Sand stress as a non-determinant of habitat segregation of indigenous (Perna perna) and invasive (Mytilus galloprovincialis) mussels in South Africa

Periodical sand inundation influences diversity and distribution of intertidal species throughout the world. This study investigates the effect of sand stress on survival and on habitat segregation of the two dominant mussel species living in South Africa, the invasive Mytilus galloprovincialis and the indigenous Perna perna. P. perna occupies a lower intertidal zone which, monthly surveys over 1.5 years showed, is covered by sand for longer periods than the higher M. galloprovincialis zone. Despite this, when buried under sand, P. perna mortality rates were significantly higher than those of M. galloprovincialis in both laboratory and in field experiments. Under anoxic condition, P. perna mortality rates were still significantly higher than those for M. galloprovincialis, but both species died later than when exposed to sand burial, underlining the importance of the physical action of sand on mussel internal organs. When buried, both species accumulate sediments within the shell valves while still alive, but the quantities are much greater for P. perna. This suggests that P. perna gills are more severely damaged by sand abrasion and could explain its higher mortality rates. M. galloprovincialis has longer labial palps than P. perna, indicating a higher particle sorting ability and consequently explaining its lower mortality rates when exposed to sand in suspension. Habitat segregation is often explained by physiological tolerances, but in this case, such explanations fail. Although sand stress strongly affects the survival of the two species, it does not explain their vertical zonation. Contrary to our expectations, the species that is less well adapted to cope with sand stress maintains dominance in a habitat where such stress is high. GI Zardi, KR Nicastro contributed equally to the work     

Ecological aspects of ophiuroids from the phytal of S. W. Atlantic Ocean warm waters

Nine species of ophiuroids — Amphipholis januarii, Axiognathus squamatus, Hemipholis clongata, Ophiocnida scabriuscula, Ophiactis lymani, Ophiactis savignyi, Ophionereis reticulata, Ophioderma appressum and Ophiothrix angulata —were collected in various habitats from the mid-littoral and upper infralittoral on the northern coast of the State of São Paulo, Brazil. The abundance of 6 species occurring in 28 species of algae was investigated. A total of 220 samples was collected at monthly intervals from April, 1969 to May, 1970. The distributional pattern of the populations, as determined by Mobisita's index, shows aggregation of the ophiuroids. No ophiuroid species was limited to one algal species. Each species prefers certain algal species and/or a different level on the shore. The most abundant species on algae was Ophiactis lymani, with a maximum density of 1,510 individuals/100 ml of algae. Four species of ophiuroids occurred inside sponges. The most abundant species in sponges was O. savignyi, with a maximum density of 1,892 individuals/100 g of dry sponge. Two species of ophiuroids occurred inside the bryozoan Schizoporella unicornis. Feeding observations have shown Axiognathus squamatus to capture its food by mucous nets. Decapod crustaceans fed on ophiuroids in aquaria. The copepod Thespesiopsyllus sp. was found in the body cavity of the 2 Ophiactis species. Simultaneous sexual and asexual reproduction were abserved in O. savignyi.     

Hematological studies on some Nile fishes,Tilapia nilotica,Lates niloticus,and Labeo niloticus

Hemoglobin content of 3 Nile fishes — Tilapia nilotica Linnaeus, Lates niloticus Linnaeus and Labeo niloticus Forskal— were examined. Clear differences in hemoglobin and hematocrit values were observed, and seasonal variation in the hemoglobin content of Lates niloticus and Labeo niloticus were noticed. There is a correlation between hematocrit value and hemoglobin content of the 3 species examined. The chloride content is higher in the species with lower hemoglobin content.     

Community responses to UV radiation. II. Effects of solar UVB on field-grown diatom assemblages of the Carribean

The effects of ambient solar UV on community productivity and structure were assessed during primary succession of benthic diatoms on artificial substrate in a coral reef of the Caribbean. Artificial substrates, partially enclosed by UV cutoff filters, were placed at a depth of 60 cm below surface. During the initial colonization stages, the assemblages exposed to the full solar spectrum had a mean productivity 43.4% lower than the assemblages exposed to PAR+UVA only. Some differences in species diversity of assemblages under the different UV treatments were also observed. Sensitive species to UVB exposure were Mastogloia angulata, M. ovata, M. paradoxa, Nitzschia longissima, Plagiogramma staurophorum, Rhopalodia musculus, and Surirella ovata. These UVB effects gradually diminished as succession proceeded; 5 to 6 weeks after the placement of the substrates in the water, no significant differences in productivity were observed between the different treatments, while after 6 weeks of growth, species diversity and evenness were higher, although not statistically significant, in the UVB-exposed assemblages. During the first 2 weeks of growth, the productivity under PAR+UVA was significantly lower than that under PAR only. Received: 12 July 1996 / Accepted: 20 November 1997     

