RECENT RADIATION OF THE PALMARIACEAE (RHODOPHYTA)1

Molecular phylogenetic studies on the evolution of the red algae indicate that this ancient division has many lineages that have recently undergone radiations. One such example is the cold–temperate family Palmariaceae. In this study, sequences from the ribosomal DNA internal transcribed spacer regions were compared among ten species in the Palmariaceae from both Atlantic and Pacific sites, Phylogenetic analyses of sequence data, in which Rhodophysema georgii Batters was used as outgroup and root, indicate a radiation into four clades, three of which contain species of “Palmaria” and the fourth species of Halosaccion. Palmaria palmata (L.) Kuntze, the type and only North Atlantic species in the genus, stands apart from all remaining species in the family and terminates the most basal branch in the rooted tree. The three more derived clades have radiated mainly in the North Pacific. Southern Ocean Palmaria and North Atlantic Devaleraea are hypothesized to have invaded from separate but closely related North Pacific ancestors. The ease with which sequences could be aligned combined with an unsaturated transition: transversion ratio and modest divergence involving predominantly point mutations suggests that the initial radiation is relatively recent (late Miocene–Pliocene) and that the Devaleraea–Palmaria clade is even more recent (late Pliocene–Pleistocene).     

The algal flora of a subtidal Zostera bed in ventry bay,Southwest Ireland

Ventry Bay in County Kerry, southwest Ireland, was surveyed in June 1977 to a depth of 17 m using SCUBA and a grid of stations which were 400 m apart. The principal substrate was sand. Zostera marina L. was dominant and widespread from 0.5 to 13 m below Chart Datum and is the deepest-growing Zostera in Northwest Europe. Sixty species of algae were found and their occurrence and that of Zostera marina was subjected to cluster analysis resulting in one distinct cluster of Zostera marina, Ulva lactuca L., Gracilaria verrucosa (Huds.) Papenf., Polysiphonia elongata (Huds.) Spreng., Ectocarpus sp., Ceramium rubrum (Huds.) C. Ag. and Brongniartella byssoides (Good. et Woodw.) Schmitz. These algae are so common and widespread in the subtidal zone that their co-occurrence with Zostera is not considered to be significant. Zostera occurs virtually as monospecific stands in sand and the co-occurrence of any macroalgae is largely fortuitous and dependent on the heterogeneity of the substrate.     

Phycodrys valentinae sp. nov. (Delesseriaceae, Rhodophyta) with Discussion of Other Species of the Phycodrys Genus from the North Pacific

A new species Phycodrys valentinae Seliv. et Zhigad. from the northwest of the Bering Sea, distinct from other species of the genus Phycodrys by the presence of a distromatic lamina and by the arrangement of basal generative prolifications, was described. Based on the analysis of the morphological and anatomic traits of species of the genus Phycodrys, P. serratiloba (Rupr.) A. Zin., which was considered as synonym of P. riggii Gardn. to the present [27, 28, 30], was reinstated as an independent species. The taxonomic status of other representatives of the genus Phycodrys of the studied region is considered. In our opinion, five species of the genus Phycodrys (P. serratiloba, P. riggii, P. amchitkensis, P. vinogradovae, and P. valentinae) grow in the seas of the Russian Far East.     

Palatability of the Antarctic rhodophyte Palmaria decipiens (Reinsch) RW Ricker and its endo/epiphyte Elachista antarctica Skottsberg to sympatric amphipods

