Observations on Furcellaria fastigiata (L.) Lam. forma aegagropila Reinke in Danish waters together with a note on other unattached algal forms

Summary The unattached populations of Furcellaria fastigiata forma aegagropila in Danish waters exhibit characteristics which are paralleled to a remarkable degree by those possessed by communities of unattached algae in other parts of the world. These characteristics include sterility, multiplication by vegetative means, and the adoption of a form, devoid of holdfast organs, which commonly has a globular or subglobular habit suited to rolling about on the sea bottom. Branching is often profuse and patent and may be accompanied by a thallus diameter far smaller than that of the attached form. Vegetative multiplication may be effected by fragmentation, and, as in F. fastigiata forma aegagropila, by a small accessory lateral branches which function as propagulae. The latter originate in a manner somewhat different from that of the axial regenerated branches produced on decapitated thalli and on thalli which have been more than superficially wounded. Evidence suggests that much of the unattached population of this latter species in the central Kattegat is made up of diploid thalli.All loose lying forms ultimately originate from attached species, which possess or can adopt certain characteristics which enable them to survive in the unattached state.

---

Résumé Les colonies sans attaches de Furcellaria fastigiata forma aegagropila dans les eaux danoises montrent des carat`eres qui sont parallèles à un degré remarquable à ceux possédés par d'autres communautés d'algues flottantes dans les autres parties du monde. Ces caractères comprennent la stérilité, la multiplication par des moyens végétatifs, et l'adoption d'une forme, dépourvue d'organes de préhension, généralement sphérique ou quasi sphérique, adapté à se mouvoir sur le fond de la mer. La formation de branches est souvent abondante et évidente et peut être accompagnée d'un diamètre thallique bien plus petit que celui de la forme fixe. La multiplication végétative peut être effectuée par fragmentation, et comme chez F. fastigiata forma aegagropila, par moyen de petites branches latérales accesoires qui fonctionnent comme des propagulae. Ces dernières prennent leur naissance d'une manière quelque peu différente de celle des branches axiales régénérées produites sur thalles décapités et sur thalles qui ont été plus que superficiellement blessés. Léxpérience suggère que beaucoup de colonies flottantes de cette dernière espèce dans le Kattegat central sont formées de thalles de forme diploide.Toutes les formes libres et flottantes prennent leur origine en dernier ressort des espèces fixes, qui poss`edent ou peuvent adopter certains caractères qui leur permettent de survivre dans l'état de liberté.

---

---

Most of the observational work in this paper was carried out in the Department of Botany, Aberystwith.

---

Personal communications with Herr Fuglede of Skiber.     

The first finding of the unattached form ofLaminaria gurjanovae in Pos’eta Bay, Sea of Japan

The unattached form ofLaminaria gurjanovae A. Zin. inhabiting silty and sandy bottoms at depths of 10–30 m in the area of the Tumen River mouth (Pos’eta Bay) is described for the first time. Laminarian thalli interlace their long (up to 50-cm) stipes and form dense aggretations.     

Regeneration of Enteromorpha

Young plants of E. intestinalis have been grown in culture. Cutting off the thalli of attached plants just above the basal rhizoids leads to the regeneration of new branches. After a short time the point of wounding is indistinguishable. The process can be repeated many times; each regeneration resulting in a more branched thallus. Segments of unattached plants kept in culture may give rise to rhizoids from the basal cut ends and to papillae from the upper ends. They may also produce “bottle brush” forms similar to those found amongst ship-fouling algae. Such forms seem to arise when “swarmers” are retained in the parent cell and germinate in situ to give aggregations of juvenile plants. Their formation can be stimulated by a temperature shock.     