Lingual and biting responses to prey chemicals by ingestively naive scincid lizards: discrimination from control chemicals,time course,and effect of method of stimulus presentation

Summary. We tested responses to prey chemicals by lizard hatchlings of an oviparous species and neonates of a viviparous species, neither of which had never eaten. Both species responded more strongly to prey chemicals than to odorous and odorless control stimuli presented on cotton swabs. Although only a few species have been examined, all that have been tested have an innate capacity for prey chemical discrimination, suggesting that this innate response to prey chemicals is widespread among lizards that use the lingual-vomeronasal system to locate and identify prey. Innate prey chemical discrimination has the great advantage of permitting lizards lacking prior experience with food to respond appropriately to chemical cues associated with food. Both species discriminated prey chemicals from control substances at age three days, earlier than previously known. Our data hint that Mabuya macularia may be capable of discrimination on its day of birth, but further study is needed to determine the exact onset. A stronger tendency to attack swabs bearing prey chemicals by Scincella lateralis than by M. macularia may be explained by differences in defensiveness near an experimenter or by differences in the importance of visual prey cues for confirmation of chemical cues in the natural habitats of these species. In M. macularia responses to the control stimuli declined over days of testing, suggesting habituation, but responses to prey chemicals did not habituate by the third day of testing, which is interpreted as a possible adaptive response to permit location of food. In the standard method of stimulus presentation, a cotton swab bearing a chemical stimulus is placed anterior to a lizard's snout. We tested a new method in which the swab was placed in continuous contact with the lizard's anterior labial scales. The new method elicited significantly stronger responses from M. macularia. We discuss reasons for this finding and applications for the new method. Received 2 September 1999; accepted 15 December 1999     

Sensitivity and acclimation to UV radiation of zoospores from five species of Laminariales from the Arctic

Spores of five Laminariales from Arctic Spitsbergen were exposed in the laboratory to photosynthetically active radiation (PAR; 400–700 nm), PAR+UVA radiation (UVAR; 320–400 nm) and PAR+UVAR+UVB radiation (UVBR; 280–320 nm). Subsequently, germination was monitored over periods of 3, 6 and 9 days. The investigated species were the upper sublittoral Saccorhiza dermatodea, the upper to mid-sublittoral Alaria esculenta and Laminaria digitata, the mid-sublittoral L. saccharina and the lower sublittoral L. solidungula. The germination capacity decreased sharply after 16 h exposure to PAR+UVAR+UVBR in all species. However, S. dermatodea was able to recover from the damaging effects of UVBR. There was also a small increase in percentage germination of A. esculenta 6–9 days after the treatment. No recovery was evident in the other species. After 8 h exposure to PAR+UVA+UVB, L. digitata recovered completely, and L. saccharina and L. solidungula, partially. The only species susceptible to PAR+UVAR was L. solidungula. One prominent cytological feature of UVR-exposed spores was the enlargement of phenolic vesicles (physodes) (particularly seen in S. dermatodea and A. esculenta), which may have a protective function against UVR. Pilot experiments under natural irradiance conditions indicate that the PAR component of solar radiation exerts an additional stress. Overall the data show that zoospores of the species from the upper sublittoral are less sensitive to UVR or have the capacity to recover from UV stress in contrast to species from deeper waters, probably due to their UV protective and repair capabilities.Communicated by O. Kinne, Oldendorf/Luhe     

Variation in composition of predator-attracting allelochemicals emitted by herbivore-infested plants: Relative influence of plant and herbivore

Summary During foraging, natural enemies of herbivores may employ volatile allelochemicals that originate from an interaction of the herbivore and its host plant. The composition of allelochemical blends emitted by herbivore-infested plants is known to be affected by both the herbivore and the plant. Our chemical data add new evidence to the recent notion that the plants are more important than the herbivore in affecting the composition of the volatile blends. Blends emitted by apple leaves infested with spider mites of 2 different species,T. urticae andP. ulmi, differed less in composition (principally quantitative differences for some compounds) than blends emitted by leaves of two apple cultivars infested by the same spider-mite species,T. urticae (many quantitative and a few qualitative differences). Comparison between three plant species — apple, cucumber and Lima bean — reveals even larger differences between volatile blends emitted upon spider-mite damage (many quantitative differences and several qualitative differences).     

