Occurrence of tri-n-butyltin-caused imposex in the North Pacific marine snail Nucella lima in Auke Bay,Alaska

Like female dog whelks (Nucella lapillus) in the Atlantic Ocean, females of the Pacific gastropod N. lima respond to low concentrations of tri-n-butyltin (TBT) by growing a penis and vas deferens. This condition, termed imposex, was found to occur in N. lima collected from August 1987 to May 1988 along a TBT pollution gradient associated with a marina in Auke Bay, Alaska. The suite of symptoms characteristic of imposex in N. lima was slightly different than in N. lapillus. Imposex, as measured by relative penis size (RPS) of females to males, increased from 0.0 to 34.27 along this gradient and as measured by vas deferens sequenceindex (VDS) increased similarly from 0.0 to 4.29. Concentrations of TBT in N. lima increased from below detection limits (about 0.010 g Sn g^-1 dry tissue wt) to 0.065 g Sn g^-1 dry tissue wt along the gradient. The gradient was determined by measuring TBT in whole-body tissues of bay mussels (Mytilus edulis). Concentrations of TBT in mussels increased from below detection limits to 0.833 g Sn g^-1 dry tissue wt in mussels from within the marina that was the major local source of TBT. Imposex, tissue TBT burden, and position along a TBT pollution gradient are significantly correlated (P<0.01) in N. lima. TBT was tested as a causative agent for imposex by exposing snails at a distant control site to TBT paint. After 1 mo exposure, 33% of the females grew a penis ranging in size from 0.2 to 0.8 mm. Our results generally corroborate those found for N. lapillus and indicate that imposex in N. lima is caused by environmental TBT exposure. We suggest that the RPS in the genus Nucella may be useful in monitoring TBT in coastal waters worldwide.     

Hinia reticulata and Nucella lapillus. Comparison of two gastropod tributyltin bioindicators

Hinia reticulata and Nucella lapillus (collected from 1989 to 1991 at Pointe de Pléneuf and Méan Mélen, France, respectively) exhibit imposex in response to tributyltin (TBT) pollution leached from antifouling paints and proved to be good TBT bioindicators. H. reticulata was kept for 18 mo under TBT-free conditions in the laboratory, but no evidence for imposex remission was found. A second series of experiments showed that food chain uptake of TBT from a contaminated diet is an important mode of TBT exposure in H. reticulata. Comparative tank experiments with H. reticulata and N. lapillus demonstrated that the same type of TBT exposure resulted in comparable TBT body burdens, biological concentration factors, and imposex development in both species [measured as increase of VDS (vas deferens sequence index), uncubed RPS (relative penis size index) and average female penis length]. Differences in imposex development of natural and laboratory populations are discussed against the background of different types of TBT contamination of their food. A statistical study, based on an analysis of natural populations of both prosobranch species, makes a comparison of the specific TBT sensitivity of the two bioindicators possible. As a consequence it is proposed that TBT biomonitoring programs in Europe should use both prosobranchs as indicator species.     

The morphological expression of imposex in Hinia reticulata (Gastropoda: Buccinidae): a potential indicator of tributultin pollution

Specimens from the prosobranch Hinia reticulata collected along the coast of Brittany and Normandy from 1988 to 1991 exhibited imposex (occurrence of male parts in addition to the female genital duct) in response to tributyltin (TBT) pollution. Four stages of imposex development (1 to 4) with two different types in the Stages 1, 3 and 4 could be distinguished and have been documented with scanning electron micrographs for the first time. Furthermore, three additional alterations of the genital tract are shown. Neither TBT-induced sterilization nor sex change occurred. TBT accumulation in whole body and selected tissues is described and sex-related differences are shown. The VDS (vas deferens sequence) index, cubed and uncubed RPS (relative penis size) index and average female penis length of a population were analysed with regard to their quality as indices for TBT biomonitoring. Based on the way imposex develops in this species the VDS is recommended as the most valid index. Only in highly polluted areas should the uncubed RPS be used as a second index.     

