Colonization of meiobenthos in oil-contaminated subtidal sands in the lower Chesapeake Bay

In-situ manipulative experiments were conducted over a 3-month period (May–August 1980) to examine the rate at which meiobenthos colonizes oiled and untreated azoic fine sands at a shallow subtidal site in the lower York River, Virginia. Three concentrations of fresh Prudhoe Bay crude oil were added to sediments: 100, 2 500 and 10000 mg oil kg^-1 dry wt sediment. Untreated azoic and natural sediments served as controls. Within 16 d, meiofauna densities in all treatments were comparable to natural populations in surface oxidized sediments, but densities fluctuated greatly during the remainder of the sampling period. Nematodes slowly colonized the subsurface anoxic sediments below the redox potential discontinuity (RPD); some less common species did not significantly recover below the RPD in the two more heavily oiled treatments. Analysis of nematode community composition by reciprocal averaging ordination and numerical classification revealed generally lower abundances, but no distinct differences, in species composition in the oiled substrates as compared to untreated and natural community controls. Ordination of sequential samples suggested that the nematode species assemblages in the untreated controls fully recovered from these small-scale disturbances by 90 d. Life history characteritics and frequent tidal transport combine to make estuarine meiobenthos highly resilient following disturbance. Contrary to prior recolonization studies, a successional sequence was found for the colonizing nematodes which may be analogous to models of macrobenthic colonization (e.g. McCall, 1975). The comesomatid nematode Sabatieria pulchra, which is frequently dominant in polluted sediments, colonized relatively late in the experiment. Consequently, stress resistance and resilience may not be as coincident in meiofauna as in macrofauna because of differences in factors affecting their dispersal.     

Inhibition of larval settlement to a soft bottom benthic community by drifting algal mats: An experimental test

The hypothesis that drifting red algal mats inhibit settlement of planktonic larvae was tested in a field experiment in 1986. Substratum free of algae (caged boxes) was compared with substratum covered with algae (natural substratum and open boxes). Whereas settling densities of 1500 to 5500 ind. m^-2 of the bivalves Macoma balthica, Cardium glaucum and Mya arenaria were observed in sediment without algal cover, no recruits of the same species were found beneath the algae during the period of peak settlement (June to July). The same difference was observed for the polychaete Nereis diversicolor, although in this case occasional individuals were found beneath the algae. The results demonstrate that algae mats may be efficient inhibitors of larval settlement to benthic soft-substratum communities. It is concluded that algae act as a larval filter.     

Microbial reaction rates and bacterial communities in sediment surrounding burrows of two nereidid polychaetes (Nereis diversicolor and N. virens)

The effects of infaunal mode of life on sediment properties, microbial reaction rates, as well as abundance and composition of bacterial communities were studied in sediment surrounding burrows (mucus lining, oxidised wall, ambient anoxic and surface sediment) of two closely related, but behaviourally different, nereidid polychaete worms: the facultative suspension-feeder Nereis (Hediste) diversicolor and the obligate deposit-feeder Nereis (Neanthes) virens. Burrow sediment of the two species was collected from two adjacent (50 m distance) shallow sandy locations (Kertinge Nor, Denmark). The burrow lining and wall of both polychaete species were enriched in organic matter originating from mucous secretions by the inhabitants and phytoplankton trapped through irrigation. This was more evident for N. diversicolor that shows a significantly higher irrigation rate than N. virens. Both the organic matter mineralisation rates (based on anaerobic incubations) and bacterial abundance were higher along the burrow linings and walls. However, accumulation of porewater TCO₂ and dissolved organic carbon in sediments adjacent to burrows increased most rapidly in the presence of N. diversicolor, suggesting higher heterotrophic activity associated with this species. Surprisingly, bacterial abundance was lower around burrows of N. diversicolor than those from N. virens indicating that burrow environments from the first species harbour a more active bacterial community. Molecular fingerprints of the 16S rRNA gene from bacterial communities showed that the composition of the burrow linings and walls resembled the ambient anoxic sediment rather than the oxic sediment surface. On the other hand, the bacterial fingerprints of the sediment surrounding the burrows of the two polychaete species were markedly different suggesting either a site-specific difference in sediment parameters or a significant species-specific impact of the burrow inhabitants.     

