Unique distribution of epiphytic Neosiphonia harveyi (Rhodomelaceae,Rhodophyta) along sargassacean hosts

Although epiphytism is generally heaviest on older portions of the host thallus and both variation and abundance of epiphytes decrease with proximity to meristematic apices, Neosiphonia harveyi (Bailey) Kim, Choi, Guiry and Saunders was frequently found on upper parts of sargassacean hosts. This study compared density, thallus length and fertility of epiphytic N. harveyi among different regions of Sargassum patens C. Agardh thalli to reveal a unique distribution pattern of this epiphyte. The majority of epiphytic N. harveyi was observed on the upper part of host thalli and was seldom seen near the basal part. Over a 3‐month period, mean epiphyte density was greater on host apices while thallus length and fertility were greater on subapical portions of host thalli, suggesting the colonization by this epiphyte near the apical part of the host. Distribution patterns of this epiphyte were similar among S. patens growing at different depths. Apical portions of S. patens appear to be more suitable substrata for N. harveyi settlement and colonization compared with other portions of the host thallus, regardless of depth.     

Effect of grazing by isopods and amphipods on growth of Ulva spp. (Chlorophyta)

Eutrophication of shallow coastal waters often leads to blooms of macroalgae. Grazing by crustaceans, such as amphipods and isopods, can reduce macroalgal biomass accumulation. At the same time, growth of the macroalgae can be stimulated by epiphyte removal. The role of grazing by isopods and amphipods on Ulva spp. biomass development was investigated in the Veerse Meer, a brackish lagoon situated in the southwest Netherlands. Exclusion of grazing in the field did not stimulate Ulva spp. growth. In fact, growth rates were higher in exclosures that allowed grazers to enter. Edibility tests identified the amphipod Gammarus locusta, and the isopods Idotea chelipes and Sphaeroma hookeri as potential grazers on Ulva spp. However, when epiphytic diatoms were present on the Ulva spp. thalli, Gammarus and Sphaeroma grazed on ephiphytes and not on Ulva tissue. Only Idotea continued to graze on Ulva spp. A laboratory growth experiment revealed a positive effect of Gammarus presence on Ulva spp. growth, probably caused by preferential removal of epiphytic diatoms from the Ulva spp. thalli. The growth stimulation by epiphyte removing grazers such as Gammarus may explain the higher growth rates in the presence of grazers observed in the field. When determining the potential role of invertebrate grazers in controlling macroalgal biomass accumulation, it is important to include an assessment of the epiphyte abundance on the macroalgae, as preferential removal of epiphytes may stimulate growth and thus have the opposite effect.     

ROLE OF SURFACE WOUNDS AND BROWN ALGAL EPIPHYTES IN THE COLONIZATION OF ASCOPHYLLUM NODOSUM (PHAEOPHYCEAE) FRONDS BY VERTEBRATA LANOSA (RHODOPHYTA)1

Ascophyllum nodosum (L.) Le Jol. forms extensive beds in wave‐sheltered, rocky intertidal habitats on the northwestern Atlantic coast. This fucoid seaweed is host to an obligate red algal epiphyte, Vertebrata lanosa (L.) T. A. Chr. [=Polysiphonia lanosa (L.) Tandy], and two facultative brown algal epiphytes, Elachista fucicola (Velley) Aresch. and Pylaiella littoralis (L.) Kjellm. Although V. lanosa can occur throughout most of the length of host fronds, it largely predominates in midfrond segments. The two brown algal epiphytes are restricted to distal segments. Through field experiments conducted in Nova Scotia, Canada, we tested the hypothesis that surface wounds are required for the colonization of distal segments of host fronds by V. lanosa. Distal tissues normally have a smooth surface because of their young age (A. nodosum fronds grow apically). By creating small wounds that mimicked grazing wounds distributed elsewhere on host fronds, we demonstrated that V. lanosa can colonize distal frond segments during the growth and reproductive season (summer and autumn). Approximately half of the artificial wounds were colonized by V. lanosa during this time. The experimental exclusion of both brown algal epiphytes from distal frond segments did not affect colonization by V. lanosa. Thus, we conclude that the absence of surface irregularities on distal segments of host fronds, specifically small wounds, is the main factor explaining the absence of V. lanosa there. We propose that further experimental work clarifying epiphyte distribution in host beds will enhance our ability to understand the functional role of epiphytes in intertidal ecosystems.     

