The effect of depth and sediment type on the spatial distribution of shallow soft-bottom amphipods along the southern Portuguese coast

The present study provides updated information on amphipod assemblages of the southern Portuguese coast, the most south-western part of continental Europe. The influence of depth and sediment type on the spatial distribution of shallow soft-bottom amphipods was tested by means of univariate and multivariate methods. Samples were collected down to 40?m depth within areas ranging from very fine sand to gravel. The ratio between density and diversity was found to increase with depth and from coarser to finer sediments. Both physical variables tested were found to play a major role in structuring amphipod assemblages. However, some dominance of depth was evident as samples were mainly separated according to this variable. Moreover, while only 15 taxa were common to all depth levels, 35 were collected from all sediment types. Both depth and sediment type may override the potential effects of geographical location, as samples from different areas along the southern Portuguese coast were biologically related. A peculiar habitat was sampled in this study: a maerl bed, which was confirmed to be a hotspot of density. Nevertheless, this habitat did not harbor a distinct amphipod assemblage but some taxa such as Caprella spp., Lembos websteri, lysianassids and Gammarella fucicola were preferentially collected there. The biogeographical analysis pointed out the occurrence of 8% of Mediterranean endemic species, confirming the influence of the Mediterranean Sea beyond the Strait of Gibraltar, and the Atlantic-Mediterranean feature of the amphipod fauna of this geographically relevant area. The present results support the need to interpret the spatial distribution patterns of marine species in relation to the combined effect of depth and sediment characteristics.     

Comparison of the epifauna spatial distribution in Posidonia oceanica,Cymodocea nodosa and unvegetated bottoms: Importance of meadow edges

In order to detect the influence of seagrass, the most important habitats of shallow soft-bottom along the Mediterranean coast, on spatial distribution of epifauna, four different types of habitat were sampled: Posidonia oceanica, Cymodocea nodosa, edge of Posidonia meadow and sandy bottoms. Sampling was carried out, using the hand net method, in various random sites and at different times (April 95, August 95 and February 96) on the Alicante coast (SE Spain). A double taxonomic approach was used in order to detect spatial and temporal changes in the abundance of the main taxonomic groups and amphipod species. The differences among habitats were detected by non-parametric multidimensional scaling (MDS) for community structure and by analysis of variance for groups and amphipod populations. The community structure defined by the abundance of taxonomic groups was not significantly modified by the habitats. Single abundance of taxonomic groups also had no significant preference for the type of habitat except in the case of Acari, a group linked to Posidonia. However, certain trends of preference, such as mysids on Posidonia edge and isopods on Cymodocea, were detectable. On other hand, the amphipod assemblage showed important differences among habitats. Some species changed significantly in abundance depending on the habitat considered, e.g. Dexamine spiniventris and Perioculodes longimanus. It was possible to conclude that there is an important influence of Posidonia and Cymodocea on epifauna distribution by diversification of habitat structure on sandy bottoms, but it was more important at the species level than when considering taxonomic groups. Furthermore, sandy bottoms and the meadow edges had a relatively high importance on fauna distribution, depending on taxa and the period of the year.     

SELF‐THINNING AND SIZE INEQUALITY DYNAMICS IN A CLONAL SEAWEED (SARGASSUM LAPAZEANUM,PHAEOPHYCEAE)1

Fronds of clonal seaweeds with extensive holdfasts relative to frond size are known not to self‐thin during growth, even in crowded stands. We tested whether frond self‐thinning would occur for such a seaweed since these traits are more similar to those of unitary seaweeds, which do self‐thin in crowded conditions. We used Sargassum lapazeanum Setch. et N. L. Gardner (Fucales, Phaeophyceae) from the Pacific coast of Mexico, for which we first confirmed its clonal nature by performing a regeneration experiment in culture tanks. During the growth season (winter to late spring), S. lapazeanum stand biomass increased, while frond density and size inequality (Gini coefficient for frond biomass) decreased. These results indicate that self‐thinning occurred at the frond level. We propose a conceptual model for frond dynamics for clonal seaweeds in general. In stands of clonal species with small fronds and relatively extensive holdfasts (particularly when holdfasts are perennial), frond dynamics would be determined mostly by intraclonal regulation, which seems to prevent excessive crowding from occurring. Such species display a positive biomass–density relationship during the growth season. On the contrary, in stands of clonal species with large fronds relative to holdfast size, frond dynamics would be determined mostly by interactions among genets. For such species, self‐thinning may be detected at the frond level in crowded stands, resulting in a negative biomass–density relationship during growth.     