An experimental study of the relationship between below-ground structure and meiofaunal taxa

An experimental study was conducted in a South Carolina, USA salt marsh to examine relationships between meiofaunal-sized tube structure and densities of burrowing meiofauna. Abundance of the tube-building polychaete Manayunkia aestuarina Bourne was increased by erecting exciosure areas from June–August 1979. Non-manipulated areas of identical dimensions were staked out in adjacent locales. Densities of burrowing meiofauna and nematode trophic groups, and volumes of Spartina alterniflora root mat and tube-builders were monitored approximately every 2 wk from each experimental treatment. Volumes of tube-builders were significantly higher inside exclosure sites on all dates in July and August. Root-mat volume and total number of nematodes were not different between treatments at any time. The dominant copepod species, Stenhelia (D.) bifidia, and the nematode species, Spirinia sp., first increased in number inside manipulated sites and then decreased in abundance as tube volume increased. No general trend was apparent for nematode trophic groups when areas with increased volumes of tubebuilders were compared to unmanipulated sites. The fact that densities of S (D.) bifidia and Spirina sp. decreased only when tube-builders attained highest abundance suggests that negative interactions between tube-builders and these meiofaunal taxa may be effective only at elevated abundances.     

Natural variations in tissue concentration of copper,zinc and iron in the polychaete Nereis diversicolor

This study was undertaken to investigate the natural variation of heavy metals in the polychaete Nereis diversicolor O. F. Müller and to examine this variation with respect to physiological and environmental parameters. During a 2 yr period from October 1980 to October 1982, concentrations of copper, zinc and iron in N. diversicolor from the Tees Estuary, North East England, were found to vary significantly with time. A seasonal influence on whole body copper concentration, in part related to the reproductive cycle, was observed. Zinc concentrations in N. diversicolor increased during periods of rapid growth and may represent a physiological demand for this metal. Whole body concentrations of iron appeared to be independent of size of worm, sexual maturation and sediment concentrations.     

Effects of physical disturbance on nematode communities in sand and mud: a microcosm experiment

A microcosm experiment was carried out to evaluate the effects of continuous and spasmodic physical disturbance of differing frequency on the structure of nematode communities of intertidal sand and mud. There was a marked, characteristic change in abundance and diversity for both sediment types. In the sand microcosms, the majority of univariate measures of community structure, including species diversity, were lowest in the sediments subjected to a high frequency of disturbance. For the mud microcosms, most univariate measures reached their highest values in the treatments with an intermediate frequency of disturbance and were lower in treatments subjected to both higher and lower frequencies. Multivariate ordinations for both nematode assemblages showed a clear separation of undisturbed controls and disturbed treatments, but only for the muddy sediment was there a graded change in community composition with increasing frequency of disturbance. These results confirmed our a priori expectation that nematode assemblages from mobile sandy sediments would be more resilient to physical disturbance than those from sheltered muds, and these observations are considered in the context of Connell's intermediate disturbance hypothesis. Received: 7 May 1997 / Accepted: 10 October 1997     

Impacts of chronic trawling disturbance on meiofaunal communities

Bottom trawling causes chronic and widespread disturbance to the seabed in shelf seas. Meiofauna may be impacted directly or indirectly by this disturbance, since the passage of trawls causes immediate mortality or displacement, changes sediment structure and geochemistry and affects the abundance of predators or competitors. Since meiofauna make a significantly greater contribution to benthic production than the larger macrofauna, there are compelling reasons to assess their response to chronic trawling disturbance. In this study, we determined the effects of trawling disturbance, season, sediment type and depth on the structure and diversity of nematode communities. Our analyses were based on comparisons between nematode communities in three beam-trawl fishing areas in the central North Sea. These areas were trawled with mean frequencies of I (low disturbance), 4 (medium) and 6 (high) times year⁻¹ respectively. Our analyses showed that trawling had a significant impact on the composition of nematode assemblages. In two sampling seasons, the number of species, diversity and species richness of the community were significantly lower in the area subject to high levels of trawling disturbance than in the areas subject to low or medium levels of disturbance. However, levels of disturbance at the ‘low’ and ‘medium’ sites may have been insufficient to cause marked longterm changes in community structure. Many of the observed changes were consistent with responses to other forms of physical disturbance. The extent to which the observed changes in community structure reflect changes in the production of the nematode community remains unknown, although overall abundance was not significantly affected by trawling disturbance. Published online. 9 August 2002     

