Sublethal effects of a rapidly spreading native alga on a key herbivore

1. Multiple anthropogenic stressors are causing a global decline in foundation species, including macrophytes, often resulting in the expansion of functionally different, more stressor‐tolerant macrophytes. Previously subdominant species may experience further positive demographic feedback if they are exposed to weaker plant–herbivore interactions, possibly via decreased palatability or being structurally different from the species they are replacing. However, the consequences of the spread of opportunistic macrophytes for the local distribution and life history of herbivores are unknown.
2. The green alga, Caulerpa filiformis, previously a subdominant macrophyte on low intertidal‐shallow subtidal rock shores, is becoming locally more abundant and has spread into warmer waters across the coast of New South Wales, Australia.
3. In this study, we measured (a) the distribution and abundance of a key consumer, the sea urchin Heliocidaris erythrogramma, across a seascape at sites where C. filiformis has become dominant, (b) performed behavioral field experiments to test the role of habitat selection in determining the local distribution of H. erythrogramma, and (c) consumer experiments to test differential palatability between previously dominant higher quality species like Ecklonia radiata and Sargassum sp. and C. filiformis and the physiological consequences of consuming it.
4. At all sites, urchin densities were positively correlated with distance away from C. filiformis beds, and they actively moved away from beds. Feeding experiments showed that, while urchins consumed C. filiformis, sometimes in equal amounts to higher quality algae, there were strong sublethal consequences associated with C. filiformis consumption, mainly on reproductive potential (gonad size). Specifically, the gonad size of urchins that fed on C. filiformis was equivalent to that in starved urchins. There was also a tendency for urchin mortality to be greater when fed C. filiformis.
5. Overall, strong negative effects on herbivore life‐history traits and potentially their survivorship may establish further positive feedback on C. filiformis abundance that contributes to its spread and may mediate shifts from top‐down to bottom‐up control at locations where C. filiformis has become dominant.

The spread of the previously subdominant Caulerpa filiformis along the coast of NSW, Australia has strong lethal and sublethal effects on a key herbivore population. We performed sea urchin surveys, behavioral experiments, and consumer experiments and found that C aulerpa has strong negative effects on their populations. These results may establish reinforcing feedbacks on the spreading algae and mediate shifts from top‐down to bottom‐up control.     

Host location and selection by the symbiotic sargassum crab <Emphasis Type="Italic">Portunus sayi</Emphasis>: the role of chemical,visual and tactile cues

The Sargassum community consists of a unique and diverse assemblage of symbiotic fauna critical to pelagic food chains. Associated symbionts presumably have adaptations to assist in finding Sargassum. In situ scattered Sargassum patches accumulate as they are pushed toward the shoreline (via wind, waves, currents or tides) and are frequently less than 1 m apart and in depths of 10 cm or less as the patches approach the shoreline Crabs, and other symbiotic fauna, must relocate to another patch that is seaward in direction or likely perish as their current patch will likely become beached. This study investigated sensory cues used for host location and selection by the Sargassum crab, Portunus sayi. Chemical detection trials were conducted with a two-chamber choice apparatus with Sargassum spp. and Thalassia testudinum as habitat source odors. Visual detection trials (devoid of chemical cues) and habitat selection trials were conducted in which crabs were given a choice between hosts. Results showed that P. sayi responded to chemicals from Sargassum spp. Crabs visually located host habitats but did not visually distinguish between different hosts. In host selection trials, crabs selected Sargassum spp. over artificial Sargassum and T. testudinum. These results suggest that crabs isolated from Sargassum likely use chemoreception; within visual proximity of a potential patch, crabs likely use both chemical and visual information.     