Characterization of the Adhesive Holdfast of Marine and Freshwater Caulobacters

Caulobacters are prosthecate (stalked) bacteria that elaborate an attachment organelle called a holdfast at the tip of the cellular stalk. We examined the binding of lectins to the holdfasts of 16 marine Caulobacter strains and 10 freshwater species or strains by using a panel of fluorescein-conjugated lectins and fluorescence microscopy. The holdfasts of all the marine isolates bound to only wheat germ agglutinin (WGA) and other lectins that bind N-acetylglucosamine (GlcNac) residues. The freshwater caulobacters showed more variability in holdfast composition. Some bound only to WGA and comparable lectins as the marine strains did. Others bound additional or other lectins, and some did not bind to the lectins tested. The binding of WGA appeared to involve the regions of the holdfast involved with adhesion; a holdfast bound to WGA was significantly less adhesive to glass. Competition experiments with WGA-binding holdfasts and oligomers of GlcNac demonstrated that trimers of GlcNac (the preferred substrate for WGA binding) were more effective than dimers or monomers in preventing WGA binding to holdfasts, suggesting that stretches of contiguous GlcNac residues occur in the WGA-binding holdfasts. In addition, differences between freshwater and marine holdfasts in the strength of WGA binding were noted. The effect of a number of proteolytic and glycolytic enzymes on holdfast integrity was examined; the proteases had no effect for all caulobacters. None of the glycolytic enzymes had an effect on marine caulobacter holdfasts, but chitinase and lysozyme (both attack oligomers of GlcNac) disrupted the holdfasts of those freshwater caulobacters that bound WGA. Despite some similarity to chitin, holdfasts did not bind Calcofluor and no measurable effects on holdfast production were detectable after cell growth in the presence of diflubenzuron or polyoxin D, inhibitors of chitin synthesis in other systems. Finally, the holdfasts of all caulobacters bound to colloidal gold particles, without regard to the coating used to stabilize the gold particles. This binding was stronger or more specific than WGA binding; treatment with colloidal gold particles prevented WGA binding, but the reverse was not the case.     

The North-American amphipods, Melita nitida Smith, 1873 and Incisocalliope aestuarius (Watling and Maurer, 1973) (Crustacea: Amphipoda: Gammaridea), introduced to the Western Scheldt estuary (The Netherlands)

The American amphipod species Melita nitida andIncisocalliope aestuarius have been found in the WesternScheldt estuary (the Netherlands). This is the first record of these species inthe north-east Atlantic. Shipping is the most likely vector of introduction.Thedistribution of both species is investigated and compared with the distributionand the microhabitat of co-occurring amphipod species. Melitanitida is known from both the east and west coast of North Americaand I. aestuarius originates from the east coast of NorthAmerica. Until now neither has been reported from other parts of the world. Inthe Netherlands both species are restricted to the mesohaline part of theWestern Scheldt. Melita nitida occurs predominantly underPacific oysters at the underside of boulders, mainly sublittorally.Incisocalliope aestuarius is associated to hydrozoans.Bothmicrohabitats are hardly utilized by other amphipod species. Therefore, thetheory that the existence of many empty niches in north-western Europeanbrackish waters make this environment particularly susceptible to invasions ofalien species is corroborated. The application of hard substrates in a regionoriginally predominated by soft bottoms moreover facilitates the introductionofexotic species. The species community on hard substrates in the mesohaline partof the Western Scheldt contains a high proportion of introduced species:approximately one third of the macrofauna species is of allochthonous origin.     

Patterns of gammaridean amphipod abundance and species composition associated with dominant subtidal macroalgae from the western Antarctic Peninsula

The communities of gammaridean amphipods associated with eight dominant macroalgal species were examined near Palmer Station, Western Antarctic Peninsula. A total of 78,415 individuals belonging to 32 amphipod taxa were identified with mean densities ranging up to 20 individuals/g algal wet wt. The most abundant amphipod taxon, Metaleptamphopus pectinatus, was found to associate predominately with the brown alga Desmarestia menziesii, while the second most common taxon, Jassa spp. occurred primarily on the red alga Gigartina skottsbergii. Non-metric multidimensional scaling analysis demonstrated that the population densities of each amphipod species and amphipod species composition were similar on the same algal species but dissimilar on different species of algae. Comparisons of amphipod communities associated with a given algal species but from different sampling sites indicated that although the structure of species-specific macroalgal-associated amphipod communities can vary across spatial scales of 3 km, 50% of the macroalgal species examined showed no significant inter-site differences in associated amphipod community structure. Spearman rank correlation analyses showed that higher abundances of amphipods occurred on the macroalgae with the highest number of branches. As many Antarctic amphipods are known consumers of macroalgae, their remarkable abundances are likely to play a significant role in mediating energy and nutrient transfer in nearshore Antarctic Peninsular macroalgal communities.     