Temporal dynamics of algal epiphytes on Leathesia marina and Colpomenia sinuosa macrothalli (Phaeophyceae)

This is the first study on the temporal dynamics of the epiphyte communities of Leathesia marina and Colpomenia sinuosa, in terms of richness, diversity and abundance, and their relationships between the hosts and the environmental factors. A total of 31 epiphyte taxa were recorded, with the two hosts sharing 64.5% of the epiphytic species found. The red algal epiphytic group had the highest abundance on both hosts and the diatoms presented the highest species richness. The epiphyte species on L. marina showed a progressive increase in abundance during the warm seasons that was related to the end of the macroscopic life stage of the host. They also showed a greater specific richness on L. marina than on C. sinuosa, but a lower diversity and evenness, and a clear dominance of the species Urospora penicilliformis and Erythrotrichia carnea. Low abundance of epiphytes was found on C. sinuosa, without temporal variation, but high diversity and evenness and without any dominant species. In addition, a clear zonation was observed on this host, where epiphytic abundance decreased on the lateral and upper part of the thallus.     

Epi-endophytic symbiosis between Laminariocolax aecidioides (Ectocarpales, Phaeophyceae) and Undaria pinnatifida (Laminariales, Phaeophyceae) growing on Argentinian coasts

The present is the first study on epi-endophytic algae on thalli of Undaria pinnatifida growing along Argentinian coasts. The main goal is to describe the nature and the morphology of this symbiosis. Individuals of Laminariocolax aecidioides were detected in both June and December 2004, growing on U. pinnatifida sporophytes. In nature, the epi-endophyte were macroscopically observed as dark zones that partially covered the hosts’ fronds. L. aecidioides vegetative thalli were irregularly branched uniseriate filaments. The life cycle is described from laboratory cultures started from Patagonian populations. Caryology revealed that the sporophytic diploid phase presented 16 chromosomes whereas the gametophytic haploid phase presented 8 chromosomes. Isolates made from thalli growing in the interior of infected hosts developed into filamentous, branched sporophytes that reproduced by both unispores and plurispores that were produced in unilocular and plurilocular sporangia, respectively. The results of this paper also allowed us to conclude that L. aecidioides uses the thalli of U. pinnatifida as a proper substrate. The penetration of endophitic filaments among the host′s cortical cells produced a lateral compression and, finally, their thalli development generated perforations in the host tissues. The effects of the epi-endophytic infection of L. aecidioides on U. pinnatifida are neither severe nor deleterious.     

EFFECTS OF THE SYMBIOTIC RED ALGA HYPNEOCOLAX STELLARIS ON ITS HOST HYPNEA MUSCIFORMIS (HYPNEACEAE,GIGARTINALES)1

The presence of the “symbiotic” red alga Hypneocolax stellaris Borgesen was found to substantially reduce the growth rate of Hypnea musciformis (Wulfen) Lamouroux. The growth rate of injected field material was 40% less than unifected thalli. Likewise, the growth rate of host thalli infected in the laboratory was reduced 70% compared to uninfected thalli. Neither the quantity nor quality of carrageenan extracted from Hypnea musciformis was directly affected by the presence of Hypneocolax stellaris. In a mariculture system the presence of a symbiotic alga on its host would be expected to reduce the host's growth rate and thereby lower its economic potential.     

Lipid characterization of red alga Rhodymenia pseudopalmata (Rhodymeniales,Rhodophyta)