Although many macroalgae that occur throughout Western Antarctic Peninsular waters are known to produce defensive secondary metabolites that deter grazing, the rhodophyte, Palmaria decipiens is palatable to several sympatric meso- and macro-grazers. It has been hypothesized that high levels of mesoherbivory by amphipods may account for the conspicuous lack of filamentous epiphytes emerging from the thalli of marcophytes in this region. Nonetheless, Elachista antarctica is a filamentous phaeophyte found growing within, and emerging from the thallus of the rhodophyte P. decipiens. It is surprising that E. antarctica occurs exclusively in association with a palatable species of macroalgae considering the standing biomass of other chemically defended unpalatable species is very high. We tested the hypothesis that E. antarctica grows on P. decipiens due to the host's overwhelming palatability compared to that of the epiphyte. That is, the hypothesis that mesograzers prefer the host over the epiphyte, grazing around emerging filaments. Choice and no choice feeding assays with live tissues of E. antarctica and P. decipiens were conducted in three different trials with four sympatric amphipod species (Prostebbingia gracilis, Gondogeneia antarctica, Oradarea bidentata, and Paraphimedia integricauda) commonly found in association with P. decipiens. G. antarctica consumed both species but ate P. decipiens at a faster rate than the epiphyte in two of three trials. P. gracilis, O. bidentata, and P. integricauda fed on the epiphyte, E. antarctica at faster rates than upon P. decipiens. Aggressive grazing of the epiphyte by this suite of amphipods indicates that differences in palatability and differences in grazing pressure on host and epiphyte do not explain the exclusive epiphytism of E. antarctica on P. decipiens.     

Community structure of amphipods on shallow <Emphasis Type="Italic">Posidonia oceanica</Emphasis> meadows off Tunisian coasts

The structure, diversity and spatial distribution of the amphipod fauna associated with Posidonia oceanica meadows were studied along the Tunisian coasts in 2007. Samples were collected in nine different meadows at 2 m depth. A total of 44 species belonging to 12 families were collected. The most common species in terms of abundance were Ampithoe helleri, Hyale camptonyx and Ericthonius punctatus. The highest values of abundance and species richness and the lowest values of diversity and equitability were found in meadows with high epiphyte biomass. Multivariate analyses of data indicated that epiphyte biomass and geographical position were major determinants of the distribution and composition of amphipod assemblages along Tunisian coasts. The presence of two lessepsian amphipod species in one of the southern Posidonia oceanica meadows modified the structure of assemblage.     

Differentiation in Coprinus lagopus

Fruit-body expansion was studied in Coprinus lagopus (sensu Buller) following surgical procedures. Elongation occurred after denuding mushroom caps of essentially all peripheral scales. Young primordia (1–5 mm) failed to develop after vertical bisection. Older primordia (e. g. 10 mm) expanded 3–4-fold after vertical bisection or quadrisection, underwent autolysis and basidiospore production. An amorphous brown gel in the stipe lumen disappeared during development of bisected primordia. Stipes isolated from primordia expanded autonomously and exhibited negative geotropism when incubated upside down or when the stipe apex was removed. Displacement of charcoal particles dusted on intact stipes revealed the most active zone of expansion to be the upper mid-region of the stipe. Segmented stipes likewise showed most active elongation in the mid-region. Vertically bisected stipes also expanded.     

Epiphytes from a deep-water characean meadow in an oligotrophic New Zealand lake: species composition,biomass and photosynthesis

1. The epiphytic flora of a characean meadow in Lake Coleridge, a deep, oligotrophic lake on the South Island of New Zealand, was dominated by diatoms, particularly Eunotia pectinalis and Achnanthes minutissima. The meadows occupied a depth range from 5 to 30 m. Adnate taxa predominated at all depths below 5 m, while increased taxonomic diversity at 5 m resulted from an increased abundance of erect taxa, including chlorophytes and stalked diatoms. 2. Seasonal changes in epiphyte biomass were followed using artificial substrata and by estimating epiphyte chlorophyll a concentration on host plants. The latter required development of a novel technique utilizing the consistent relationship between fucoxanthin and chlorophyll a concentrations in the epiphyton. Epiphyte chlorophyll a on host plants varied with depth and host species between 0.1 and 0.3 mg g^–1 dry weight. Maximum epiphyte biomass was at 10–15 m depth. At depths of 15 m and less, epiphyte chlorophyll a reached a maximum of ≈ 200–300 mg m^–2 in mid-summer, while at greater depths maximum biomass was less and coincided with a period of clear water in spring. 3. Photosynthetic carbon fixation was estimated from photosynthesis–radiation curves and estimates of radiation flux at sampling depths. At depths greater than 10 m, variability of the vertical extinction coefficient of lake water rather than seasonal fluctuations in incident radiation were responsible for determining the temporal pattern of production. Chlorophyll a-specific photosynthesis was estimated to peak in summer at 5 m (8 mg mg^–1 day^–1), and in spring at all other depths. 4. Annual epiphyte production was estimated as 27 g C m^–2 year^–1 at 5 m depth, falling to 15 g C m^–2 year^–1 at 15 m and 1 g C m^–2 year^–1 at 30 m. Areal biomass changes tended to be temporally but not quantitatively coupled to estimated in situ photosynthesis, and we hypothesize that epiphyte biomass may have been controlled by grazing gastropod snails.     