Relationships between the spread of Caulerpa filiformis and fish communities on temperate rocky reefs

The previously sub‐dominant native marine macrophyte Caulerpa filiformis is now dominant on many sub‐tidal rocky reefs in New South Wales (NSW), Australia and is expanding its distribution. As C. filiformis is highly chemically defended and structurally different to co‐occurring habitat‐forming macrophytes, two key attributes that govern fish assemblages, we hypothesized that fish assemblages, particularly herbivorous fishes, would be different at sites where C. filiformis occurred from where it was previously absent and within sites, fish community structure would be correlated to the cover of C. filiformis. We investigated these hypotheses by determining reef‐associated fish assemblage attributes (assemblage structure, species richness, total abundance, Shannon‐Weiner diversity, abundance of herbivorous species) along transects within sites where C. filiformis was present and absent. Surprisingly, despite large patches and very high densities of C. filiformis on the reefs we sampled, at larger spatial scales (i.e., among sites) no fish assemblage metrics differed between sites with large stands of C. filiformis and sites without the alga. Moreover the abundance of one dominant herbivore, the rock cale Aplodactylus lophodon, was greater at sites within large beds of C. filiformis. At smaller spatial scales, however, i.e. within sites where C. filiformis was present, fish assemblages did vary as a function of C. filiformis cover along transects, although this was not consistent across sampling times. Overall, our results suggest that the potential effects of the spread of this alga on faunal communities warrants further investigation.     

Habitat formed by the invasive macroalga Caulerpa filiformis (Suhr) Hering (Caulerpales,Chlorophyta) alters benthic macroinvertebrate assemblages in Peru

The green macroalga Caulerpa filiformis has been spreading on shallow soft sediment habitats along the Peruvian coast, colonizing previously unvegetated sediments to create monospecific meadows. We examined the nature of the impact of C. filiformis meadows on the density, taxonomic richness and assemblage structure of epifaunal and infaunal benthic macroinvertebrates. Specifically, we tested whether the spread of C. filiformis has resulted in different macroinvertebrate assemblages than those formed by the dominant native macroalgae (i.e., Rhodymenia spp.) and unvegetated sediments. Surveys were undertaken in two bays in each of two locations, in central and southern Peru, during winter 2017 and summer 2018. In general, our results show that macroinvertebrate assemblages were similar across all three habitats, although there were some differences, related to location and time, but with no clear patterns observed. Taxonomic richness and density was generally higher in the vegetated habitats than the unvegetated habitat, and where there were differences between the two vegetated habitats there was no consistent pattern of which habitat supported the highest richness or density. Given invading C. filiformis is primarily colonizing unvegetated habitats it would appear that this species is creating a new niche which supports similar assemblages, but higher taxonomic richness and density than unvegetated habitats. While our study suggests that C. filiformis is having a limited ecological impact we recommend that actions be put in place to limit the spread of this invasive species at the same time as increasing monitoring of the ecological impacts of this species as lags in the ecological impacts of invasive species are common.

     

Structure and composition of grassland habitats used by wintering Smith's Longspurs: the importance of native grasses

ABSTRACT Smith's Longspurs (Calcarius pictus) are a species of concern in North America because of their limited range and apparent low population size. To better understand winter habitat needs and guide management of this species, we examined habitat associations of Smith's Longspurs in eastern Arkansas by comparing grassland patches where Smith's Longspurs flushed to randomly located patches in the same area. Smith's Longspurs were found in sparse grassland patches of relatively low height adjacent to airport runways where the native grass prairie three‐awn (Aristida oligantha) dominated ground cover and vertical structure. Smith's Longspurs were not found in vegetation plots dominated by non‐native Bermuda grass (Cynodon dactylon). Prairie three‐awn grass may provide concealment from predators and their seeds may be an important food source. Occurrence of Smith's Longspurs was also correlated with less litter, perhaps because deeper litter could make walking and searching for seeds more difficult. Availability of suitable habitat for Smith's Longspurs along airport runways may be declining due to natural succession of grassland habitat in the absence of disturbance and recent changes in management that favor Bermuda grass. Conversion and degradation of native prairie has resulted in the decline in abundance and distribution of Chestnut‐collared Longspurs (C. ornatus) and McCown's Longspurs (Rhynchophanes mccownii). Our findings suggest that conversion of native grasslands to non‐native grasslands results in loss and degradation of habitat for wintering populations of Smith's Longspurs.     