Minimal predation upon meiofauna by endobenthic macrofauna in the Exe Estuary,south west England

Field manipulation experiments were performed in the Exe Estuary, south west England, in October 1988, to investigate the importance of the meiofaunamacroinfauna trophic link in benthic trophodynamics. Four hypothetically meiofauna-predacious endobenthic macrofauna species were selected for manipulation using the criteria of high abundance and different modes of feeding: Cerastoderma edule (filter-feeder), Nereis diversicolor (omnivorous scavenger), Ophelia bicornis (sand-ingester), Scrobicularia plana (deposit-feeder). Enclosures constructed from plastic tubing, 63 m nylon monofilament mesh and galvanized steel were deployed, containing adult members of these taxa at densities raised to approximately four times that of the surrounding sediment. The experiments ran for 12 tidal cycles. Differences in phyletic meiofaunal abundance between treatment and control enclosures at the termination of the experiment were assessed using both uni- and multivariate dataanalysis techniques. Only two univariate significant differences (p<0.05) existed for the N. diversicolor treatment and two for the S. plana treatment. Annelida, Turbellaria and copepod nauplii were the only meiofauna taxa affected. No univariate significant differences were recorded for either the C. edule or O. bicornis treatments. Multidimensional scaling ordination of the data revealed no consistent changes in community composition between treatments and controls. It is concluded that the experiments provide evidence of minimal predation by macroinfauna upon meiofauna.     

Investigations on epibenthic meiofauna I. Abundances on and repopulation of the tube-caps of Diopatra cuprea (Polychaeta: Onuphidae) in a subtropical system

Cryptic meiofauna populated the imbricate, shell-sediment matrix of the tube-caps produced by the polychaete Diopatra cuprea (Bose) in Tampa Bay, Florida, USA. Nematodes, copepods (adults and nauplii), polychaetes (adults and juveniles) and amphipods (adults and juveniles) were the most abundant taxa found on tube-caps. Meiofaunal densities on tube-caps were 4 to 19 times higher than equivalent volumes of sediment in cores taken adjacent to D. cuprea tubes. Recruitment onto tube structure occurred within 1 to 2 d after defaunated tube-caps were replanted into sediments in the field (February–November 1980). Repopulation of copepods (adults and nauplii) attained levels equal to or exceeding natural abundances on tube caps within 1 to 5 d; nematode recovery rates were inconsistent. Short-term experiments using a variety of defaunated tube treatments indicated that immigration onto above-sediment tube-caps proceeds via both water column and sediment pathways. Based on data on tube-cap longevity and construction as well as meiofaunal recruitment rates, we conclude that the generation of new tube-cap structure is exploited rapidly by meiofauna.     

Transfer of 65Zn from sediments by marine polychaete worms

Silty marine sediments spiked with ⁶⁵Zn lose only small fractions of their radioactivity when exposed to slowly flowing seawater for several weeks. However, polychaete worms (Nereis diversicolor), burrowing through the sediment, cause ⁶⁵Zn losses 3 to 7 times higher than in sediment without worms. Long-term experiments on the uptake and loss of ⁶⁵Zn by the polychaete Hermione hystrix indicate that 60 or more days exposure are required for this worm to approach steady state with ⁶⁵Zn in the sediment. Biological half-life estimates for ⁶⁵Zn accumulated from sediment by H. hystrix are extremely variable (52 to 197 days), depending on the loss-time interval chosen for the calculation. Following 5 days exposure to 16 cm³ of radioactive sediment, N. diversicolor individuals contained an average of 0.2% of the total ⁶⁵Zn in the sediment. When these worms were transferred to non-radioactive sediment, estimates of biological half-life for ⁶⁵Zn averaged 14 to 17 days during the loss period Day 3 to Day 15. Based on these experimental results, it is estimated that a population of N. diversicolor could cause an annual loss of 3% or more of the ⁶⁵Zn in the upper 2 cm of the sediment of a hypothetical radioactive estuary.     