Biomass accumulation and shading effects of epiphytes on leaves of the seagrass, Heterozostera tasmanica, in Victoria,Australia

A method is described for estimating the rate of accumulation of epiphyte biomass on leaves of the seagrass, Heterozostera tasmanica (Martens ex Aschers.) den Hartog and for estimating the effect of epiphyte biomass on photosynthesis of the seagrass. Epiphyte biomass was determined by comparison of the weight per unit area of epiphyte-covered and epiphyte-free leaf blades. Epiphyte weight increased as age of the seagrass leaves increased. Linear regression on epiphyte biomass vs. leaf age estimated the rate of biomass accumulation. Rates varied from 5.7 to 104 μg epiphyte dry weight per cm² of leaf surface per day at three sites in Western Port and Port Phillip Bay, Victoria. Rates of accumulation of epiphyte biomass were generally higher during December through March (summer) than in May (autumn), August (winter) or October (Spring). Light attenuation by epiphytes increase linearly with biomass. The rate of biomass accumulation of epiphytes was compared with leaf growth rate, ambient photon flux density in H. tasmanica beds and the photosynthesis—photon flux density curve of H. tasmanica. This comparison demonstrated that epiphyte biomass can accumulate fast enough to shade H. tasmanica leaves and significantly reduce the time (to less than one half of the leaf life span) in which positive net photosynthesis of the leaf blade is possible.     

The reproductive phenology of nine species of Rhodophyta in the subtidal region of the Isle of Man

Monthly samples of about 40 separate plants of each species were collected from 1 to 3 m below lowest astronomical tide on Port Erin breakwater, Isle of Man, Irish Sea. In three species growing on rock, Plocamium cartilagineum, Cryptopleura ramosa and Callophyllis laciniata, about 90% of the plants were fertile in late summer but less than 10% in spring although some fertile plants were always present. Delesseria sanguinea and Odonthalia dentata, also epilithic, had a winter sporing season, Odonthalia extending into late spring, and all plants were sterile in summer. Three species growing epiphytically, Palmaria palmata, Membranoptera alata and Phycodrys rubens, reproduced maximally in the first half of the year at the time when the stipes of the host species, Laminaria hyperborea, grow fastest. Only Palmaria had a sterile season, late summer. The encrusting Cruoria pellita showed little seasonality. The first three species, which reproduce mainly when the sea temperature is above average, are in the northern part of their geographical range. The remaining species (apart from Cruoria) reproduce mainly at low temperatures and are in the southern half of their ranges. Male plants appear to be in a minority in all species, presumably because they were manifest for a shorter period than carposporic plants. They appeared first after sterile periods and were absent as sporing declined. Plocamium and to a lesser extent Cryptopleura show an extremely high preponderance of tetrasporophytes in the population. This is attributed to perennation and some factor disallowing the survival of most of the tetraspores.     

Ecological effects of harvesting Lessonia (Laminariales,Phaeophyta) in central Chile

Lessonia nigrescens and L. trabeculata are economically important canopy-forming kelps in Chile. Experimental harvesting of stipes above the first dichotomy reduces stipe movement and inter-stipe friction, allowing the development of a heavy epiphytic load and increased grazing. Complete stipe removal leads to holdfast death as neither species is able to simultaneously regenerate all stipes. The invertebrate fauna inside the holdfast does not respond to upper canopy changes, but mortality does occur in partial or complete plant removals. Kelp removal also affects inter-plant distances, results in increased access of grazers to the outside and inside of kelp holdfasts, reduces recruitment of other algal species, and modifies the morphology of L. trabeculata such that the plants become more susceptible to removal by water movement.     