Long-Term Field Study Reveals Subtle Effects of the Invasive Alga Sargassum muticum upon the Epibiota of Zostera marina

Invasive species can alter coastal ecosystems both directly, e.g. through competition for substratum and nutrients, and indirectly. Indirect effects may be mediated by creation of dissimilar or inimical habitats, changes in predator and/or prey assemblages, alterations in associated biota, and perturbations of water movement and thermal regimes. Previous studies have shown that invasive algae can modify native habitat architecture, disrupt intricately linked food webs and alter epibiotic assemblages. In the UK, the seagrass Zostera marina supports a diverse epibiotic assemblage, influencing key factors such as sediment dynamics, depositional regime and trophic linkages. Increasing encroachment of the invasive alga Sargassum muticum into seagrass meadows changes the physical and chemical characteristics of the local environment and creates the potential for changes in the epibionts associated with the seagrass blades, threatening the integrity of the seagrass ecosystem. We investigated the effects of S. muticum invasion upon the epibiota of Z. marina in a drowned river valley in SW England seasonally from spring to autumn over four years in an in-situ manipulative experiment, comparing permanent quadrats with and without artificially introduced S. muticum. Epibiota were weighed, identified to the most detailed operational taxonomic unit (OTU) possible, and unitary organisms were enumerated. Multivariate PERMANOVA+ analysis revealed significant differences in epibiont assemblages between Sargassum treatments. Linear mixed effects models indicated that differences in epibiota assemblage composition were not reflected as significant differences in mean biomass per sample, or number of epibiont OTUs per sample. We conclude that S. muticum invasion into Z. marina meadows may significantly alter the species composition and abundance distribution of epibiotic assemblages found on the blades of the seagrass. Thus S. muticum invasion could have more wide-reaching effects on processes within coastal ecosystems than predicted purely by direct effects.     

Mediterranean Species of Caulerpa Are Polyploid with Smaller Genomes in the Invasive Ones

Caulerpa species are marine green algae, which often act as invasive species with rapid clonal proliferation when growing outside their native biogeographical borders. Despite many publications on the genetics and ecology of Caulerpa species, their life history and ploidy levels are still to be resolved and are the subject of large controversy. While some authors claimed that the thallus found in nature has a haplodiplobiontic life cycle with heteromorphic alternation of generations, other authors claimed a diploid or haploid life cycle with only one generation involved. DAPI-staining with image analysis and microspectrophotometry were used to estimate relative nuclear DNA contents in three species of Caulerpa from the Mediterranean, at individual, population and species levels. Results show that ploidy levels and genome size vary in these three Caulerpa species, with a reduction in genome size for the invasive ones. Caulerpa species in the Mediterranean are polyploids in different life history phases; all sampled C. taxifolia and C. racemosa var. cylindracea were in haplophasic phase, but in C. prolifera, the native species, individuals were found in both diplophasic and haplophasic phases. Different levels of endopolyploidy were found in both C. prolifera and C. racemosa var. cylindracea. Life history is elucidated for the Mediterranean C. prolifera and it is hypothesized that haplophasic dominance in C. racemosa var. cylindracea and C. taxifolia is a beneficial trait for their invasive strategies.     