Form-specific fragances fromOphrys insectifera L. (Orchidaceae) attract species of different pollinator genera. Evidence of sympatric speciation?

Two closely related forms ofOphrys insectifera were observed in the field to attract different pollinator species selectively.O. i. ssp.insectifera attracted males of two species ofArgogorytes (Sphecidae, Hymenoptera Aculeata) andO. i. ssp.aymoninii attractedAndrena combinata males (Andrenidae, Apoidea, Hymenoptera Aculeata). A third form,O. aff.i. ssp.insectifera, attracted none of these three species. Volatile compounds from flowers and inflorescences of the three forms (originating from Öland, Sweden, and Aveyron, France) were collected, using entrainment, enfleurage, and solvent extraction techniques, and identified by gas chromatography — mass spectrometry. Scent differences between the three forms were confirmed in the amounts of aliphatic hydrocarbons (C₁₁–C₁₉), methyl esters (C₁₄–C₁₈), short chain aliphatic 1-alcohols (C₆–C₁₂), and monoterpene alcohols (C₁₀).     

Distribution and abundance of marine nematodes on the kelp Macrocystic integrifolia

The species and distribution of nematodes on the kelp Macrocystic integrifolia Bory, in the Bamfield region of Barkley Sound, British Columbia, Canada were examined. Nine species (belonging to six families) of nematodes were found on the kelp blades. Three species (Monhystera disjuncta, M. refringens and Prochromadorella neapolitana) comprised 91–99% of the nematode fauna and occurred in all monthly samples from July 1978 to November 1979. The three dominant species exhibited seasonal density differences. P. neapolitana occurred mostly in the summer, M. refringens abundance peaked from July to October and M. disjuncta was relatively abundant throughout the year. The other species contributed little to the overall abundance and distribution patterns. All three species occurred in greatest abundance on the lower and middle blades of M. integrifolia in the deep end of the kelp bed. Very few individuals occurred on the upper blades. Nematode distribution on M. integrifolia appeared to be related to blade age and the associated food sources on the blade.     

Gastropod predation on egg cases of skates (Chondrichthyes,Rajidae) in the southwestern Atlantic: quantification and life history implications

Mean predation rates (± SD) on egg cases of the skates Bathyraja macloviana, B. albomaculata, Amblyraja doellojuradoi, and Psammobatis spp. from the southwestern Atlantic were estimated to be 0.151 (±0.230), 0.423 (±0.344), 0.254 (±0.390), and 0.150 (±0.288), respectively. These estimates are within the ranges reported earlier (14–40%). Egg cases of B. albomaculata were preyed on in higher proportion than expected from their abundance and suffered a heavier predation rate where the snail Trophon acanthodes was present. Predation rates were not correlated with the thickness of the egg case wall, which indicates that other factors (ecological or chemical) could explain this pattern. Five types of boreholes were found in the egg cases: one was attributable to muricid gastropods, one to naticid gastropods, and a third type to an unknown gastropod (probably Fusitriton magellanicus); the remaining were of unknown origin. Cladistic analyses showed that skates are secondarily oviparous and have maximized adaptations for living in deep water. We suggest that oviparity in skates appeared as an adaptation to maximize fecundity (40–160 eggs per year, as compared to 2–18 pups annually or biannually in viviparous guitarfishes, the plesiomorphic sister clade of skates). If a predation rate of 24% (the mean of predation rates of all skate species studied to date) is applied to the range of fecundities reported for skates, the result is that 18–114 viable pups are produced annually per female skate. Even with a high mortality rate of 64% (the only direct estimate of natural mortality for any elasmobranch), each female skate produces 17–54 eggs annually. These values are higher than most elasmobranch fecundities. This maximization of fecundity is possible mainly because the fecundity of oviparous species is not limited by body size, as in viviparity. The protracted egg-laying season (4–12 months) of most skates (as in many other deep-sea fishes) maximizes the number of eggs laid.Communicated by P.W. Sammarco, Chauvin     