Rhodymenia pseudopalmata is a red alga that grows at the Caribbean coast of the Yucatan Peninsula and has been proven successful in cultivation. In this study we present the lipid composition of R. pseudopalmata collected from wild populations during three different seasons of 2013. Cultured material was also analyzed and compared in order to evaluate its value as feedstock for biotechnological uses. Thin layer chromatography, ¹ H and ¹³ C NMR spectroscopy and gas chromatography‐mass spectrometry were used to assess variations in their lipid composition. Our results showed that the dominant lipid classes were phospholipids both in wild and cultured materials. The phospholipids phosphatidylcholine and phosphatidylglycerol and the glycolipid monogalactosyldiacylglycerol were present in both wild and cultured R. pseudopalmata, whereas the phospholipid lysophosphatidylcholine was only found in wild material. Fatty acids (FAs) showed a high monounsaturated FAs (MUFAs) content with oleic acid (C18:1ω9) as the dominant compound (78 and 94% of the MUFAs for wild and culture materials, respectively). Saturated FAs (SFAs) represented approximately 90% of the total fatty acid content, with palmitic acid (C16:0) reaching approximately 83% of the SFA content. Rhodymenia pseudopalmata was low in polyunsaturated FAs when compared to other red algae. Other compounds such as 1‐heptadecene, 1‐hexadecene, 15‐heptadecenal, 3‐eicosene 6,10,14‐trimethyl‐2‐pentadecanone, phytol, and heptadecane were also found. Lipid composition differences between the wild and cultured algae suggest that light and nutrients can be manipulated to modify lipid composition. Based on its lipid composition and cultivation feasibility, R. pseudopalmata could be a potential source for nutraceuticals and biofuels production.     

Research note: Field evaluation of epiphyte recruitment (Vertebrata lanosa,Rhodophyta) in different microsite types on host fronds (Ascophyllum nodosum,Phaeophyceae)

The seaweed Ascophyllum nodosum (Phaeophyceae, Fucales) often forms extensive beds in wave‐sheltered, rocky intertidal habitats on northern Atlantic shores. Since this alga is an ecosystem engineer that influences benthic biodiversity, it is important to understand the factors that regulate its performance. Epiphytism is known to affect the performance of macroalgal hosts. In this study, we investigated the effects of surface irregularities on A. nodosum fronds (wounds, branch axils, and lateral pits resulting from receptacle shedding) on the recruitment rate of its obligate epiphyte Vertebrata lanosa (=Polysiphonia lanosa, Rhodophyta, Ceramiales). For this purpose, we performed a field experiment in Nova Scotia, Canada. In June–July 2007, we created wounds on the surface of host fronds that mimicked the wounds that result from invertebrate grazing. At that time, we also mapped the position of epiphyte‐free lateral pits and branch axils on host fronds. In October 2007, after the reproductive season for V. lanosa, the percentage of microsites colonized by this epiphyte was statistically similar for wounds and lateral pits, but significantly lower for branch axils, although by a small difference (mean recruitment rates ranged between 37 and 50%). Since V. lanosa is known not to colonize smooth frond surfaces, our study suggests that the degree of epiphyte load on A. nodosum beds should largely be affected by the overall amount of surface irregularities, with little influence of the relative availability of each microsite type.     

Epiphytism in a subtidal natural bed of Gracilaria gracilis of southwestern Atlantic coast (Chubut, Argentina)

The diversity of epiphytes, their temporal abundance variation and the anatomical structure of host–epiphyte interfaces were studied in the agarophyte Gracilaria gracilis from a natural bed in Bahía Bustamante, Chubut Province, Argentina. Twenty-nine algal species were recorded as epiphytes during 2 years of monthly sampling (March 2006–February 2008). Total epiphyte density ranged between 0.037 ind. cm^?2 (November 2006) and 39.37 ind. cm^?2 (April 2007), with higher density values throughout the second sampling year. Ceramiales species were the most abundant epiphytes. The density of Ceramium rubrum ranged from 0.09 ind. cm^?2 (52 % of the total amount) in September 2006 to 17.4 ind. cm^?2 (44.18 % of the total amount) in April 2007. Epiphyte infection was more dependent on spore recruitment and sporeling development, especially on thalli derived from fragmentation, than on seasonal environmental variations. The different infections were analysed, taking into account the epiphytic attachment strength and invasiveness and the degree of damage inflicted on the host. Calothrix confervicola was one of the most abundant species. This epiphyte, weakly attached to the host surface, generated no host tissue damage. In contrast, C. rubrum, Polysiphonia abscissa and other Ceramiales were the species that caused more damage to the host because their rhizoids penetrated the cortical portion of the host thallus, sometimes reaching the medullary tissue. Some generalisations and characterisations of the different epiphyte groups in relation to their consequences to Gracilaria spp. are presented.     