Benthic marine algal flora of the north shore of Prince Edward Island (Gulf of St. Lawrence), Canada

A seaweed community along 60 km of shoreline in the southern Gulf of St. Lawrence has been studied for 5 years. Because of severe ice action along the shores, the algal flora is largely restricted to subtidal levels, which are accessible only during the ice-free months, April to early December. A total of 52 Rhodophyta, 50 Phaeophyta, 19 Chlorophyta and 5 Cyanophyta was identified, mostly occurring between mean low water and 10 m depth. The normally intertidal dominants Focus vesiculosus L. and Chondrus crispus Stackh. flourish at depths of 0–2.5 m and 2.5–5 m, respectively. Chondrus and Furcellaria lumbricalis (Huds.) Lamour. co-dominate the algal benthos at 5–7.5 m, and F. lumbricalis becomes prevalent at 7.5–10 m. Phyllophora truncata (Pallas) A. Zin. and Phyllophora pseudoceranoides (Gmel.) Newr. & A. Tayl. are the main perennials at 10–15 m, and often at 15–20 m as well. Below 15 m, the flora diminishes markedly, and at 20–25 m consists chiefly of Agarum cribrosum (Mert.) Bory, crustose corallines and Polysiphonia urceolata (Lightf. ex Dillw.) Grev. Laminarians other than Chorda occur only as scattered plants or small patches and do not exceed 1 m length. The algal community is relatively undiversified and is especially deficient in green algae. Despite higher surface-water temperatures during summer, the flora is essentially a cold-water one as is indicated by both composition and low R/P and (R+C)/P ratios of 1.06 and 1.45, respectively.     

Variations in the alginate content and composition of Durvillaea antarctica and D. willana from southern New Zealand

The brown seaweeds Durvillaea antarctica andD. willana are dominant components of the lowerlittoral and upper sublittoral of exposed rocky shoresin southern New Zealand. Tissue samples of bothspecies, harvested from a site on the south-east coastof South Island over a period of 2 years, wereanalysed for alginate content and composition.Individuals of both species were further separatedinto different blade (lamina and palm) and stipe(cortex and medulla) fractions to assess variationwithin the thallus. On average the alginate contentand frequency of mannuronic acid (F_m) was higherin D. antarctica than in D. willana. Blades contained more alginate than stipes, laminaeand stipes were rich in mannuronic acid whereasholdfasts were rich in guluronic acid. Variations incomposition are considered to reflect the functionaldifferences of the tissue, giving flexibility to bladeand stipe and rigidity to the holdfast. Despitefluctuations in content and composition betweencollection times no seasonal trends in eithercomponent were apparent.     

Long-Term Observations of Epibenthic Fish Zonation in the Deep Northern Gulf of Mexico

A total of 172 bottom trawl/skimmer samples (183 to 3655-m depth) from three deep-sea studies, R/V Alaminos cruises (1964–1973), Northern Gulf of Mexico Continental Slope (NGoMCS) study (1983–1985) and Deep Gulf of Mexico Benthos (DGoMB) program (2000 to 2002), were compiled to examine temporal and large-scale changes in epibenthic fish species composition. Based on percent species shared among samples, faunal groups (≥10% species shared) consistently reoccurred over time on the shelf-break (ca. 200 m), upper-slope (ca. 300 to 500 m) and upper-to-mid slope (ca. 500 to 1500 m) depths. These similar depth groups also merged when the three studies were pooled together, suggesting that there has been no large-scale temporal change in depth zonation on the upper section of the continental margin. Permutational multivariate analysis of variance (PERMANOVA) also detected no significant species changes on the limited sites and areas that have been revisited across the studies (P>0.05). Based on the ordination of the species shared among samples, species replacement was a continuum along a depth or macrobenthos biomass gradient. Despite the well-known, close, negative relationship between water depth and macrofaunal biomass, the fish species changed more rapidly at depth shallower than 1,000 m, but the rate of change was surprisingly slow at the highest macrofaunal biomass (>100 mg C m⁻²), suggesting that the composition of epibenthic fishes was not altered in response to the extremely high macrofaunal biomass in the upper Mississippi and De Soto Submarine Canyons. An alternative is that the pattern of fish species turnover is related to the decline in macrofaunal biomass, the presumptive prey of the fish, along the depth gradient.     