Influence of sponges on invertebrate recruitment: A field test of allelopathy

We manipulated live sponges in Belize, Central America, Big Pine Key, Florida (USA), and Indian River lagoon, Florida (USA) in summer/autumn, 1988. At each location, live sponges of three species were placed within 0.5 cm of ceramic tiles. Tiles with synthetic sponges positioned in the same manner and tiles with no sponges served as controls. Of 26 recruiting species analyzed, only one (Sponge sp. 6 — Indian River) was inhibited by living sponges. Four species (Perophora regina — Belize;Aiptasia pallida — Big Pine Key; andCrassostrea virginica andAscidia nigra — Indian River) recruited in greater numbers in the presence of sponges, suggesting that some larvae may be attracted rather than repelled by sponge allelochemicals. Allelopathic effects were less important than small-scale flow effects and patchy larval supply in determining recruitment patterns on surfaces adjacent to sponges.     

Mechanisms of habitat segregation between an invasive (Mytilus galloprovincialis) and an indigenous (Perna perna) mussel: adult growth and mortality

The invasive mussel Mytilus galloprovincialis and the indigenous mussel Perna perna coexist intertidally on the south coast of South Africa through partial vertical habitat segregation: M. galloprovincialis dominates the upper shore and P. perna the lower shore. Recruitment patterns can explain the zonation of P. perna, but not the invasive species. We examined the role of post-recruitment interactions by measuring spatial and temporal differences in adult growth and mortality rates of the two species. Specifically, we tested the hypothesis that interspecific differences in growth and mortality reflect adult distribution patterns. The two study locations, Plettenberg Bay and Tsitsikamma, are 70 km apart with two sites (separated by 300–400 m) per location, each divided into three vertical zones. Growth was measured seasonally using different marking methods in 2001 and 2003. Cumulative adult mortality was measured through summer in 2003/2004. Both species generally grew more slowly upshore, but they showed different effects of season. For P. perna, growth was significantly reduced in winter in the low zone, but unaffected by season in the high zone. For M. galloprovincialis, growth was either unaffected by season or increased in winter, even in the high zone. Thus, growth of P. perna and M. galloprovincialis was reduced under cool winter and warm summer temperatures, respectively; and while growth was more similar between species in summer, M. galloprovincialis grew much faster than P. perna in winter. Mortality of P. perna increased upshore. For M. galloprovincialis, mortality was not zone-dependent and was significantly greater than for P. perna on the low-shore and (generally) across the shore in Tsitsikamma. Both species had higher growth and mortality rates in Plettenberg Bay than in Tsitsikamma. Thus, P. perna seems able to maintain spatial dominance on the low-shore and at certain sites because of higher mortality of M. galloprovincialis. We conclude that seasonality in growth of the two species reflects their biogeographic affinities and that coexistence is possible through pre-recruitment effects that limit the vertical distribution of P. perna and post-recruitment effects that limit M. galloprovincialis.     

Shearwater diet during migration along the coast of Rio Grande do Sul,Brazil

The coastline of the state of Rio Grande do Sul (Brazil) is used annually as a migratory route by a number of seabird species. Thirteen surveys were carried out between July 1997 and July 1998 to obtain information on the diet of Puffinus gravis, Puffinus griseus, and Puffinus puffinus along the coast of Rio Grande do Sul. Sampling efforts focused on dead beach-cast specimens. One hundred twenty-one stomach content analyses were performed on P. gravis from a total of 238 specimens; 17 stomach content analyses were performed on P. griseus from a total of 54 specimens; and 25 stomach content analyses were performed on 25 P. puffinus from a total of 61 dead specimens. Cephalopods were the main prey found in the stomachs of the three species and were present in 96% of the P. gravis stomachs. The following cephalopods were observed: Ommastrephidae, Gonatus antarcticus, Gonatus sp., Lycoteuthis lorigera, Lycoteuthis sp., Histioteuthis sp., Cycloteuthis sp., Loligo plei, Loligo sanpaulensis, Sepiidae, and Argonauta nodosa. The latter was the most abundant species in the three species of Puffinus. The following fish were identified from otoliths: Porichthys porosissimus, Ctenosciaena gracilicirrhus, Paralonchurus brasiliensis, Cynoscion guatucupa, and Trichiurus lepturus. There was also a high incidence of synthetic materials such as plastic, polyethylene and rubber.     