Life histories of two ichneumonid parasitoids of Cyclosa octotuberculata (Araneae): Reclinervellus tuberculatus (Uchida) and its new sympatric congener (Hymenoptera: Ichneumonidae: Pimplinae)

In Japan, two species of the genus Reclinervellus were found to attack a single host spider, Cyclosa octotuberculata (Araneae). One of these, Reclinervellus tuberculatus comb. nov., winters as a tender larva and has at least two generations a year. This species laid its egg on the anterior face of the host's abdomen. Prior to laying an egg the female repeatedly rubbed her ovipositor over the dorsal and lateral surfaces of the host's abdomen near the base for more than 25 min. The hatched larva fixed itself at the position where the egg was laid, and developed by consuming the host's body fluids. The other species, R. masumotoi sp. nov., is closely related to R. tuberculatus but is easily distinguished from it by having a very weakly reclivous Cu vein in the hindwing, a less developed carina of the propodeum, a shorter ovipositor and more blackish coloration of the mesoscutum. Although this species is sympatric with R. tuberculatus and attacks the same host, the position of the egg and larva on the host's body is quite different from that of R. tuberculatus. The egg of this species was laid on the posterior surface of the host's abdomen and the larva matured 10–20 days earlier than those of R. tuberculatus. Via these modes of parasitism R. masumotoi seemed to escape from competition with R. tuberculatus at the stage of oviposition and out‐competed it when a single host individual bore larvae of both species. Previous records of parasitoids reared from C. octotuberculata were reviewed and found to include records of both species.     

Reproduction of the colonial hydroid Obelia geniculata (L., 1758) (Cnidaria,Hydrozoa) in the White Sea

Populations of the colonial hydroid Obelia geniculata in the White Sea reproduce asexually by frustule formation. Young medusae appear in the plankton during July and August. The number of medusae rarely exceeds 36 per m³, and the average number varies every year from 0.4 to 10 per m³. The size of medusae is smaller than reported from other regions. The umbrella of the largest recorded medusa was only 0.57 mm in diameter and the specimen had just 35 tentacles. Only a few mature medusae were found during the study. The colonies in the White Sea are epiphytic and grow only on laminarian thalli. At the beginning of July there are no colonies on thalli from the upper subtidal zone. By the end of August, colonies of O.␣geniculata had increased in density to 30 per m². Hydroid recruitment was attributed to active frustule production by colonies living below that zone. The frustules detach from the stems of the hydroids and are found in plankton. Production of frustules on branches occurs continuously during colony growth until water temperatures climb above 0 °C. We found that water temperature in this Arctic environment is generally too low for medusa maturation and planula development in the species. Propagation by frustule formation is the principal means of reproduction in Obelia geniculata within the White Sea, and this phenomenon accounts for the species being a dominant epiphyte on laminarian thalli there.     

Gametophyte-sporophyte coalescence in populations of the intertidal carrageenophyte <Emphasis Type="Italic">Mazzaella laminarioides</Emphasis> (Rhodophyta)

Mazzaella laminarioides has consistently been reported as a typical coalescent/clump species with a triphasic life history of the Polysiphonia-type in which the haploid gametophyte is the predominant phase with respect to the diploid sporophyte. Preliminary observations of intertidal populations revealed that, in some instances, cystocarpic and tetrasporic fronds emerged from the same clump (G-T clumps), implying a coalescent process of haploid and diploid thalli by fusion of their corresponding adjacent basal holdfasts. Population surveys at three sites in Coliumo Bay, central Chile, were carried out to characterize frond demography as well as to asses the frequency of gametophyte-tetrasporophyte (G-T) coalescence. Visual and resorcinol methods were employed to determine the phases of the fronds collected over central transects of 15 randomly sampled clumps. Coalescence of G-T clumps was infrequent, with gametophytes dominating over tetrasporophyte thalli.     