Experimental prey species preferences of Hexaplex trunculus (Gastropoda: Muricidae) and predator–prey interactions with the Black mussel Mytilus galloprovincialis (Bivalvia: Mytilidae)

Hexaplex trunculus is one of the most widespread Mediterranean species of muricid gastropod and lives on rocky, sandy-mud and mud substrata. Although common in the Adriatic Sea, relatively little is known about its ecology especially feeding behaviour. The aim of this study was to explore the aspects of the feeding behaviour of H. trunculus using Arca noae, Modiolus barbatus and Mytilus galloprovincialis as experimental prey. Prey species preference, predator size, prey size choice, feeding rates, handling times and mode and place of attack were analysed. Typically, only M. galloprovincialis was attacked: A. noae rarely at the byssal gape and M. barbatus never. Small (40 mm) H. trunculus could not easily attack large M. galloprovincialis (65 mm) and preferred small (20 mm) and medium (35 mm) sized prey. Conversely, medium (55 mm) and large (70 mm) H. trunculus fed randomly on M. galloprovincialis of all three sizes. The feeding strategy adopted by H. trunculus individuals varied with respect to size. A tendency to drill the prey shell was recorded for small predators, whereas marginal chipping was adopted more frequently by large individuals. On average small, medium and large H. trunculus consumed 2.4±1.6 (range 0–4), 1.2±1.6 (range 0–4) and 2.0±2.1 (range 0–6) M. galloprovincialis, respectively. There was a statistically significant difference in prey-handling time with respect to the method of access adopted, predator and prey sizes. The time required to access a M. galloprovincialis individual by marginal chipping was considerably less than that required for drilling. H. trunculus consumed an average of 0.60±0.80 g M. galloprovincialis tissue dry weight over a 5-week period, that is, ~40% of its own tissue body weight. This translates to an average-sized (55 mm shell height) H. trunculus consuming ~18 M. galloprovincialis of 50 mm shell length (minimum marketable size) per year. H. trunculus showed no preference to drill either the left or right valves of M. galloprovincialis but generally attacked the posterior shell margin.     

Rapana venosa as an indicator species for TBT exposure over decadal and seasonal scales

Imposex and decline in reproductive output in marine gastropods have been linked to tributyltin (TBT) exposure. This study describes Chesapeake Bay, USA, veined rapa whelk Rapana venosa imposex incidence and sex ratios from 1998 to 2009. Tissue TBT concentrations (ng g^?1) were examined with respect to whelk sex, size, and water temperature at the time of collection, and also to egg case size, hatching success, and veliger diameter. Imposex incidence declined and population sex ratios moved closer to parity from 1998 to 2009. Observed TBT concentrations (ng g^?1) were higher in James River than in Ocean View whelks with higher TBT concentrations observed in males than in females. Exponential declines in TBT concentrations from female-specific first to last clutches within a reproductive season were observed, indicating that whelks depurate TBT through egg case deposition. Egg capsule hatching success and veliger size were similar for female and imposex whelks. The R. venosa imposex levels observed in Chesapeake Bay apparently do not affect the production, release, or viability of larvae.     

Monitoring the effectiveness of the European tributyltin regulation on the Basque coast (northern Spain) by assessing imposex in two gastropod species (Nassarius reticulatus and Nassarius nitidus)

Imposex and butyltin body burden were assessed in 2011 along the Basque coast (northern Spain) in two gastropod species (Nassarius reticulatus and Nassarius nitidus) four years after an initial survey in 2007. The aim of this re-survey was to monitor the effectiveness of the European ban on the use of tributyltin (TBT) based antifouling paints on ships’ hulls (EC 782/2003). Imposex levels in 2011 were lower than those determined in 2007 at most of the sampling sites. Accordingly, TBT concentrations in the female body burden of Nassarius reticulatus varied from 43 to 250 ng Sn/g in dry weight in 2011, which was a lower maximum than in 2007. Nevertheless, the results for the butyltin degradation index suggest that there have been recent inputs of TBT within the two main Basque harbours. Overall, the legislative measure is contributing to the reduction of TBT effects on the Basque coast although its presence is still evident.     

Population structure and growth of Donax trunculus (Bivalvia: Donacidae) in the western Mediterranean

A comparison of shell growth in Donax trunculus (collected between 1988 and 1990 of Cullera, Spain) has been carried out using an analysis of cohort progression in monthly length frequency distributions, hyaline surface shell growth rings and internal microgrowth bands. In the Mediterranean there are two periods of recruitment of D. trunculus, one in the summer (July to September) and the other in winter (December to February). Clams recruited to the population in winter display a clear cessation in shell growth during the following summer which may possibly be correlated with spawning, whereas individuals of the summer recruited cohort show no growth cessation the following summer and continue to deposit shell during this period. The normally opaque shell of D.trunculus reveals the presence of translucent hyaline growth rings when the shells are backlit by a strong light source, and these have been shown to be laid down in the shell during summer months. Formation of a hyaline ring is accompanied by a narrowing of the microgrowth patterns present in shell sections. Both the hyaline rings and the length frequency distributions have been used to determine the age and growth rate of D. trunculus.     