Tide-model analysis of the zonation of intertidal prosobranchs. I. Four species of littorina (L.)

Experiments were carried out in a tide model to determine the patterns of zonation of four species of periwinkles under laboratory conditions. Both upper shore littorinids, Littorina neritoides (L.) and L. saxatilis (Olivi), showed the same zonation in relation to the tidal cycle as observed on the shore at Plymouth. In the model, the animals adopted a zone above the level of high water and then remained inactive. L. littoralis (L.) showed the same distribution unless fucoid algae were provided, in which case the periwinkles clustered on them. L. littorea (L.) showed the ability to adopt a pattern of zonation in the model, which was at, and just below, high tide level, but higher than that normally observed on the shore.     

Stipe wall extension of Flammulina velutipes could be induced by an expansin-like protein from Helix aspersa

Expansin proteins extend plant cell walls by a hydrolysis-free process that disrupts hydrogen bonding between cell wall polysaccharides. However, it is unknown if this mechanism is operative in mushrooms. Herein we report that the native wall extension activity was located exclusively in the 10 mm apical region of 30 mm Flammulina velutipes stipes. The elongation growth was restricted also to the 9 mm apical region of the stipes where the elongation growth of the 1st millimetre was 40-fold greater than that of the 5th millimetre. Therefore, the wall extension activity represents elongation growth of the stipe. The low concentration of expansin-like protein in F. velutipes stipes prevented its isolation. However, we purified an expansin-like protein from snail stomach juice which reconstituted heat-inactivated stipe wall extension without hydrolytic activity. So the previous hypotheses that stipe wall extension was resulted from hydrolysis of wall polymers by enzymes or disruption of hydrogen bonding of wall polymers exclusively by turgor pressure are challenged. We suggest that stipe wall extension may be mediated by endogenous expansin-like proteins that facilitate cell wall polymer slippage by disrupting noncovalent bonding between glucan chains or chitin chains.     

Reproductive ecology of four subtidal red algae

The reproduction and stature of the red algae Callophyllis cristata (C. Ag.) Kuetz., Membranoptera alata (Huds.) Stackh., Phycodrys rubens (Huds.) Batt., and Ptilota serrata Kuetz. were recorded from subtidal populations at Appledore Island, Maine, U.S.A., with respect to time and depth. Only Membranoptera alata exhibited a conspicuous seasonal fluctuation of reproduction. A vertical gradient of reproduction was evident, with reduced levels of reproduction in shallow populations of Phycodrys rubens and Ptilota serrata, as well as deep populations of Phycodrys rubens, Membranoptera alata and Callophyllis cristata. Differential stratification of the reproductive phases of Ptilota serrata occurred with higher frequencies of tetrasporic plants in deep populations and of cystocarpic plants in shallow populations. In contrast, the haploid and diploid plants of the other three species showed similar distributional patterns. Membranoptera alata, Phycodrys rubens, and Callophyllis cristata showed a conspicuous decrease in stature during maximum reproduction.     

