copepodPseudocalanus minutus in the presence of phytoplankton is characterized by a decrease in average swimming speed and an increase in pause behaviors compared to its swimming behavior in filtered seawater. Copepods exposed to chemosensory stimulation alone (filtered phytoplankton exudate) exhibited an increase in average swimming speed and an increase in the number of burst swimming behaviors. When exposed to a novel, non-food chemosensory stimulus (morpholine), no change in swimming behavior was observed unless the copepods had been conditioned to this odor in the presence of phytoplankton. Copepods exposed to mechanosensory stimulation alone (plastic spheres) exhibited a decrease in swimming speed and an increase in pause behaviors. When exposed to both forms of stimulation simultaneously (phytoplankton exudate and plastic spheres), a further decrease in swimming speed and increase in pause behaviors occurs, yielding a swimming pattern similar to that found in the presence of phytoplankton. This analysis of swimming pattern indicates that both chemoreception and mechanoreception contribute to the recognition of food inP. minutus.Contribution No. 406 of the US EPA Environmental Research Laboratory; Narragansett, Rhode Island 02882, USA     

Locomotion,feeding, grooming and the behavioural responses to gravity,light and hydrostatic pressure in the stage I zoea larvae of Ebalia tuberosa (Crustacea: Decapoda: Leucosiidae)

The stage I zoeae of Ebalia tuberosa swam by sculling with the exopodites of the 1st and 2nd maxillipeds and flexed the abdomen to brake or change direction. The larvae gained depth by stopping all natatory movements and sinking passively at rates of 6 mm s^-1. The zoeae refused both living and dead nauplii of Artemia spp., as well as two species of diatoms, but fed readily on detritic material on the bottom which they scooped up using the endopodites of the maxillipeds and pressed against the mouthparts using the telson. The setae on the posterior border of the telson were used for grooming the maxillipeds and the anterior mouthparts. Day-old stage I zoeae were negatively geotactic, positively phototactic and responded to pressure increases by swimming upwards and by high barokinesis. By the third day some larvae had become positively geotactic but were photopositive, and the majority responded to pressure increases as in the day-old larvae. Five-day old larvae were still photopositive but the majority had become positively geotactic and fewer himbers responded to pressure. Seven-day old larvae failed to respond to any of the stimuli used and assumed a predominantly benthic lifestyle. It is suggested that this anomalous behaviour is related to the dispersal of the larvae and to the specialized habitat requirements of the adults while the rather unusual morphology of the larvae is related to their feeding behaviour and semi-benthic lifestyle.     

Between-individual variation in haemocyanin concentrations in the Norway lobster<Emphasis Type="Italic"> Nephrops norvegicus</Emphasis> following exposure to hypoxia and manganese

Environmental hypoxia and the release of reduced manganese (Mn) are often combined stress factors in marine sediments. Previous investigations have shown that hypoxia induces variable responses in the concentration of the respiratory pigment haemocyanin (Hc) of crustaceans. A recent study demonstrated that the magnitude and direction of the changes was dependent on the initial level of Hc in individuals of the Norway lobster Nephrops norvegicus. The changes also took place within 24 h rather than days as suggested by previous studies. In this present study we investigated changes of Hc in individual N. norvegicus in uncontaminated sea water after exposure to and release from realistic hypoxic stress (PO ₂=6.0 kPa). Furthermore we investigated how concomitant exposure to realistic concentrations of manganese (20 mg l ^-1) modified the responses we observed. We confirmed a between-individual variation in response and a change in Hc towards an optimum level after hypoxia, but also showed that changes that took place post-hypoxic exposure occurred over a timescale similar to that of the hypoxic response itself. Manganese exposure resulted in no significant changes in Hc whereas Mn exposure combined with hypoxia resulted in a significant decrease (15%) in all individuals independent of initial Hc. N. norvegicus was thus unable to compensate for the effects of hypoxia by synthesis of Hc after exposure to naturally occurring Mn concentrations.     