Effect of epiphytism on reproductive and vegetative lateral formation in the brown, intertidal seaweed Ascophyllum nodosum (Phaeophyceae)

In the brown alga Ascophyllum nodosum (L) Le Jolis, a common species on sheltered Northern temperate rocky shores, gametes are produced in receptacles that emerge from small depressions (lateral pits) along the branched frond. These lateral pits are also the preferred settling site for the obligate epiphyte Polysiphonia lanosa (L) Tandy. Therefore, epiphytism can be expected to interfere with host reproductive output. The present study investigated the potential impact of the epiphyte on A. nodosum in two series of laboratory experiments that measured: (i) the direct shading of the host plant underneath an epiphyte canopy; and (ii) the development of receptacles in clean and epiphytised A. nodosum segments (excised from individual fronds) over a 6 month period. These experiments showed that light reaching emerged fronds underneath a dense epiphyte cover was reduced by 40%, and this was independent of the degree of desiccation the epiphyte experienced. Concurrently, in the growth study with epiphytised A. nodosum segments (segments with one clean and one epiphytised lateral pit) total receptacle biomass per epiphytised fragment was significantly reduced compared with clean segments (0.52 g and 1.25 g per gram of frond segment, respectively), although this effect was only significant in A. nodosum from sheltered shores. However, expressed as biomass per lateral pit, receptacle biomass in the remaining clean lateral pits in epiphytised segments was significantly increased in segments from both shores, demonstrating that A. nodosum can at least partially compensate for the loss of production resulting from epiphytism.     

Porphyra yezoensis bei Helgoland — eine entwicklungsgeschichtliche Studie

APorphyropsis-like epiphytic specimen found in the harbour of Helgoland was grown in culture and proved to be identical with the JapanesePorphyra yezoensis. Life history studies on this economically important alga resulted in some interesting and hitherto unknown details. The variability of the adult frond is fundamentally determined by the pattern of spore germination. Settled onChaetomorpha filaments, monospores elongate within 20 minutes; the epiphytic germlings are attached to the substrate by a typical basal cell and give rise exclusively to elongated fronds provided with a cuneate base. Unattached spores, however, germinate into buds with rhizoids; they develop into elongated elliptical to oval fronds provided with round or cordate bases. Only plants with male areas were observed in the cultures, butConchocelis was abundantly produced from cells of aged thalli. Grown in mussel-shells, the filamentous phase liberated conchospores for a long time.

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Herrn Paul-Heinz Sahling zum 75. Geburtstag gewidmet, mit Anerkennung und Dank für 50jährige Zusammenarbeit.     

SURFACE INTERACTIONS OF THE EPIPHYTIC MACROALGA HINCKSIA MITCHELLIAE (PHAEOPHYCEAE) WITH THE SHOALGRASS HALODULE WRIGHTII (CYMODOCEACEAE)1

Meadows of Halodule wrightii (Cymodoceaceae) underwent a decline in a tidal flat located at Paranaguá Bay (Parana, SE Brazil). This decline appeared to be related to an overgrowth of the epiphytic macroalga Hincksia mitchelliae (Harv.) P. C. Silva (Phaeophyceae). In order to characterize the type of epiphytism between the alga and its plant host, we compared two samples from the beginning and end of the algal overgrowth via electron and optical microscopes. The investigation revealed that at both sampling periods, there was an epiphytism of type II, which is due to an infection of epiphytes strongly attached to the surface of the host but not associated to any apparent direct host‐tissue damage. The presence of plasmodesmata between the cells of Hincksia only in the late stage of the host–epiphyte interaction indicated a change in the vegetative organization of Hincksia in relation to its host to improve nutrient absorption and distribution through the epiphyte cells. This is the first report on plasmodesmata in H. mitchelliae. The proposed mechanisms with which the algal epiphytes lead seagrasses to death are shadowing by adhesion on Halodule surface and disruption of its osmoregulatory system. Our findings have implications for the conservation and management strategies of seagrass ecosystems.     

Ceramialean epiphytism in an intertidal Gracilaria chilensis (Rhodophyta) bed in southern Chile