Recolonization of vascular epiphytes in a shaded coffee agroecosystem

Aim: Shaded coffee plantations constitute an important refuge for biodiversity. Despite the fact that epiphytic plants form a significant component of these agroecosystems, their removal from the shade trees is commonplace in Latin America. To what extent does the epiphyte community recover from this severe disturbance? Location: Shaded coffee agroecosystem in Veracruz, Mexico (19°28′03″ N, 96°55′58″ W; 1200 m asl). Methods: We assessed the diversity, biomass and recolonization patterns of vascular epiphytes in shade trees, 8‐9 yr after complete epiphyte removal (E^?), and in control ‘non‐removal’ sites (E⁺). In order to evaluate the effects of prior epiphyte removal, all vascular epiphytes were completely removed from 10 trees per treatment (E^? and E⁺); all epiphyte species collected were identified and dry biomass measured. Results: Eight to nine years after removal, epiphyte biomass in the E^? shade trees was 35% of that found in the control sites. A total of 55 epiphyte species, belonging to 12 families, were registered; 40 in E^?, and 48 in E⁺. Six species belonging to Bromeliaceae, Orchidaceae, Cactaceae and Araceae accounted for 75% of the biomass in E⁺ while six species of bromeliads accounted for 76% of the biomass in E^?. Some bromeliads proliferated following disturbance; however, ferns showed lower recovery. Conclusions: Epiphyte community recovery, in terms of biomass and diversity, is considerably higher in the coffee plantation than has been previously reported for other tropical ecosystems. Epiphyte recolonization patterns reflected both the abundance of species in the surrounding matrix and certain species‐specific traits. For such agroecosytems to function as effective reservoirs of epiphyte diversity, epiphyte stripping should be avoided.     

Biomass and primary production of a 8–11 m depth meadow versus <3 m depth meadows of the seagrass Cymodocea nodosa (Ucria) Ascherson

Current knowledge about the abundance, growth, and primary production of the seagrass Cymodocea nodosa (Ucria) Ascherson is biased towards shallow (depth <3 m) meadows although this species also forms extensive meadows at larger depths along the coastlines. The biomass and primary production of a C. nodosa meadow located at a depth of 8–11 m was estimated at the time of maximum annual vegetative development (summer) using reconstruction techniques, and compared with those available from shallow meadows of this species. A depth-referenced data base of values at the time of maximum annual development was compiled to that end. The vegetative development of C. nodosa at 8–11 m depth was not different from that achieved by shallow (depth <3 m) meadows of this species. Only shoot density, which decreased from 1637 to 605 shoots m⁻², and the annual rate of elongation of the horizontal rhizome, which increased from 23 to 71 cm apex⁻¹ year⁻¹, were different as depth increased from <3 to 8–11 m. Depth was a poor predictor of the vegetative development and primary production of C. nodosa. The biomass of rhizomes and roots decreased with depth (g DW m⁻² = 480 (±53, S.E.) − 32 (±15, S.E.) depth (in m); R² = 0.12, F = 4.65, d.f. = 35, P = 0.0381) which made total biomass of the meadow to show a trend of decrease with depth but the variance of biomass data explained by depth was low. The annual rate of elongation of the horizontal rhizome showed a significant positive relationship with depth (cm apex⁻¹ year⁻¹ = 18 (±5.1, S.E.) + 5.0 (±1.33, S.E.) depth (in m); R² = 0.50, F = 14.07, d.f. = 14, P = 0.0021). As shoot size and growth did not change significantly with depth, the reduction of shoot density should drive any changes of biomass and productivity of C. nodosa as depth increases. The processes by which this reduction of C. nodosa abundance with depth occur remain to be elucidated.     

Effects of fish nests on pattern and zonation of submersed macrophytes in a softwater lake

Nests of centrarchid fishes maintain patches of diaspore-propagated submersed aquatic plants (principally Elatine minima (Nutt.) and Isoetes braunii Dur.) between 0.9 and 2.4 m depth in a softwater oligotrophic lake. Nests are cleared of vegetation in May–June, and abandoned by fish in June. By August, many nests are colonized by diaspore-propagated species that do not spread vegetatively on horizontal stems. About 75% of the nests are reoccupied by fish the following May, and some new nests are formed. Nests not re-occupied by fish are colonized by rhizomatous plants of 4 species that have not been observed to flower or set seed in the lake. Rhizomatous plants reach frequencies near 100% by 14 months after abandonment. Pattern analyses show that patch sizes of diaspore-propagated species correspond to the size of fish nests. Conventional explanations of macrophyte zonation based on depth, exposure, and competition are insufficient in this lake; effects of disturbance must be considered as well.