Observations on the biology of Donax trunculus and D. vittatus at Ile d'Oléron (French Atlantic coast)

Observations have been made on the biology of two species of the bivalve genus Donax, on two beaches, Vertbois and St. Trojan, on the Ile d'Oléron, French Atlantic coast. The two species displayed different vertical distribution: D. trunculus Linné in the intertidal; and D. vittatus (da Costa) between low-water spring tide (LWST) and 5 to 6m depth, with only the fringe of the population extending to the shore. D. trunculus also showed differential distribution by size or age, the youngest individuals being highest on the shore, and the oldest near LWST. The two species exhibited a similar pattern of seasonal recruitment and growth, but in both there were differences in growth characteristics between the two beaches, individuals at St. Trojan growing to a larger size than those at Vertbois. The mean pattern of mortality in the two areas was similar, and productivity per 100 recruits surviving to the first winter following settlement was greater at St. Trojan than at Vertbois. For D. trunculus the mean population density at Vertbois was higher and the production per unit area on the two beaches was similar. Production to biomass (B) and elimination to biomass (B) ratios were similar for the two beaches for both species, and while production was greater for D. trunculus at both beaches, the B and B ratios were higher for D. vittatus. Comparisons with other published data reveal that the rate of growth of D. trunculus is similar throughout its range from Brittany to the Mediterranean Sea, while that of D. vittatus shows more variation. In both species, there is a trend of increasing P/B and E/B ratios from north to south throughout the geographical range.     

Origin and nature of spermatophoric mass of the spiny lobster Panulirus homarus

In the spiny lobster Panulirus homarus (Linnaeus) the spermatophoric mass consists of a highly convoluted tube, containing the sperm mass and a gelatinous matrix. The spermatophoric components have their origin in the proximal vas deferens and the completed spermatophoric mass is stored within the distal dilated part of the vas deferens. In the proximal vas deferens the inner glandular epithelial cells give rise to leaf-like typhlosoles which gradually diminish in size and finally disappear in the distal vas deferens. The secretions originating from the typhlosole form the outer gelatinous matrix of the spermatophoric mass. Histochemical observations reveal that the wall of the spermatophore tube consists of neutral mucopolysaccharide, whereas the sperm mass and the gelatinous matrix are rich in acidic mucopolysaccharides. Both polyacrylamide gel electrophoresis and agarose gel electrophoresis were used to separate the acidic mucopolysaccharide (AMPS) complexed with the proteins and the simple proteins. In addition, AMPS have been isolated following the method of Rahemtulla and Lovtrup (1974) and used for electrophoresis. In both the polyacrylamide gel electrophoresis and agarose gel electrophoresis, the extracted acid mucopolysaccharides give only a single fraction which, in comparison with the standard acid mucopolysaccharides, corresponds to chondroitin sulphate. The functional significance of the chondroitin sulphate and neutral mucopolysaccharides in the spermatophoric components of P. homarus is discussed in relation to their functional role in spermatophore hardening and protection of the delicate spermatozoa during their prolonged storage on the sternum of the female.     

Male reproductive biology of the red frog crab,Ranina ranina,off Hachijojima,Izu Islands,Japan