Grazing of the invasive alga Codium fragile ssp. tomentosoides by the common periwinkle Littorina littorea: Effects of thallus size, age and condition

We examined the potential of herbivory by the common periwinkle Littorina littorea to limit recruitment and vegetative re-growth of the invasive green alga Codium fragile ssp. tomentosoides in a series of manipulative field experiments in tidepools on a wave-exposed rocky shore in Nova Scotia, Canada. Snails were excluded or included from circular plots (14 to 20 cm diameter) with cages to compare growth and survival of C. fragile against procedural (partial cages) or natural (uncaged) control plots. Our results show that L. littorea may restrict growth and survival of C. fragile by grazing new recruits (< 2 cm thallus length), fronds of adult thalli that are bleached and necrotic, and residual holdfasts (< 2 mm thickness) of detached thalli (artificially severed to mimic wave dislodgement). Once recruits grow beyond a critical size (~ 3 cm), or damaged tissues or holdfasts regenerate, grazing of C. fragile by L. littorea appears to be limited and ineffectual. Our experimental results corroborate correlative evidence from previous studies that herbivory by L. littorea limits the abundance of C. fragile in tidepools on the Atlantic coast of Nova Scotia, particularly pools in the high intertidal zone where these snails are abundant. Lower on the shore, littorinid grazing and physical stressors that render algae more vulnerable to grazers (e.g. UV radiation and freezing) are less intensive, and probably have less of a regulatory effect on populations of C. fragile.     

Marine brown algae (Phaeophyta) from the north coast of Papua New Guinea,with a description of Dictyota magneana sp. nov.

The marine benthic brown algae of the north coast of Papua New Guinea (mainly from Madang province) are documented, based on collections made by the first author, between 1980 and 1990. All records [34 identified taxa (+Sargassum spp.)] are listed with bibliographic, taxonomic, nomenclatural and biogeographical notes. The specimens belonging to the genus Dictyota are identified according to recent species definitions. D. magneana De Clerck et Coppejans is described as new to science. Only some representatives of the genus Sargassum have been identified to species level. The phaeophycean flora of the north coast is very similar to that of the south coast (Port Moresby area); nevertheless some differences can be observed.     

The ecology of Mytilus edulis L. in exposed rocky intertidal communities

Information is given to show that Mytilus edulis L. is an important component of exposed rocky coast communities of the Western Hemisphere. It is abundant on the exposed coast of Washington State with a potential distribution extending from + 10.6 ft ( + 3.2 m) down to at least 10 m depth: it is not continuous along this entire gradient. It dominates a band from +10.6 ft down to +9.6 ft ( + 3.2 to + 2.9 m) tidal height where it usually intergrades with its congener M. californianus Conrad. Its upper limit appears to be determined by desiccation stress and the lower probably by competition and or predation.In the mid-intertidal zone, from + 5.6 to + 9.6 ft ( + 1.7 to 2.9 m), where M. californianus is dominant, M. edulis occurs in patches of cleared space which usually form in the M. californianus bed matrix in winter months. Here, it acts as an opportunistic species by colonizing quickly and growing rapidly to reproductive size. It is preyed upon by Thais spp. which prefer it to Mytilus californianus and eventually is eliminated from these patches. It also occurs, sometimes abundantly, in tufts of the red alga Endocladia muricata (Postels & Ruprect) J. Agardh at high and mid-intertidal heights.In low intertidal and subtidal regions, to ≈ 10 m depth, Mytulis edulis can be found in refuges of filamentous or intricately arranged substrata which generally offer protection from predation. These are represented by a wide variety of forms such as hydroids. bryozoans, filamentous algae, coralline algae, and kelp with convoluted holdfasts or complex stipes.     