Evidence that creation of a Pinus radiata plantation in south-eastern Australia has reduced habitat for frogs

Loss and fragmentation of habitat resulting from the clearing of forests for agriculture and urban development threaten the persistence of thousands of species worldwide. The clearing of native forest to plant a monoculture of exotic trees may also reduce and fragment the habitat available for indigenous plants and animals. Metacommunity theory suggests that the species richness of a community in a patch of habitat will increase with patch size but decrease with patch isolation. We investigated whether replacement of native Eucalyptus forest with a plantation of Pinus radiata has reduced and fragmented habitat for frogs, leading to a lower species richness of frog communities in the pine plantation and in small and/or isolated remnant patches of native forest. We surveyed frogs at 60 sites at streams and wetlands in the pine plantation, remnant patches of native forest surrounded by pines, and adjacent areas of contiguous native forest near Tumut in New South Wales, Australia. Only two of eight species of frogs were recorded in the pine plantation, and regression modelling indicated that streams and wetlands in the pines supported fewer frog species than those in remnant patches or the intact native forest. In addition, species richness tended to be higher at wide, shallow swamps and marshes near the headwaters of streams, with herbs, grasses, shrubs, reeds, sedges and rushes in the emergent and fringing vegetation. There was little evidence to suggest that larger eucalypt remnants supported more species of frogs, or that remnants isolated by greater expanses of pines supported fewer species, but we had low power to detect these effects with our data set. Our results support the preservation of all remnants of native forest along drainage lines and around swamps, soaks and bogs, regardless of size. Where new pine plantations are established, for example, on cleared agricultural land, care should be taken to maintain the structural and vegetative characteristics of water bodies to ensure that they continue to provide suitable breeding habitat for frogs.     

Protonephridia in the larvae of the paleonemertean species Carinoma mutabilis (Carinomidae,Nemertea) and Cephalothrix (Procephalothrix) filiformis (Cephalothricidae,Nemertea)

During spiralian development, the first pair of nephridia forms anterior to the mouth. Each organ consists of a few cells, which is characteristic for spiralian larvae. In nemerteans, one of the unambiguously spiralian taxa, so far protonephridia, has been reported only in advanced pilidium larvae, where they likely persist as juvenile and adult nephridia. These organs have not been recorded in larvae of the basally branching nemertean taxa. In search for these organs, we examined the ultrastructure of pelagic planuliform larvae of the palaeonemerteans Carinoma mutabilis and Cephalothrix (Procephalothrix) filiformis. In both species, a pair of protonephridia is located at the level of the stomodaeum. Each protonephridium of C. mutabilis consists of two terminal cells, two duct cells and one nephropore cell, while that of C. filiformis consists of three terminal cells, three duct cells and one nephropore cell. In C. mutabilis and in C. filiformis, all terminal cells contribute to forming a compound filtration structure. In both species, the protonephridia seem to develop subepidermally, since in C. filiformis, the nephropore cells pierce the larval epidermis and in C. mutabilis, the nephropores are initially covered by the binucleated multiciliated trophoblast cells. On the fifth day, these cells degenerate, so that the protonephridium becomes functional. The occurrence of protonephridia in the larvae of both paleonemertean species is in accordance with the hypothesis that a common ancestor of Nemertea and Trochozoa had a larval stage with a pair of protonephridia. This does not contradict previous hypotheses on placing the Nemertea as an ingroup of the Trochozoa or Spiralia (= Lophotrochozoa). Whether these protonephridia are restricted to the larval phase or whether they are transformed into the adult protonephridia, like those of the pilidium larva, remains to be answered.     

Disturbance history influences the distribution of stream invertebrates by altering microhabitat parameters: a field experiment