Male red frog crabs, Ranina ranina, were collected year round in 1990 and 1991 off Hachijojima for histological study of the reproductive system and cycle. The testis containing the lobules and seminiferous ducts is surrounded by connective tissue. The seminiferous duct connects to the anterior end of the vas deferens, which can be histologically divided into three portions similar to one another in appearance. It was surrounded by fibrous connective tissue, muscle fibrils and columnar epithelium. Muscle fibrils were absent in the anterior portion. Multiple sperm masses were not formed in the vas deferens and ejaculatory duct, but the sperm mass was covered with a capsule composed of two layers. The outer layer of the capsule was periodic acid Schiff (PAS)-positive, but the inner layer was negative. Both layers were Alcian Blue negative, except the vacuoles in the outer layer that were stained blue. The small round androgenic gland was attached to the posterior end of the vas deferens of the coxa of the eighth thoracic appendage. The ejaculatory duct was distinguishable from the vas deferens by the absence of columnar epithelium and the presence of thick longitudinal muscle fibers. Spermatogenesis was histochemically examined. The acrosomal vesicle appeared to be derived from PAS-positive vesicles in the cytoplasm of the spermatid at the early stage of spermiogenesis. The arms were positive to the Feulgen reaction and the subacrosomal region was negative to PAS. Seasonal changes in reproductive cycle were inconspicuous histologically and microscopically. Sperm were always present in the testis and vas deferens throughout the year and occupied 5.1 to 19.6% of testis observed in cross sections. The minimum size of maturity is less than 39 mm carapace length, but the minimum size capable of successful mating was estimated to be ca. 55 mm.     

Potential histopathological and molecular changes in rat vas deferens inhaled by Boswellia papyrifera and Boswellia carterii

Boswellia papyrifera and Boswellia carterii released from smoke contaminate indoor environment and consequently adversely affect humans as evidenced by respiratory disturbances. The aim of this study was to determine the effects of these plants on pathological and biochemical changes in vas deferens of albino rats. Animals were administered 4 g/kg body weight B. papyrifera and B. carterii daily for 120 days along with controls. Significant changes were observed in epithelial cell types and some cells showed signs of degeneration. The ultrastructural studies revealed marked changes in cytoplasmic organelles. Microvilli were missing and lysosomes were found in the cytoplasm. In addition, all treated groups plasma fructose and other biochemical parameters were decreased indicating reduced energy necessary for motility and contractility of spermatozoa. Many spermatozoa were disorganized and agglomerated. Data suggest that smoke from these plants adversely affects vas deferens.     

An updated scheme of imposex for<Emphasis Type="Italic"> Cantharus cecillei</Emphasis> (Gastropoda: Buccinidae) and a new mechanism leading to the sterilization of imposex-affected females

The morphological expressions of imposex in gastropods have been classified into seven stages (S₀–S₆) with three types (a, b and c). This typical imposex scheme, however, fails to cover some new findings such as specimens with the proximal vas deferens section passing by the vaginal opening (VDSP) and aborted egg capsules. In the present study, Cantharus cecillei was sampled from the coastal waters of China from May to November in 2003. This species showed all the morphological expressions in the typical imposex scheme with the exception of S_1b and S_2b. In addition, specimens with VDSP occurred in all stages other than stage 4, and were documented for the first time with scanning electron micrographs or in histological sections. Besides VDSP, which was observed exteriorly, excrescent tissue, e.g. the vas deferens, also originated in the ventral channel or the inner wall of the pallial oviduct was extruded. Based on these new findings, the typical imposex scheme was modified to obtain one valid for C. cecillei. Types a*, b* and c* were added as new types corresponding to types a, b and c, respectively. S_4* was also rearranged to correspond to S₄; S_4*, thus, no longer represented the end of imposex development. The updated scheme was not only valid for describing diverse expressions of imposex development in C. cecillei and Nassarius reticulatus, but could also be generalized to define various forms of VDSP in other species. Histological observation indicated that egg capsules would be blocked by the tissues of the vas deferens section or the extrusion of the pallial oviduct wall. This new mechanism, leading to the sterilization of imposex-affected females, was first conclusively revealed and described in the present scheme.Communicated by T. Ikeda, Hakodate     

Donax trunculus and Venus verrucosa as bioindicators of trace metal concentrations in Mauritanian coastal waters