Determining shark size from forensic analysis of bite damage

Bite damage patterns have long been used to estimate shark species and body size, with somewhat limited success. The lack of fit between damage patterns and shark size is partially due to variation in tooth size and shape within an individual. The ability to accurately predict body size from bite patterns is important for better understanding the ecological and behavioral underpinnings of shark bites/attacks on marine organisms, humans, and submarine equipment. To this end, we measured interdental distance (IDD) between the most labial teeth in the first six tooth files on both the upper and lower jaws, as well as the circumference of the portion of each jaw that bears teeth, for prepared jaw sets from fourteen shark species and regressed these data against total length. IDD is allometric as well as an accurate predictor of total length in all species examined, except Carcharhinus acronotus. Tooth-bearing circumference is also allometric and predictive of total length in all species. Though considerable overlap exists in IDD and circumference ranges among species for the total length ranges examined, Carcharodon carcharias and Isurus sp. can be differentiated from Carcharhinus limbatus, Carcharhinus brevipinna, and C. acronotus based on these values alone. When combined with knowledge of species-specific feeding behavior, geographic distribution, and habitat preferences, these simple measures from bite damage patterns allow quick, accurate assessment of shark size and potential species.     

The abundance and life histories of terrestrial isopods in a salt marsh of the Ria Formosa lagoon system,southern Portugal

Four species of isopod characteristic of salt marsh habitats, Tylos ponticus, Porcellio lamellatus, Halophiloscia couchii and Armadillidium album coexist in the upper reaches of the Ria Formosa lagoon salt marsh system in southern Portugal. In this locality, T. ponticus is the most abundant of the four species with mean annual densities of 2,950 m⁻² and a peak density of 10,387 m⁻² in July 1998 which is very much higher than what has previously been recorded for any isopod in any habitat. The mean annual densities for the other species were lower: P. lamellatus 36 m⁻², A. album 19 m⁻² and H. couchii 3 m⁻², indicating a less significant role in this ecosystem. Tylos ponticus and A. album started to breed on May, 24 and 12 months after release from the marsupium, respectively, where as other species start to breed in March, 12 months after their release from the marsupium. Tylos ponticus has a relative growth rate (RGR) of 0.23 between release from the marsupium and time of first breeding in July of its second year and breeds at a mature mass of 3.6 mg AFDM whereas the other three species mature after 10–12 months, have more than double this RGR but because of the shorter pre-reproductive period breed at masses of 1.8 mg AFDM for P. lamellatus, 1.0 mg AFDM for H. couchii, and 1.1 mg AFDM for A. album, respectively. The mass specific fecundity of all three of the less abundant species was higher than that of T. ponticus but the offspring of T. ponticus were ten times heavier than those of the next largest species, P. lammellatus. The difference in abundances between the species is interpreted as being due to the larger mass of the offspring of the most successful species. This larger mass confers an adaptive advantage due to larger size being associated with reduced juvenile mortality for isopods under abiotically stressful conditions. M. Sprung died in an accident on June 18, 2003     

Vertical distribution,abundance and community structure of oncaeid copepods in the Oyashio region,western subarctic Pacific

The vertical distribution, abundance and community structure of oncaeid copepods were investigated in the Oyashio region, western subarctic Pacific. Samples were collected with a 0.10 mm mesh closing type net from five discrete layers down to a maximum depth of 2,000 m in September and December 1996 and in April and October 1997. The copepods were widely distributed from epipelagic to bathypelagic zones, and showed prominent peaks of abundance above the thermocline and/or between 250 and 1,000 m depth. Standing stocks of total oncaeid copepods ranged from 1.5 to 2.5×10⁵ inds m^–2 at 0–2,000 m in the water column, which are the same order of magnitudes as those reported in tropical, subtropical and polar regions by previous workers. A total of 38 species and two forms belonging to the genera Oncaea, Triconia, Spinoncaea, Conaea and Epicalymma, and two provisionally classified species of the family Oncaeidae were identified in this study. Of these, 14 species have already been recorded from the eastern subarctic Pacific. Several warm-water species were also found in December 1996 and/or October 1997, when the effect of warm-core rings originating from the Kuroshio Current was evident in the epipelagic zone. Dominant species throughout the whole water column were T. borealis, T. canadensis, O. grossa, O. parila, O. rimula, O. lacinia, Epicalymma spp. and Oncaea sp. A, and these eight species together always made up about half of the total oncaeid numbers. Community analysis revealed that species composition in epipelagic and upper mesopelagic zones varied temporally due to the changes of hydrographic conditions, whereas those in lower mesopelagic and bathypelagic zones were relatively stable, reflecting the constancy of deep environments.Communicated by O. Kinne, Oldendorf/Luhe     