1. We investigated the effects of local disturbance history and several biotic and abiotic habitat parameters on the microdistribution of benthic invertebrates after an experimental disturbance in a flood‐prone German stream. 2. Bed movement patterns during a moderate flood were simulated by scouring and filling stream bed patches (area 0.49 m²) to a depth of 15–20 cm. Invertebrates were investigated using ceramic tiles as standardized substrata. After 1, 8, 22, 29, 36 and 50 days, we sampled one tile from each of 16 replicates of three bed stability treatments (scour, fill and stable controls). For each tile, we also determined water depth, near‐bed current velocity, the grain size of the substratum beneath the tile, epilithic algal biomass and standing stock of particulate organic matter (POM). 3. Shortly after disturbance, total invertebrate density, taxon richness and density of the common taxa Baetis spp. and Chironomidae were highest in stable patches. Several weeks after disturbance, by contrast, Baetis spp. and Hydropsychidae were most common in fill and Leuctra spp. in scour patches. The black fly Simulium spp. was most abundant in fill patches from the first day onwards. Community evenness was highest in scour patches during the entire study. 4. Local disturbance history also influenced algal biomass and POM standing stock at the beginning of the experiment, and water depth, current velocity and substratum grain size throughout the experiment. Scouring mainly exposed finer substrata and caused local depressions in the stream bed characterized by slower near‐bed current velocity. Algal biomass was higher in stable and scour patches and POM was highest in scour patches. In turn, all five common invertebrate taxa were frequently correlated with one or two of these habitat parameters. 5. Our results suggest that several ‘direct’ initial effects of local disturbance history on the invertebrates were subsequently replaced by ‘indirect’ effects of disturbance history (via disturbance‐induced changes in habitat parameters such as current velocity or food).     

INTRODUCED MACROALGAE IN THE AUSTRALIAN REGION: CURRENT STATE OF KNOWLEDGE

Introductions of non‐native macroalgae and the subsequent displacement of native species are globally becoming more frequent. The algal genera Undaria, Sargassum, Caulerpa and Codium have been identified as being particularly invasive. An overview on the present knowledge on macroalgal introductions in the Australian region is presented and options for management are discussed, mostly using examples from studies on the introduced Japanese kelp, Undaria pinnatifida. Undaria pinnatifida was first detected in Tasmania, Australia in the early 1980's. Since then, its range has expanded despite eradication efforts. Long distance jumps appear to be the major mode of spread of U. pinnatifida in Tasmania. Studies are underway to distinguish the relative importance of spore dispersal, drift of adult plants and anthropogenic factors in spreading this invasive kelp. Although information on the real impacts of U. pinnatifida and other introduced macroalgae is sparse, the development of management and control strategies is of vital importance to prevent further spread and translocation of these “pest” species.     

Simulation of hurricane-like disturbances on a Caribbean seagrass bed

We tested the hypothesis that hurricanes cause changes in the plant community structure of Caribbean seagrass beds by acting selectively on populations of rooted macrophytes, which include seagrasses and rhizophytic algae. We also tested the hypothesis that susceptibility to elimination of the rooted macrophytes by the disturbance depends on differences in their growth forms. Two commonly registered disturbances by hurricanes in shallow seagrass beds are burial and sediment removal, which were simulated in marked plots of 1.1 × 1.1 m, at two randomly selected stations in Puerto Morelos tropical reef lagoon. The treatments consisted of control (no disturbance), two levels of burial, and two levels of sediment removal, with four replicates per treatment per station. The experiment was initiated in July 2002 and, 2 months afterwards, the densities of the populations of macrophytes were measured in experimental units of 0.8 × 0.8 m within the plots. MDS analysis showed that both sediment removal and burial caused changes in the species composition of the seagrass community. At one station, burial had a greater impact than sediment removal, whereas at the other station, the degree of impact of both types of disturbance was similar. Some macrophytes were consistently removed more than others, supporting the selective elimination hypothesis. Populations of Thalassia testudinum Banks ex König, Halimeda spp., and spongy algae (Avrainvillea spp. and Cladocephalus spp.) were, in almost all cases, undamaged by experimental manipulations. The populations of Syringodium filiforme Kütz., brush-like algae (Penicillus spp. and Rhipocephalus spp.), and Udotea spp. were reduced by more than 70%, when averaged across all manipulations and stations. A comparative analysis of growth forms of the above-mentioned macrophytes suggested that a solid, deeply anchored root-rhizome or rhizoid system, combined with a flexible or modular above-ground structure, is an advantageous characteristic to resist perturbation by hurricanes or storms.     