Different ammonia tolerances may facilitate spatial coexistence of Gammarus roeselii and the strong invader Dikerogammarus villosus

The introduction of invasive species that can replace native species is one of the most critical threats to the biodiversity of aquatic systems. Here we investigated the potential contribution of one factor to the coexistence of the indigenous amphipod Gammarus roeselii and the invasive amphipod Dikerogammarus villosus in the same ecosystem (Lake Constance) within different microhabitats. We quantitatively studied the influence of ambient ammonia concentrations on the distributions of the two amphipod species. We also assessed the ammonia tolerance ranges of both species in laboratory experiments by measuring mortality rate, precopula disruption, egg mortality, and microhabitat choice. The proportion of G. roeselii among the two amphipod species was significantly positively related to the ammonia concentration in the water, which indicated that the distribution of the invasive D. villosus was limited at high ammonia concentrations. Although the mortality rates of the two species did not significantly differ, G. roeselii was more tolerant to ammonia with regard to precopula disruption, egg mortality, and microhabitat choice. The effective ammonia concentrations that led to a significantly reduced direct reproductive success in D. villosus were within the range of the highest field concentrations measured, where only G. roeselii occurred. D. villosus may have a smaller range than the indigenous G. roeselii partially because of its lower tolerance to higher ammonia concentrations, which lead to reduced reproductive success. Beside other habitat parameters differences on ammonia tolerance between the two amphipods might allow their coexistence along a gradient of microhabitats in Lake Constance.     

Comparative study of the antennal sensilla of five species of root maggots: Delia radicum L., D. floralis F., D. antiqua Mg., D. platura MG. (Diptera : Anthomyiidae) and Psila rosae F. (Diptera : Psilidae)

A comparative study of the antennal sensilla of Delia radicum L., D. floralis F., D. antiqua Mg., D. platura Mg. (Diptera : Anthomyiidae) and Psila rosae F. Diptera Psilidae) is undertaken. For both sexes of each species, the type, distribution, and density of sensilla are determined. All 5 species have trichoid (olfactory) and grooved (olfactory) sensilla. Basiconica I (blunt) sensilla (olfactory) are found on each of the species examined, except D. platura. Basiconica II (tapered) (olfactory) and clavate (olfactory) sensilla are found only on Delia species. Also, only Delia species have single-chambered, dorsal pits, and these contain basiconic II pit sensilla (olfactory). Common to all 5 species is a multi-chambered ventral pit (olfactory). In the ventral pit, all 5 species have grooved pit sensilla (olfactory). In addition to this type of sensillum the Delia species have smooth-walled conical pit sensilla (hygro-/thermosensitive) and P. rosae has granular pit sensilla (hygro-/thermosensitive). Smooth-walled tapered pit sensilla (hygro-/thermosensitive) are found in D. radicum. Similarities and differences in the density of surface sensilla between dorsal and ventral funicular surfaces, male and female flies, and oligophagous (D. antiqua, D. radicum, D. floralis and P. rosae) and polyphagous (D. platura) species are compared. Several differences in sensillum density between the dorsal and ventral funicular surfaces are observed, but these do not fit into a consistent trend. Except for D. radicum, there are differences in sensillum density between male and female flies. For the oligophagous species, females have a greater sensillum density, whilst for the polyphagous D. platura males have a greater sensillum density. Comparisons between species show the greatest differences between the Delia species and P. rosae, and within the 4 Delia species, differences in sensillum density do not correlate with host range or body size.     

Patterns of abundance and assemblage structure of epifauna inhabiting two morphologically different kelp holdfasts

The mobile fauna associated with two sympatric kelp species with different holdfast morphology (Saccorhiza polyschides and Laminaria hyperborea) was compared to test for differences in the assemblage structure of holdfast-associated mobile epifauna. A total of 24,140 epifaunal individuals were counted from 30 holdfasts of each kelp species. Overall epifaunal abundances exceeded faunal abundances previously reported from holdfasts of other kelps. Three taxonomic groups, Amphipoda, Mollusca, and Polychaeta, accounted for ca. 85% of all individuals. Total abundances increased with the amount of habitat available, quantified either as the volume or the area provided by the holdfasts. The multivariate structure of the epifaunal assemblage did not differ between holdfasts of the two kelp species. However, epifaunal assemblages responded differentially to the habitat attributes provided by each type of kelp holdfast: multivariate variation in the assemblage structure of epifauna was mostly explained by holdfast area and volume for L. hyperborea, and by the surface-to-volume ratio for S. polyschides holdfasts. Therefore, the physical attributes of biogenic habitats, here kelp holdfasts that better predict patterns in the assemblage structure of associated fauna can differ according to their different physical morphology, even though the overall assemblage structure of associated fauna was similar.     