Some polyphagous Homoptera gain protection from their natural enemies by feeding on the toxic plantsSpartium junceum andErythrina corallodendrum (Leguminosae)

Summary Interactions were studied among alkaloid-containing legumes (Erythrina corallodendrum andSpartium junceum) and non-toxic plants (Citrus sinensis, Cucurbita moschata andEuphorbia tirucalli), several polyphagous homopterans,Aphis craccivora (Aphididae),Icerya purchasi, I. aegyptiaca (Margarodidae),Lepidosaphes ulmi (Diaspididae) andPlanococcus citri (Pseudococcidae), and some major natural enemies of these homopterans. Significant reductions in survival due to negative effects of alkaloid containing as compared with non-alkaloidal plants were recorded for the predatorsRodolia cardinalis andChilocorus bipustulatus, but not forCryptolaemus montrouzieri (Coleoptera: Coccinellidae),Chrysoperla carnea (Neuroptera: Chrysopidae) andSympherobius sanctus (Neuroptera: Sympherobiidae). The development time of the larvae or pupae ofR. cardinalis, C. carnea andS. sanctus was longer on the toxic plants than on the non-toxic ones. The percentage of parasitism ofA. craccivora collected from the non-alkaloidal plantsVicia palaestina andMelilotus albus was much higher than that onS. junceum. The parasitoid complexes ofA. craccivora differed between both plant groups. The nutritive value of honeydew ofI. purchasi andA. craccivora, as expressed by the life span ofEncyrtus infelix (Hymenoptera: Encyrtidae) adults, was also investigated. Life spans were significantly longer when the wasps fed on honeydew produced on non-alkaloidal plants (C. sinensis andPittosporum tobira) than on alkaloid containing plants whenI. purchasi — but notA. craccivora — was the producer. It is suggested that the chemical defense ofE. corallodendrum andS. junceum is exploited by polyphagous phytophages to reduce predation. In nature, population growth and density of four of the investigated homopterans are conspicuously high when they developed on the alkaloid containing plant species, and very low on non-alkaloid plants. The efficiency of their natural enemies may be reduced by sequestration of alkaloids (or other toxic plant compounds) or their transfer into excreted honeydew. Therefore it is assumed that a generalist phytophagous homopteran may be protected from its natural enemies, although at different rates of efficiency, if it can safely sequester the host allelochemical when it develops on toxic species within its host range.     

Megabenthic decapod crustacean assemblages on the Galician continental shelf and upper slope (north-west Spain)

The structure of megabenthic decapod crustacean assemblages on the Galician (north-west Spain) continental shelf (100 to 200 depth) and upper slope (200 to 500 m) was analyzed based on surveys carried out in autumn and spring, from 1980 to 1987. Forty species belonging to 19 families were caught. The portunid crab Polybius henslowii, a species with pelagic phases, was the most abundant species, but displayed strong spatial and temporal fluctuations. Other dominant species were the Norway lobster Nephrops norvegicus, the portunid Liocarcinus depurator, the galatheids Munida intermedia and M. sarsi, and the shrimps Solenocera membranacea, Plesionika heterocarpus, Pasiphaea sivado and Dichelopandalus bonnieri. Total abundance and biomass (average values excluding Polybius henslowii = 255 individuals and 2.06 kg/30 min tow) and species richness and diversity, H′ (6.85 species and H′ = 1.45 per tow) displayed a significant positive correlation with depth, and strong interannual fluctuations. The factors determining community organization were depth and, to a lesser extent, spatial structure. There was clear evidence of bathymetric zonation, differentiating between species characteristic of the slope (D. bonnieri and Pasiphaea sivado), shelf-slope edge (Macropipus tuberculatus, Pontophilus spinosus, Munida sarsi, S. membranacea, Processa spp.) and shelf (L. depurator, Macropodia tenuirostris, Paguridae and Chlorotocus crassicornis). The spatial zonation was related to changes in oceanography and sediment along the continental margin. Goneplax rhomboides, N. norvegicus, C. crassicornis and Alpheus glaber are benthic species which generally exhibit burrowing behaviour, and they were found mainly in the southern area where there are fine sediments due to the outwelling from the Rías Baixas. Different benthopelagic shrimps (Pontophilus spinosus, Plesionika heterocarpus,Processa spp. and Pasiphaea sivado) were typical of the zone just north of Fisterra, characterized by a convergence of water masses bringing about an increase in productivity due to upwelling. The benthic anomuran and brachyuran crabs Munida intermedia, M.␣sarsi, L.␣depurator and Macropipus tuberculatus were characteristic of the northwestern zone between Fisterra and Estaca, where the infauna reaches high biomass despite coarser sediments with a lower concentration of organic material than in the southern area. Lastly, both the Paguridae and Macropodia tenuirostris were species typically found in the waters in the northern shelf. Based on interannual changes in assemblage structure, two periods could be distinguished: between 1980 and 1984, when Polybius henslowii, D. bonnieri and Pasiphaea sivado had abundance peaks; and another period from 1985 to 1987 when L. depurator, Munida intermedia, M.␣sarsi and Macropipus tuberculatus increased in abundance. Received: 21 May 1996 / Accepted: 9 August 1996     