Tracking the origin of the invading Caulerpa (Caulerpales, Chlorophyta) with Geographic Profiling, a criminological technique for a killer alga

The application of the Geographic Profiling technique (with the “Rossmo formula”) proved to be effective in assessing the spreading origin of invading species of Caulerpa in the Mediterranean. Geoprofiling is a technique more frequently used in criminology. We applied this method to an algal invasion for the first time. The method was calibrated with the distribution data of Caulerpa taxifolia, whose spreading in the Mediterranean Sea started from the aquarium of Monaco. This is the first time that Geographic Profiling is calibrated on a data set of sites of presence of a biological invader, of which the spreading origin is known. The application on Caulerpa racemosa var. cylindracea showed that the probable spreading center of the alga should be located in Western Sicily for the Mediterranean and in the southern part of Tenerife for the Canary Islands (Las Galletas, a fishing village). The spreading centers correspond to areas with ports, indicating that the spreading of this alga should be related to (fishing) boats. This result confirmed the opinion of other authors. The parameters used in the Rossmo Formula, obtained through calibration with the known origin of C. taxifolia invasion for identifying the origin of the spread of C. racemosa var. cylindracea may be extended to other algae with similar propagation mode and similar habitat requirements.     

Native species behaviour mitigates the impact of habitat-forming invasive seaweed

Habitat-forming invasive species cause large, novel changes to the abiotic environment. These changes may elicit important behavioural responses in native fauna, yet little is known about mechanisms driving this behaviour and how such trait-mediated responses influence the fitness of native species. Low dissolved oxygen is a key abiotic change created by the habitat-forming invasive seaweed, Caulerpa taxifolia, which influences an important behavioural response (burrowing depth) in the native infaunal bivalve Anadara trapezia. In Caulerpa-colonised areas, Anadara often emerged completely from the sediment, and we experimentally demonstrate that water column hypoxia beneath the Caulerpa canopy is the mechanism instigating this “pop-up” behaviour. Importantly, pop-up in Caulerpa allowed similar survivorship to that in unvegetated sediment; however, when we prevented Anadara from popping-up, they suffered >50% mortality in just 1 month. Our findings not only highlight the substantial environmental alteration by Caulerpa, but also an important role for the behaviour of native species in mitigating the effects of habitat-forming invasive species.     

Epibiosis of Oxygenic Phototrophs Containing Chlorophylls a, b, c, and d on the Colonial Ascidian Cystodytes dellechiajei

The external surfaces of marine animals are colonized by a wide variety of epibionts. Here, we study the phototrophic epibiotic community attached to the colonial ascidian Cystodytes dellechiajei collected in the Mediterranean Sea. Epifluorescence microscopy analysis showed abundant filamentous cyanobacteria on the upper and basal parts of the ascidian that displayed autofluorescence, as well as some unicellular cyanobacteria, diatoms, and structures, which could belong to microscopic rhodophyte algae. In addition, high-performance liquid chromatography of the photosynthetic pigments confirmed that the phototrophic epibionts possess chlorophyll (Chl) d, as well as Chl a, b, and c, which enable them to use far-red light for photosynthesis in that peculiar microenvironment. Furthermore, laser scanning confocal microscopy showed the presence of a few small patches of cells on the basal part of the ascidian displaying fluorescence between 700 and 750?nm after excitement with a 635-nm red laser, typically within the range of Chl d. Denaturing gradient gel electrophoresis of the 16S rRNA gene polymerase chain reaction amplified using specific primers for Cyanobacteria detected sequences related with the genera Planktothricoides, Synechococcus, Phormidium, and Myxosarcina, as well as sequences of chloroplasts of diatoms and rhodophyte algae. Remarkably, only the sequences related to the filamentous cyanobacteria Planktothricoides spp. and some chloroplast sequences were found in almost all specimens collected under different macroecological conditions and geographical areas, suggesting thus certain specificity in the epibiotic association. On the other hand, Prochloron spp. and Acaryochloris marina, typically associated to tropical ascidians, were not detected by denaturing gradient gel electrophoresis. However, given the low abundance of cells displaying Chl d in C. dellechiajei and the fact that molecular fingerprinting techniques not always recover low abundance groups, the presence of these cyanobacteria cannot be ruled out. Nevertheless, our data indicate that tropical ascidians and C. dellechiajei differ in their phototrophic communities, although Chl d-containing cells are present in both microenvironments.     