Short-term predator avoidance behavior by invasive and native amphipods in the Great Lakes

Understanding predator avoidance behavior by prey remains an important topic in community and invasion ecology. Recently, the Ponto-Caspian amphipod Echinogammarus ischnus (Stebbing 1898) was accidentally introduced into the Great Lakes. Since its introduction, it has displaced the native amphipod, Gammarus fasciatus (Say 1818), from several locations in the lower lakes. To assess whether behavioral differences in predator avoidance might be a causal mechanism increasing the success of the invasive amphipods, two experiments were conducted examining (1) native and invasive amphipod behavioral responses to five fish species with different foraging behaviors, and (2) amphipod responses to different densities of round gobies, a hyper-abundant benthic invertivore. Echinogammarus reduced its distance moved in the presence of all fish species tested, whereas Gammarus reduced its distance moved only after exposure to round gobies, black crappies, and rainbow darters. Both amphipod species increased the time spent motionless following exposure to round gobies, but not after encountering the scent of most of the remaining fish predators. The exception was that Echinogammarus also responded to black crappie scent whereas Gammarus did not. Although both amphipod species exhibited behavioral responses to many of the fish predators, the magnitude of their responses differed only after exposure to the brown bullhead. In the bullhead trials, Echinogammarus reduced its distance traveled significantly more than Gammarus. Both amphipod species increased their avoidance response to increasing goby density, however, the pattern of avoidance behavior was different. Invasive E. ischnus exhibited a consistently strong avoidance response to round gobies over the test duration. Native G. fasciatus initially avoided goby scent, but then either ceased their avoidance response or showed a hyper-avoidance response, depending on goby density. These results suggested (1) both species of amphipods were able to differentiate and react to a variety of fish predators, (2) invasive Echinogammarus amphipods avoided a larger range of fish predators than the native Gammarus, (3) increased avoidance behavior was associated with an increased density of fish, and (4) the avoidance response patterns of invasive Echinogammarus when faced with round goby predators might lead to increased predation on native Gammarus in habitats where they co-occur.     

Tube building by the amphipods Lembos websteri Bate and Corophium bonnellii Milne Edwards

Lembos websteri Bate and Corophium bonnellii Milne Edwards are tubicolous. The morphology of tubes collected from Laminaria holdfasts is described and contrasted with that of tubes built amongst filamentous algae in a wave-sheltered environment. In both species tube length is a function of animal length. Corophium build tubes with openings raised above the substratum more commonly than Lembos, and will construct mats of tubes away from solid surfaces; Lembos tend to build against solid surfaces. Tube construction appears to depend upon the ‘knitting’ activity of the first two pairs of peraeopods, utilizing a secretion containing acid and sulphated mucopolysaccharides. The process of tube construction on substrata of known granulometry and the morphology of tubes built in the laboratory are described. The burrowing capabilities of both species are compared. On sand, Lembos tend to build on the sediment surface, whilst Corophium tend to burrow into the substratum. Corophium begin tube building after transference to a new situation much sooner than Lembos, with female Lembos starting to build sooner than male Lembos, Corophium can build a covering tube in < 15 min. In the absence of extraneous material, both species line glass tubes with a meshwork of mucoid filaments. In empty dishes, mucoid tubes are rarely built. The ability of Lembos to colonize wave-exposed sites may be related to its preference for more sheltered spaces within kelp holdfasts.     

Entomophthora brevinucleata sp. nov. [Zygomycetes,Entomophthoraceae], a pathogen of gall midges [Dip.: Cecidomyiidae]

Entomophthora brevinucleata, a fungal pathogen of gall midges, is described. The species occurs on at least 3 host species, including the wheat blossom midgeSitodiplosis mosellana Géhin andMycodiplosis sp.E. brevinucleata produces rhizoids, either monohyphal or comprising bundles of a few hyphae, without holdfasts. The hyphal bodies are subspherical with mean dimensions of 27.5×22.1 μm. The primary conidia are campanulate with mean dimensions from individual hosts of 11.2–18.9×8.7–15.8 μm. They contain 3–13 nuclei with a mean diameter of between 2.7 and 3.7 μm stained with lactophenol-aceto-orcein. The secondary conidia are of similar shape but smaller. Cystidia and resting spores were not observed.