Wave action and competitive interaction between the invasive mussel Mytilus galloprovincialis and the indigenous Perna perna in South Africa

On the south coast of South Africa, the invasive alien mussel Mytilus galloprovincialis shows partial habitat segregation with the indigenous mussel Perna perna. P. perna predominates in the lower mussel zone and M. galloprovincialis in the upper zone, with mixed beds where the two overlap. We examined competitive interactions between these species by translocating mussels into small plots at high densities. Treatments involved different combinations of species and densities placed in each zone. Mortality was monitored regularly and at the end of each experiment, growth and condition index were measured. The experiment was attempted three times. The first two attempts were disrupted by wave action, especially winter storms, but provided information on species-specific effects of wave action. In experiment one, wave induced mortality decreased from a mean for both species of approximately 90% on the low shore to ca. 50% on the high shore, and was 15–30% lower for P. perna than M. galloprovincialis in each zone. In experiment two, M. galloprovincialis mortality was not affected by zone (Kruskal–Wallis test, P > 0.05), but was higher than P. perna mortality in the low zone (P < 0.05). P. perna survival was significantly (P < 0.05) lower on the high than mid and low zones, apparently due to the effects of greater emersion. Condition index showed a similar pattern, being lowest in the low zone for M. galloprovincialis and in the high zone for P. perna (3-way ANOVA, P < 0.05). Growth rates were fastest for both species in the low zone (Kruskal–Wallis, P < 0.05 in both cases). The third experiment was run for 12 months in the low zone only and provided evidence of intraspecific competition for P. perna and of interspecific competition. Condition was significantly greater for P. perna in all treatments (2-way ANOVA), as was growth (Kruskal–Wallis P < 0.05). Significant treatment effects indicated that P. perna had a negative effect on M. galloprovincialis survival (Kruskal–Wallis, P < 0.05). Again wave action was important; by the end of the experiment all mussels had been removed from plots stocked only with M. galloprovincialis. Thus P. perna improves survival of M. galloprovincialis on the low shore in the short term, by providing protection against wave action, but excludes it competitively in the longer term. The results show that partial habitat segregation is likely to be a permanent feature on the south coast, with M. galloprovincialis unable to dominate the low shore due to the effects of waves and competitive exclusion by P. perna. This study is the first attempt to examine the mechanisms of interaction between invasive and indigenous marine mussel species and provides evidence of the importance of environmental conditions in the mediation of this interaction.     

Reproductive strategies of Mycalecontarenii (Porifera: Demospongiae)

The sexual and asexual phases of the reproductive cycle of the sponge Mycalecontarenii (Martens, 1824) were studied from samples collected over a period of 2 years (June 1994 to May 1996) in a Mediterranean coastal basin (Porto Cesareo, southwestern Apulia). The species is viviparous; year round it shows a discontinuous oocyte production. No males were found, possibly owing to the very short period of spermatogenesis. M.␣contarenii produces asexual buds during autumn and winter, and this process seems to be related to changes in water temperature. Even though, at present, we cannot evaluate the incidence of larval recruitment, we postulate that asexual reproduction in M.contarenii plays a crucial role in maintaining the sponge population. Received: 8 July 1997 / Accepted: 5 November 1997