Biremis blandi (Polychaeta: Terebellidae), new genus,new species,caught by D.S.R.V. “Alvin” in the Tongue of the Ocean,new providence,Bahamas

Biremis, n. gen., characterized by an enlarged cephalic lobe, achaetous thoracic segments, and abdominal bilobed uncinigerous pinnules, lacks eyes and branchiae and is, thus, grouped with the subfamily Polycirrinae of the Terebellidae. Biremis blandi, n. sp., has prominent longitudinal muscular ventral ridges, lacking ventral glandular scutes. The avicular uncini, arranged in a single row on each lobe of each abdominal pinnule, are very similar to those of Polycirrus. The new species, collected from D.S.R.V. Alvin, was observed resting on and swimming over the sloping walls of the Tongue of the Ocean (25° N; 77°35 W), at 600 m depth. Its ability to swim may be a requirement for survival on an unstable substratum.Contribution No. 2891 from the Woods Hole Oceanographic Institution.     

Wave exposure,swimming performance,and the structure of tropical and temperate reef fish assemblages

We examined the relationship between swimming performance, wave exposure, and the distribution patterns of labrids on temperate rocky reefs, in comparison with previous functional analyses of a tropical assemblage. Visual censuses of the distribution and abundance of labrids across two major gradients of wave exposure (depth and aspect to prevailing winds) were made at two offshore islands near Port Stephens, New South Wales, Australia. Distinct shifts in species composition and abundance were evident between high and low wave exposure habitats on temperate rocky reefs, particularly between deep and shallow habitats on exposed reef fronts. The swimming performances of temperate labrids were assessed through examination of pectoral fin shape (aspect ratio) and in situ swimming speeds. A diversity of pectoral fin morphologies was exhibited within this temperate assemblage, ranging from rounded to tapered fins (aspect ratios of 0.52 and 1.43, respectively). Fin shape was strongly correlated (Pearsons correlation 0.884, P<0.001) with swimming speed (ranging from 1.05 and 3.06 body lengths s^–1), in a relationship comparable to that observed in tropical labrids. Inter-specific differences in swimming ability provided some explanation for differences in the distribution and abundance of temperate labrids in relation to wave exposure. However, our findings suggest that although coral reef labrids appear to predominantly use high aspect-ratio fins to successfully occupy wave-exposed habitats, temperate labrids appear to be using an enhanced swimming ability through increased body size.Communicated by G.F. Humphrey, Sydney     

Temperature and plankton

The relationship between temperature and metabolism was studied in Artic copepods with regard to the concept of metabolic cold adaptation of polar poikilotherms. Temperature tolerance and respiration rates of the dominant copepods Calanus finmarchicus (Gunnerus), C. glacialis (Jaschnov), C. hyperboreus (Krøyer) and Metridia longa (Lubbeck), collected in Fram Strait, Greenland Sea, in July 1983, were studied at different temperatures. Temperature tolerance in the boreal C. finmarchicus was slightly higher than in the three Arctic species. Respiration rates at lower temperatures followed the Arrhenius equation in all species, with values for (temperature characteristics) between 11.05 and 22.95, corresponding to a Q₁₀ between 2.05 and 4.5. This increase in metabolic rate with rising temperature was not related to an increase of swimming activity, as was shown by videoanalysis. Activity was determined as average swimming speed and as frequency of certain locomotor patterns. Average swimming speed remained unchanged at all temperatures and was ca 1 cm s^-1 for all species, when only periods of active swimming were considered. The time spent with active swimming did not change with temperature in M. longa and C. finmarchicus, but decreased in c. glacialis. In C. hyperboreus it increased at 5°C and decreased again at higher temperatures. It is suggested that the increase in oxygen consumption is fully accounted for by the basal metabolism.     