Allozyme electrophoretic analysis of the Hediste limnicola — H. diversicolor — H. japonica species complex (Polychaeta: Nereididae)

Three morphologically similar species of the polychaete genus Hediste [H. limnicola (Johnson, 1901), H. diversicolor (O. F. Müller, 1776), and H. japonica (Izuka, 1908)] have been described, respectively, from the west coast of North America, Europe, and Japan. Although these taxa are geographically isolated by oceans, the morphological criteria by which they can be separated are sufficiently variable as to make identification difficult. We have compared these three taxa biochemically using ten allozyme loci and have found that they are genetically distinct and constitute valid species. H. japonica (collected in June 1990 from Kyushu, Japan) appears more closely related to H. limnicola (collected in July 1989 from two sites in California and two in Oregon, USA) than it is to H. diversicolor (collected in October-November 1989 from the Weser estuary and the Jadebusen in Germany). H. limnicola has been recognized previously as a self-fertilizing hermaphrodite, but it is quite polymorphic in the four populations examined and we suggest that some cross-fertilization must occur in the field. Marked allele frequency differences between the two H. diversicolor populations examined, which were separated by only about 30 km, suggested that there was restricted gene flow between populations, or that there had been strong short-term selection.     

Benthic polychaete assemblages and their relationship to the sediment in Port Madison,Washington

Two indices of community association and a 3 dimensional ordination of stations were used to elucidate the relationship between small-scale changes in species, composition of polychaete assemblages and changes in the physical character of the sediment, at 9 stations along a 1.5 km subtidal transect. There were no faunal or physical discontinuities that might have been used to delineate boundaries of polychaete communities, except at a very shallow station where the effects of extreme fluctuations of temperature and salinity were evident. The degree of similarity between assemblages was related to the similarity of the sediments at the different stations. In particular, the changes in species composition appeared to correspond most clearly to differences in the clay content of the sediment.     

Modification of sediment permeability and shear strength by two burrowing invertebrates

Laboratory experiments have been conducted on the effects of bioturbation by the amphipod Corophium volutator (Pallas) and the polychaete Nereis diversicolor (O.F. Müller) on the physical properties of estuarine muddy sand. Invertebrates and muddy sediment were collected from Langbank, Firth of Clyde, Scotland. Sediment columns were seeded with low, medium and high densities of invertebrates in single-and mixed-species experiments. Permeability decreased with increasing density of C. volutator, but increased with increasing density of N. diversicolor. Water content decreased with increasing density of both species. Shear strength increased with increasing density of both species, but more so with C. volutator than with N. diversicolor. Mixed-species effects on shear strength were additive, but some of the permeabilities were lower than predicted. Bivariate and multivariate correlation, and partial correlation analyses were applied to the data. Shear strength was negatively correlated with water content. The correlations between shear strength and permeability and between permeability and water content were affected differently by the two species. There were few partial correlations between the variables.     

Uptake of 55Fe from contaminated sediments by the polychaete Nereis diversicolor

The polychaete Nereis diversicolor O. F. Müller was exposed to marine sediments which had been contaminated with ⁵⁵Fe either through the testing of nuclear devices or by the release of liquid waste effluent from a nuclear fuel reprocessing plant. The amount of ⁵⁵Fe taken up reached a concentration in the worms about 6% of the concentration in both types of sediment. Bioavailability of ⁵⁵Fe was shown to depend on its concentration in sediment and not on sediment type, stable iron concentration, or the source of contamination. N. diversicolor may not extract from sediment the same fraction of ⁵⁵Fe as does 0.1N HCl and caution should be exercised in using such measurements to represent the bioavailable fraction.     

Distribution quantitative de la méiofaune dans l'aire d'épandage de l'égout de Marseille