The cultivation of the agarophytic red alga Gracilaria has become an activity of major importance in several parts of the world. However, Gracilaria cultivation in Chile still faces problems such as epiphytism. We report ceramialean epiphytism abundance, recruitment patterns and the microscopic stages fixed on the Gracilaria thalli in an intertidal bed (Metri bay) of southern Chile. Using a factorial field experiment, we analyze the effect of intertidal elevation and the use of epiphyte free inoculum on the abundance of ceramialean epiphytic algae. This evidence is used to provide management and control recommendations for ceramialean epiphytes. The results show a summer recruitment and increase in abundance of ceramialean epiphytes. A seasonal and spatial pattern of abundance of microscopic stages of this epiphytic algae was also found, showing a significant increase from the apical (new tissues) to the central parts of the thalli (older tissues). Apical tissues are free of epiphytic propagules. The epiphyte-free tissue zone decreases significantly from winter to summer. Also, a significantly higher density of microscopic stages of ceramialean epiphytes was found on the thalli collected in the farming are a during the summer at low tidal levels. The experimental results indicated that the abundance of epiphytes was greater at the lower tidal levels than at the higher distribution limits of Gracilaria and it was not possible to control epiphytism by manual cleaning. This revised version was published online in September 2006 with corrections to the Cover Date.     

QUANTIFYING GROWTH AND CALCIUM CARBONATE DEPOSITION OF CALLIARTHRON CHEILOSPORIOIDES (CORALLINALES,RHODOPHYTA) IN THE FIELD USING A PERSISTENT VITAL STAIN1

Growth and calcium carbonate deposition rates of the coralline alga Calliarthron cheilosporioides Manza were quantified by monitoring fronds in the intertidal zone that had been chemically labeled with the nontoxic fluorescent brightener Calcofluor white. This vital stain effectively labeled apical meristems of coralline thalli in the field: fronds exposed for only 5 min had detectable chemical marks at least 1.5 years later. By distinguishing portions of thalli that developed before and after exposure, this methodology permitted accurate measurement of growth and calcium carbonate deposition at each meristem. In Calliarthron, meristematic activity declined with increasing frond size. However, because growing fronds dichotomize, the total number of meristems and the deposition rate of new calcified tissue both increased with frond size. Growth rates reported here suggest that large fronds may not be as old as previously estimated. The Calcofluor white method may improve demographic studies of corallines by resolving growth and age of fronds in the field and may facilitate studies of climate change on calcium carbonate deposition in these ecologically important, calcifying algae.     

Seasonal Occurrences of Epiphytic Algae on the Commercially Cultivated Red Alga Kappaphycus Alvarezii (Solieriaceae, Gigartinales, Rhodophyta)

Common problems faced in farming of the red algal genus Kappaphycus/Eucheuma are “ice-ice disease” and the occurrence of epiphytes. Considerable work has been documented on “ice-ice disease” and it's mode of infection but limited information is available on the emergence of epiphytes. The present study addresses the phenomenon of epiphyte infection, its prevalence in commercially cultivated red alga, Kappaphycus alvarezii, and their variability associated with seasonality. Cultured seaweed became susceptible to epiphytes in the dry seasons (1) between March – June and (2) September – November. Findings revealed Neosiphonia savatieri (Hariot) M. S. Kim et I. K. Lee, as the dominant infecting epiphyte, representing up to 80–85% of the epiphyte present during peak seasons. Besides N. savatieri, Neosiphonia apiculata, Ceramium sp., Acanthophora sp. and Centroceras sp. were observed in smaller quantities. SEM (Scanning Electron Microscope) micrographs revealed the epiphyte's attachment to the host. Further histological study showed the extent of penetration of epiphytes into the host's cortex tissues and condition of its surrounding tissues. The outbreak of epiphytic filamentous red algae also correlated with drastic changes in seawater temperature and salinity during March– June and September – November.     

DEFORMATIVE DISEASE IN IRIDAEA LAMINARIOIDES (RHODOPHYTA): GALL DEVELOMENT ASSOCIATED WITH AN ENDOPHYTIC CYANOBACTERIUM1

This study is the first report of an algal disease, developed in close association with an endophytic organism, documented for the southeastern Pacific. We describe a disease affecting wild populations of the red alga Iridaea laminarioides Bory in central Chile, characterized by gall development on the surface of sporophytic, cystocarpic, and immature thalli. These abnormal growths result in severe morphological alterations of the affected thalli. Diseased fronds display an aggregated spatial distribution and occur throughout the year, with a maximum in summer followed by a decline in winter. The presence of galls was not associated with broken or torn fronds. Although causality has not been unequivocally demonstrated, our field and laboratory observations indicate a strong association of the galls with infections by an endophytic cyanobacterium, probably belonging to the genus Pleurocapsa.