Trace-metal concentrations (Cd, Cu, Fe, Mn and Zn) were investigated in two species of abundant filter-feeding molluscs from Mauritania, the wedge shell Donax trunculus L. and the clam Venus verrucosa L. D. trunculus were collected on the beach of Nouakchott at low tide and V. verrucosa were sampled at the Banc d'Arguin at depths varying from 10 to 20 m, in mid-February 1987. D. trunculus contains significantly higher Fe and Zn concentrations than the clam. Total trace-metal content per individual (Y) in D. trunculus could be related to body weight (W) as the power function Y=aW ^b . The subsequent regression coefficient b was >1 for Cu, Fe, Mn and Zn and >1 for Cd. Metal concentrations in both species decreased in the order: Fe>Zn>Mn>Cu>Cd. Comparison of trace-metal concentrations in the small D. trunculus and the large V. verrucosa revealed lower contents in V. verrucosa, except for Cd. The trace-metal distribution in the various organs of V. verrucosa indicates that gills seem to concentrate metals, except Cu, to a greater degree than the other organs (visceral mass and remainder). The relatively elevated content of Fe in D. trunculus and of Cd, Fe and Zn in the gills of V. verrucosa seem to be of natural origin. In view of the scarcity of Mytilus sp. along the Mauritania coast, the molluscs D. trunculus and V. verrucosa are proposed as useful bioindicators of trace-metal concentrations.     

Increased sperm numbers in the vas deferens of meadow voles, <Emphasis Type="Italic">Microtus pennsylvanicus,</Emphasis> in response to odors of conspecific males

Sperm competition occurs when the sperm of two or more males compete to fertilize the egg/s of a particular female. Males of some species respond to a high risk of sperm competition by increasing the number of sperm in their ejaculates. Males may accomplish such a response by increasing the intensity or duration of contraction of the cauda epididymidis and vas deferens. During emission (first phase of the ejaculatory process), the vas deferens receives sperm from the cauda epididymidis and propels the sperm to the urethra. In this paper, we tested the hypothesis that males exposed to a high risk of sperm competition mobilize larger numbers of sperm from the cauda epididymidis to the vas deferens before initiation of copulatory behavior. This accumulation of sperm in the vas deferens would result in a larger number of sperm in the ejaculate. To test this hypothesis, we exposed male meadow voles, Microtus pennsylvanicus, to either low or high risks of sperm competition using soiled bedding of conspecific individuals. At three different times after this exposure (15, 30, or 60 min), we removed both vasa deferentia and counted the sperm within them. We found a significant increase in sperm numbers in the vas deferens of males after 30 min of being exposed to a high risk of sperm competition. The lower sperm numbers after 15 and 60 min of exposure suggest that the observed response is relatively slow and that sperm mobilized to the vasa deferentia may return to the cauda epididymides if ejaculation does not occur some time after the observed response. Our results indicate that the physiological response that may result in high sperm numbers in the ejaculate in relation to high risk of sperm competition can occur before initiation of copulatory behavior.     

Survival of Nucella lapillus in a tributyltin-polluted area in west Brittany: a further example of a male genital defect (Dumpton syndrome) favouring survival

A male genital defect was reported in a Nucella lapillus (L.) population at Dumpton Gap (England). This defect was termed Dumpton syndrome (DS) and appears to be a genetic feature. Its main characteristic is the absence of penis (aphally) in males and in females. In 1992, such a phenomenon was discovered in populations in the vicinity of Brest (Brittany, France). DS-affected females exhibit fewer tributyltin-induced imposex characteristics than expected in normal individuals. The percentage of female sterilization is thus lower, favouring population survival. In consequence, the DS is considered to be a pollution-resistance feature. Comparison with the Dumpton population revealed similarities and differences in the DS characteristics. It is thus hypothesized that the DS observed at the two locations is due to two different biological mechanisms. Indeed, aphallic males with a split prostate were observed but no underdevelopment of their vas deferens and testis was noted in the present study. In this gonochoristic gastropod species, the most DS-affected males in Brest possess an ovotestis and it is thus hypothesized that feminity is remnant in N. lapillus. Incidence of abnormality is ten times higher in females than in males. This suggested that a sex-difference operates in the Brest phenomenon.     

The androgenic gland of the shrimp Palaemon dayanus

The androgenic gland of Palaemon dayanus Henderson differs in morphology and anatomy from that of all other known natantian decapods. The gland is attached to the distal extremity of the was deferens as a thin, plate-like structure made up of numerous parallel strands in close juxtaposition. The gland is clearly separated by intervening thick muscle bands from the epithelium of the vas deferens. The structure of the gland shows distinct zones of activity such as growth and degenerating zones. The presence of degenerating zones together with zones of activity in the same gland at the same time is attributed to a holocrine mode of secretion in the androgenic gland.     