A quantitative analysis of meiofauna was carried out at ten sample stations distributed along a transect off Marseilles' (France) sewage outfall in November, 1981. The meiofauna was counted and average and total biomass of nematodes and copepods were calculated for each station. Comparison of data between stations or groups of stations was made using statistical tests. The results revealed three sectors in which effects of pollution on meiofauna differed: (1) A heavily polluted coastal zone (at a distance of 30 to 150 m from the outfall), relatively poor in meiofauna, which, however, was not completely absent-as were macrobenthic organisms; the meiofauna was composed mainly of nematodes and copepods, but acarians and naupliar larvae (probably copepods) were also numerous; the copepods were uncommonly large in size, and alone consituted most of the total benthic biomass in this zone. (2) An intermediate zone (400 to 1 150 m away from the coast), also polluted and much richer in meiofauna than the coastal zone; the meiofauna groups were more diversified, annelids (mainly polychaetes) increasing in number, while acarians became scarce and copepods decreased in size; at 400 m away from the outfall, where the sediment is strongly polluted, the nematode population consisted of large individuals which contributed greatly to the biomass; at 400 and 800 m, the distribution of both meiofauna and macrobenthos was heterogeneous; at 1 150 m away from the outfall, maximal meiofaunal densities were recorded, due primarily to an increase in nematode numbers; here, unlike all the stations nearer to the coast, total nematode biomass was greater than copepod biomass. (3) An off-shore zone (1.8 to 4 km distant from the coast), slightly polluted, where meiofauna densities were reduced and individuals decreased in size with increasing depth. Generally, an enrichment in the meiofauna was evident from the coastal to the intermediate zone. Enrichments induced by urban pollution have been recorded previously, but not, as in the present study, at a distance of 1 km away from the outfall, succeeding to a highly impoverished coastal zone. Thus, by supplying organic matter and nutrients, Marseilles' sewage can enrich sediments, but only to a certain distance away from the outfall, where the deleterious effects of pollutants are attenuated.     

Some factors affecting the colonization of a dredged channel

Standing crop, population size, and species diversity of the macrobenthic organisms in an estuarine channel were studied before and after dredging. A new suction-corer which sampled an area 0.1 m² to a depth of 30 cm was used in order to insure the inclusion of large, deep-dwelling animals. Eleven months after dredging, biomass and number of species and specimens had not recovered to pre-dredging levels. Colonization began with relatively large, swiftly moving forms such as the errant polychaete Nereis succinea and the crab Neopanope texana sayi. Stations in silt and mud regions recovered more slowly than those in sandier sediments. Most of the dominant and subdominant species had not recovered 11 months after dredging, and the previously abundant polychaetes Notomastus latericeus and Clymenella torquata had virtually disappeared. Only relatively uncommon lamellibranchs such as Tellina agilis, Lyonsia hyalina and Mulinia lateralis increased after dredging. Distribution of sediment types changed as the result of modified tidal velocities in the channel. Mud and silt were removed by the dredge, exposing the sand underneath, and sandy stations became muddier as the result of lowered current velocities. Marked changes in species composition reflected this change in sediment character. Eleven months after dredging no evidence of succession was found, but colonization had begun. Values of all three parameters studied were reduced to small fractions of pre-dredging levels, although species diversity in sandy sediments exceeded pre-dredging levels.     

Oxygen/sulfide regime and nematode fauna associated with Arenicola marina burrows: new insights in the thiobios case

Profiles of oxygen and sulfide around the burrows of the lugworm, Arenicola marina, from a North Sea tidal flat were examined with microelectrodes, and the steep gradients were related to the microdistribution of nematodes. Around the tail shaft free oxygen penetrated only 2 mm into the burrow wall, coinciding with a bright zone sharply limited by the ambient black sediment. Contrastingly, in normal bottoms of the tidal flat (controls) only the surface of the bright zone was supplied with free oxygen. Here, the dark colouration coincided with the presence of free hydrogen sulfide. Around the tail shaft the nearest free hydrogen sulfide was detected 6 mm from the burrow wall leaving several millimetres of black sediment without measurable free sulfide. We discuss how these divergencies may relate to the stability of the oxygen/sulfide gradients and the course of time involved in their formation. A total of 54 nematode species were identified. Based on non-metric Multidimensional Scaling Ordination, four nematode assemblages corresponded to four microhabitats of the A. marina burrow: the funnel, the feeding pocket, the tail shaft and the feacal cast. The tail shaft assemblage (oxic plus partly anoxic zones) was similar to that of the anoxic zone of the control sediment. It was dominated by the most abundant nematode in the present study, Metalinhomoeus biformis (mean abundance in tail shaft 202 indx10 cm^-3). Adults of common nematode species from sulfidic microhabitats had a significantly higher length/diameter ratio than those inhabiting the oxic zone of the control sediment (p<0.001). The chemical recordings and metric analysis indicate that these slender nematodes around the A. marina tail shaft and in the reduced horizons of the reference sites represent thiobiotic assemblages, as compared to the shorter and stouter oxybiotic species characterising the assemblages from the surface zone and (partly) the funnel.     