Local disturbance history and habitat parameters influence the microdistribution of stream invertebrates

1. We investigated the effects of local disturbance history and habitat parameters (abiotic and biotic) on the microdistribution of benthic invertebrates during several floods in two streams, the Schmiedlaine in Germany (four events) and the Kye Burn in New Zealand (two events). 2. Bed movement patterns were quantified using metal‐link scour chains. Before and after each flood, quantitative invertebrate samples were taken from replicate bed patches that had experienced sediment scour, fill or remained stable. 3. Patterns of invertebrate density in the different bed stability types (i.e. scour, fill, stable) varied between floods, sampling dates and streams, but invertebrate density was highest in stable patches in >50% of all the patch type effects detected and lowest in fill patches in 75% of all detected effects. Stable bed patches acted as a refugium for Liponeura spp. and Leuctra spp. in the Schmiedlaine and for Hydracarina and Deleatidium spp. in the Kye Burn. 4. Averaged across both streams, only near‐bed current velocity was correlated with invertebrate distribution on the streambed more often than disturbance history. In the Kye Burn, disturbance history and water depth were the most influential habitat parameters. 5. Our results suggest that a thorough understanding of the microdistribution of benthic invertebrates requires knowledge of disturbance history, as well as more readily measured habitat parameters such as current velocity or water depth.     

The structure of biogenic habitat and epibiotic assemblages associated with the global invasive kelp <Emphasis Type="Italic">Undaria pinnatifida</Emphasis> in comparison to native macroalgae

Kelp forests dominate temperate and polar rocky coastlines and represent critical marine habitats because they support elevated rates of primary and secondary production and high biodiversity. A major threat to the stability of these ecosystems is the proliferation of non-native species, such as the Japanese kelp Undaria pinnatifida (‘Wakame’), which has recently colonised natural habitats in the UK. We quantified the abundance and biomass of U. pinnatifida on a natural rocky reef habitat over 10 months to make comparisons with three native canopy-forming brown algae (Laminaria ochroleuca, Saccharina latissima, and Saccorhiza polyschides). We also examined the biogenic habitat structure provided by, and epibiotic assemblages associated with, U. pinnatifida in comparison to native macroalgae. Surveys conducted within the Plymouth Sound Special Area of Conservation indicated that U. pinnatifida is now a dominant and conspicuous member of kelp-dominated communities on natural substrata. Crucially, U. pinnatifida supported a structurally dissimilar and less diverse epibiotic assemblage than the native perennial kelp species. However, U. pinnatifida-associated assemblages were similar to those associated with Saccorhiza polyschides, which has a similar life history and growth strategy. Our results suggest that a shift towards U. pinnatifida dominated reefs could result in impoverished epibiotic assemblages and lower local biodiversity, although this could be offset, to some extent, by the climate-driven proliferation of L. ochroleuca at the poleward range edge, which provides complex biogenic habitat and harbours relatively high biodiversity. Clearly, greater understanding of the long-term dynamics and competitive interactions between these habitat-forming species is needed to accurately predict future biodiversity patterns.     