Vitellogenesis in the deep-sea shark Centroscymnus coelolepis

At present, information on the reproductive physiology of Centroscymnus coelolepis, which is one of most important and widespread deep-sea shark species, is completely lacking. In this study, we investigated vitellogenesis, a key step in the reproduction biology of fishes. Specimens of C. coelolepis were collected at 2850 m depth in the Western Mediterranean Sea. The size of the collected sharks (range: 35.5–65.0 cm TL) was much lower than those typically reported for the Atlantic and Pacific Oceans. The marked distinctiveness of Mediterranean and Atlantic/Pacific populations was reflected by the achievement of sexual maturity at a smaller size in Mediterranean specimens. The examination of cytoplasmatic components of oocytes indicated that vitellogenin uptake in the ovary started when oocytes reached 14 mm in diameter. Only reproductive females displayed a significant relationship between plasmatic vitellogenin and gonadal development, suggesting that vitellogenesis in C. coelolepis is a discontinuous process. Oestradiol levels were tightly coupled with gonadal development, underlining the importance of this hormone in controlling vitellogenesis. All these findings suggest that vitellogenesis in this yolk-sac viviparous shark might occur with similar mechanisms of oviparous vertebrates.     

Organotin pollution in Malta coastal zone

The speciation of organotins in seawater, surface microlayer and sediments in the marine environment of Malta is described. Concentrations of tributyltin (TBT) in bulk seawater inside the harbours were as high as 0.3 μg Sn L^‐1 but were below detection limits (5 ng Sn L^‐1) in open sea, 1.6 km offshore. In sediments, TBT levels were highest for the yacht marinas and ranged between 0.03 and 1.5 ug Sn g^‐1. Dibutyltin is more common in the surface microlayer than TBT. Diphenyl and monophenyltin were found infrequently in bulk seawater and in sediments. Tetrasubstituted organotins, namely, Me_nBu₍₄__n)Sn, where n = 1, 2, and 3, were found frequently in TBT‐contaminated sediments (0.1–9μg Sng^‐1), in seawater and in the microlayer where concentrations as high as 140 μg Sn L^‐1 (Me₃BuSn) were measured. Direct environmental methylation of TBT and that of its debutylated analogues may play a significant role in the geochemical cycling of tin under certain environmental conditions.     

Growth parameters of deep-water decapod crustaceans in the Northwestern Mediterranean Sea: a comparative approach

 Relative and absolute growth were studied in 17 species of deep-water decapod crustaceans, spanning nine families of six different infra-orders, in the Northwestern Mediterranean Sea. The overall maximum abundance of these species lay between 200 m and 750 m (i.e. upper- and mid-slope species). Relative and absolute growth rates were compared by contrasting the slopes of the size–weight relationships for the different species and calculating the von Bertalanffy growth-equation parameters asymptotic length (L _∞ ) and growth rate (k). The size–weight relationships differed significantly as function of the species' life habits. The results revealed a significant decrease in weight relative to size in mesopelagic species (which carry out diel vertical migrations), an almost isometric relationship between size and weight in the less mobile nektobenthic species, and a significant increase in weight relative to size in strictly benthic species. The mean allometric coefficient for each group increased significantly from mesopelagic to benthic species. However, no general trend was observed in the growth-performance index, Φ (an index used to compare absolute growth rates between species, as a function of habit and depth of maximum abundance for all species combined), suggesting that the deep-water decapod crustaceans studied have similar absolute growth rates. Nevertheless, comparison of growth-parameter and growth-performance index values within families did reveal differences. Mesopelagic species of the families Sergestoidae and Pasiphaeidae showed slightly increased growth rates with increasing depth of distribution. Nektobenthic species of the genus Plesionika followed a trend opposite to that shown by mesopelagic species, with a higher growth rate for the shallowest-dwelling species (P. heterocarpus) than the deepest-dwelling species (P. acanthonotus). Taking growth as one of the major components of an organism's energy budget, the growth rates for the decapod crustacean species in this study were significantly lower than those reported in the literature for shallow-water penaeid crustacean species (which are distributed in higher-temperature habitats than deep-water Mediterranean crustaceans) and higher than those reported for mesopelagic myctophid fish species. Hence, the well-defined growth trends shown by deep-water decapod crustacean species in the Northwestern Mediterranean Sea, compared to the less well-defined trends in the other taxa, is discussed in the framework of the overall dynamics of their ecosystem. Received: 25 May 1998 / Accepted: 27 September 1999