The effect of hypoxia on behavioural and physiological aspects of lesser sandeel, <Emphasis Type="Italic">Ammodytes tobianus</Emphasis> (Linnaeus, 1785)

Lesser sandeel (Ammodytes tobianus) is abundant in near-shore areas where it is a key prey. It exhibits the behaviour of alternating between swimming in schools and lying buried in the sediment. We first determined the species’ standard metabolic rate (SMR), critical partial pressure of oxygen and maximal oxygen uptake The sandeel were then exposed to an acute stepwise decline in water oxygen pressure (18.4, 13.8, 9.8, 7.5, 5.8, 4.0, and 3.1 kPa ). Swimming speed and routine- and post-experimental blood lactate levels were measured, in addition to levels associated with strenuous exercise. The SMR was 69.0 ± 8.4 mg O₂ kg⁻¹ h⁻¹ and the about seven times as high. The was found to be 4.1 kPa. A rapid decrease (within 1 h) in from 18.4 to 3.1 kPa had no significant effect on routine swimming speed (0.9 ± 0.06 bl s⁻¹), but steady levels at the lowest (3.1 kPa) gradually reduced the swimming speed by 95% after 40 min. The routine blood lactate levels were 2.2 ± 0.6 mmol l⁻¹, while the levels in the strenuously exercised groups were significantly higher with 5.4 ± 1.6 and 5.8 ± 1.3 mmol l⁻¹. The highest levels were observed in post-experimental fish with 7.5 ± 2.7 mmol l⁻¹. We argue that, as sandeel showed no decrease in swimming speed (to offset stress) nor an increased speed to escape the hypoxia, the fish either rely on a low SMR and being a reasonable strong oxygen regulator as a mean to cope when exposed to acute hypoxia, or that the hypoxia simply developed too fast for the fish to decide on an appropriate strategy. Not showing a behavioural response may in the present case be maladaptive, as the consequence was major physiological stress which the fish however appears tolerant towards. The high routine blood lactate levels suggest that anaerobic metabolism is associated with swimming in sandeel, which may be related to the specific lifestyle of the fish where they regularly bury in the sediment.     

Behaviour and tolerance to hypoxia in juvenile Norway lobster (Nephrops norvegicus ) of different ages

The annual occurrence of hypoxia (<25% oxygen saturation) in the bottom waters along the Swedish west coast coincides with the postlarval settlement of Norway lobster, Nephrops norvegicus (L.). This study investigates behaviour and the experimental effects of low oxygen concentrations in juvenile N. norvegicus of different ages. All experimental individuals were reared to the juvenile (postlarval) stage in the laboratory and then given sediment as a substratum. Behavioural responses to low oxygen concentrations were tested in early and late Postlarvae 1 exposed to normoxia (>80% oxygen saturation, pO₂ > 16.7 kPa), moderate hypoxia (30% oxygen saturation, pO₂ = 6.3 kPa) and hypoxia (25% oxygen saturation, pO₂ = 5.2 kPa). The experiments were run for a maximum period of 24 h or until individuals died. Behaviour was studied using sequential video recordings of four behavioural activities: digging, walking, inactivity or flight (escape swimming up into the water column). Behaviour and mortality changed with lowered oxygen concentrations; energetically costly activities (such as walking) were reduced, and activity in general declined. In normoxia, juveniles initially walked and then burrowed, but when exposed to hypoxia they were mainly inactive with occasional outbursts of escape swimming. To increase oxygen availability the juveniles were observed to raise their bodies on stilted legs (similar to adults in hypoxic conditions), but oxygen saturations of 25% were lethal within 24 h. The results suggest that the main gas exchanges of early postlarval stages occur over the general body surface. Burrowing behaviour was tested in Postlarvae 1 and 2 of different ages held in >80% oxygen saturation for 1 wk. The difference in time taken to complete a V-shaped depression or a U-shaped burrow was measured. The results showed a strong negative relationship between postlarval age and burrowing time, but all individuals made a burrow. Juveniles were more sensitive to hypoxia than adults. Thus, the possible consequences of episodic hypoxia for the recruitment of Nephrops norvegicus and for the recolonization of severely affected areas are discussed. Received: 4 August 1996 / Accepted: 11 October 1996