The species composition,occurrence and temporal stability of submerged aquatic macrophyte patches along the main channel border of pool 5A,Upper Mississippi River

Species composition, relative abundance, distribution and physical habitat associations of submerged aquatic macrophytes in the main channel border (MCB) habitat of Pool 5A, Upper Mississippi River (UMR) were investigated during the summers of 1980 and 1983. The submerged aquatic macrophytes in Pool .5A MCB were a small and stable component of the river ecosystem. Submerged plants occurred primarily in small, monospecific clumps. Clumps in close proximity to each other formed plant patches. Plant patches were stable in location and number between 1980 and 1983; 82.5% of the patches first observed in 1980 were present in 1983. Submerged macrophytes covered about 10–12 ha of the 201 ha MCB in Pool 5A. Submerged plants were most common in the lower two-thirds of the pool. Ten species of aquatic macrophytes occurred on rock channel-training structures and eleven occurred on non-rock substrates in the MCB. The most common submerged plants, in order of abundance, were Vallisneria americana Michx., Heteranthra dubia Jacq., Potamogeton pectinatus L., Ceratophyllum demersum L. and Potamogeton americanus C. & S.     

The effects of an invasive habitat modifier on the biotic interactions between two native herbivorous species and benthic habitat in a subtidal rocky reef ecosystem

Range expanding species can have major impacts on marine ecosystems but experimental field based studies are often lacking. The urchin Centrostephanus rodgersii has recently undergone a southerly range expansion to the east coast of Tasmania, Australia. We manipulated densities of C. rodgersii and algal regrowth in urchin barrens habitat to test effects of the urchin on biotic interactions between two native herbivores, black-lip abalone (Haliotis rubra) and another urchin (Heliocidaris erythrogramma), and their benthic habitat. After 13 months, removals of only C. rodgersii resulted in overgrowth of barrens habitat by algae and sessile invertebrates. Densities of abalone increased (+92 %) only in patches from which C. rodgersii was removed and algal regrowth allowed. In contrast, densities of H. erythrogramma increased in all treatments (+45, +28, +25 %) in which C. rodgersii was removed, irrespective of the algal regrowth manipulations. These results suggest that C. rodgersii has a negative influence on the densities of abalone through competition for food and on densities of H. erythrogramma through competition for preferred habitat. Densities of abalone (+65 %) but not H. erythrogramma (+25 %), were lower in the patches from which C. rodgersii and canopy algae regrowth were removed relative to patches from which only C. rodgersii was removed (+92 and +28 %, respectively). These results suggest that C. rodgersii overgrazing of canopy-algae results in loss of structural complexity which could increase abalone susceptibility to predation, cause abalone to seek shelter in cryptic microhabitats and/or prevent their return to patches where canopy algae are absent. The ongoing spread of C. rodgersii and expansion of barrens habitat in eastern Tasmania will continue to negatively affect populations of these two native herbivores and their associated fisheries at a range of spatial scales. This example shows that habitat modifying species which become highly invasive can have disproportionate negative impacts on the structure and dynamics of the recipient community.     

Dispersion of the horned grebe Podiceps auritus (L.) (Aves) on Lake Myvatn, Iceland, in late summer

Studies on habitat selection by waterbirds usually describe large-scale distribution of populations or species but the dispersion within lakes is an understudied aspect. Detailed mapping of horned grebes (Podiceps auritus (L.)) on the North Basin of Lake Myvatn, Iceland, was carried out in late summer 2000 in order to compare their dispersion with water depth and submerged vegetation. Depth and vegetation were measured on sampling sites that were evenly distributed over the North Basin, and the information obtained compared to the distribution of the grebes. Four species of submerged macrophytes dominated in the North Basin; Myriophyllum spicatum, M. alterniflorum, Potamogeton filiformis and P. perfoliatus, mainly at depths <2.5 m. The grebes seemed to prefer to dive in shallow areas vegetated with stands of Myriophyllum spp. Densities of three-spined sticklebacks Gasterosteus aculeatus were compared in stands of M. spicatum and P. filiformis in order to estimate food availability. No difference was found, so the sticklebacks could not explain the preference of the grebes for areas vegetated with Myriophyllum spp. Nevertheless, this study indicates that shallow areas, vegetated with Myriophyllum spp., are important feeding areas for the horned grebe.