Impact of heavy metals of industrial plant wastewater on benthic communities of Bizerte Lagoon (Tunisia)

ABSTRACT

The aim of this study was to assess the consequences of human impact on the characteristics of sediments heavy metal concentration, grain size and its influence on the structure of the microbial and meiofaunal community assemblages. A survey was carried out in July 2013 within six sites located in the Bizerte Lagoon (Tunisia), both downstream and upstream of industrial effluents. The highest total sediment metal concentrations were detected in stations located close to the industrial sewage discharge points. In these stations, the lowest densities of the total meiofauna (33?±?13?ind/10?cm^?2) and conversely the highest densities of cultivable bacteria that are heavy metal resistant have been reported (16?±?80.34?CFU?g^?1). Univariate (ANOVA) and multivariate (MDS/CCA) analyses demonstrate high dissimilarity (0.06) in meiofaunal and bacterial community structures between downstream and upstream industrial sewages. Furthermore, canonical correspondence analysis CCA results indicated that heavy metal sediment contamination promoted bacteria that are resistant to heavy metals, while heterotrophic bacteria supported the development of meiofauna taxa. The results highlight the importance of bacteria/meiofauna interactions, as both meiofaunal and microbial communities give indications of the ecological impact of heavy metal contamination in sediment.     

Feeding grounds for waders in the Bay of the Mont Saint-Michel (France): the Lanice conchilega reef serves as an oasis in the tidal flats

The tube-building polychaete Lanice conchilega can form dense populations, often called reefs, which promote benthic community change and constitute feeding grounds for secondary consumers. The aim of this study was to quantify the role of the L. conchilega reef of the Bay of the Mont Saint-Michel (BMSM) for feeding waders, by combining macrobenthos data, bird counts and bird diet information. Wader densities in the reef were on average 46.6 times higher than in non-reef areas. According to faecal analyses, waders in the reef mainly selected the accompanying fauna and especially crustaceans. The attractiveness of the reef to feeding birds may be largely explained by the high abundance, richness and biomass of macrobenthic species in the reef compared with the rest of the BMSM.     

Benthic fauna of a shallow-water gaseohydrothermal vent area in the Aegean Sea (Milos, Greece)

The benthic community of shallow-water hydrothermal vents (10 m water depth) in a sandy bay on the south coast of Milos, Greece was studied. The macro- and meiofauna, as well as environmental factors such as temperature, salinity, sulfide concentration, grain size composition, ATP concentration, organic carbon, chlorophyll a and phaeopigments were evaluated. Samples were taken along transects from a seagrass meadow into a hydrothermally active area. Hydrothermally active spots were distinguished from the surrounding sediment by their black sediment coloration and an overlying white bacterial mat. For comparison, a control transect from a seagrass meadow into bare sand in a non-vent area in the same bay was studied. Overall, we found decreasing faunal diversity from the seagrass bed towards the hydrothermally active area. Along this gradient, four different zones could be distinguished. (a) The seagrass area had a relatively low diversity compared to the control stations, but much higher diversity than in the areas directly influenced by hydrothermal activity. (b) The border zone between the seagrass and the hydrothermal area, with moderate sulfide concentrations, pH values, seawater salinity and temperatures, was inhabited by an impoverished community dominated by the polychaete Capitella capitata, the nematodes Oncholaimus campylocercoides and Sabatieria sp., and an as yet undescribed chromadorid nematode (Chromadorina sp.). (c) The transition zone at the border of the hydrothermal area proper, with temperatures and sulfide concentrations higher than in the border zone, was characterized by high salinity (60‰) and low pH values. Along with the gastropod Cyclope neritea, single specimens of C. capitata and O. campylocercoides occurred. (d) The hydrothermal area proper, characterized by high sulfide concentrations, had high sediment temperatures and high salinities. The pH was as low as 6. Here, the gastropod Cyclope neritea was the only species occurring in considerable numbers. The gaseohydrothermal activity seems to largely exclude the surrounding fauna in a graded zonal pattern. The increasing hydrothermal stress entrains a change of the ambient biota towards a depauperate consortium of tolerant, opportunistic species, e.g. Capitella capitata (Polychaeta) and Oncholaimus campylocercoides (Nematoda). Although the hydrothermal area proper was characterized by a thick bacterial mat, the benthic community was characterized not by bacteria feeders but by scavenging species. Received: 13 September 1996 / Accepted: 24 October 1996