Grazing of attached bacteria by heterotrophic microflagellates

Four species of heterotrophic microflagellates were examined for their ability to graze attached and unattached bacteria. The species tested displayed pronounced differences in their ability to graze the bacteriumPseudomonas halodurans attached to chitin particles. Two species of microflagellates (Monas andCryptobia sp.) efficiently grazed unattached bacteria but showed little or no ability to graze attached or aggregated cells. In contrast,Rhynchomonas nasuta andBodo sp. showed marked preferences for attached and aggregated bacteria and a limited ability to graze unattached cells. The density of attached bacteria was reduced by an order of magnitude due to grazing byBodo andR. nasuta, even though the density of unattached bacteria was 5–90× the density of attached cells. The maximum densities attained by microflagellates in the cultures were related to the density of unattached bacteria forMonas andCryptobia but not forBodo andR. nasuta. Growth of the latter two species appeared to be related to the density of attached or aggregated bacteria. Based on the results of these experiments, it is concluded that the pelagic existence of microflagellates that graze attached bacteria may be strongly linked to the distribution of suspended particles and their associated bacteria. In addition, the removal of attached bacteria by microflagellates can significantly affect the density of bacteria attached to particles in the plankton. This activity may have important implications for the controversy concerning the relative importance of attached and free-living bacteria in the plankton.     

No more lake balls (<Emphasis Type="Italic">Aegagropila linnaei</Emphasis> Kützing,Cladophorophyceae, Chlorophyta) in The Netherlands?

Aegagropila linnaei, a freshwater green macroalga, had been abundant in several locations in The Netherlands before the 1960s. Both the ‘lake ball’ form of this alga and dense unattached mats floating over the sediment have been described from these locations. After 1967, this species has not been recorded anymore from The Netherlands. In 2007, several historical collection sites were surveyed for extant populations of A. linnaei. All habitats have changed drastically during the last 50 years and were affected severely by eutrophication. Populations of A. linnaei seem to have become extinct in all but one location (Boven Wijde, province Overijssel), where we found very small amounts of attached filaments. The attached form had not been reported previously from The Netherlands. Environmental conditions do not seem suitable anymore to maintain extensive unattached growth forms including the enigmatic lake balls, and the species must be regarded as threatened in The Netherlands and we propose to include A. linnaei in a national red list. The decline of populations elsewhere is reviewed and discussed in this paper. In addition to morphological identification of the attached filaments, partial sequences of the nuclear large subunit rDNA were generated and compared with different growth forms and habitats from several other locations outside The Netherlands. The sequences confirm the identity of the Dutch material and indicate very little divergence both between populations in different locations and between different growth forms.     

Distribution and ecological role of the non-native macroalga Gracilaria vermiculophylla in Virginia salt marshes

Intertidal salt marshes are considered harsh habitats where relatively few stress-resistant species survive. Most studies on non-native species in marshes describe terrestrial angiosperms. We document that a non-native marine macroalga, Gracilaria vermiculophylla, is abundant throughout Virginia’s Atlantic coastline. We sampled eight marshes, characterized by low slopes and by the presence of the tube-building polychaete Diopatra cuprea on adjacent mudflats, which have been shown previously to be associated with G. vermiculophylla. G. vermiculophylla was found in 71% of the sampled quadrats on the border between the mudflat and tall Spartina alterniflora, 51% within the tall S. alterniflora zone, and 12% further inland. We also tagged G. vermiculophylla from two habitats: (1) unattached G. vermiculophylla within marshes and (2) G. vermiculophylla ‘incorporated’ onto D. cuprea tubes on the adjacent mudflats. Of the incorporated thalli, 3–9% ended up in the marsh, demonstrating connectivity between habitats. In addition, 21% of unattached thalli remained for 2 weeks within the marsh, suggesting that entanglement around marsh plants reduces tidal drift. Growth experiments in mesh bags indicate that most of the G. vermiculophylla transferred from the lagoon to the marsh decomposed there, potentially enhancing local nutrient levels. Finally, we document that G. vermiculophylla in marshes had a reduced associated flora and fauna compared to G. vermiculophylla on the adjacent Diopatra mudflats. In conclusion, unattached G. vermiculophylla is abundant along marsh borders in the tall S. alterniflora zone in Virginia, and we hypothesize that this non-native species has significant impacts in terms of marsh habitat complexity, species abundance and diversity, nutrient dynamics, productivity, and trophic interactions.     

POPULATION STRUCTURE OF THE EPIPHYTIC LICHENUSNEA LONGISSIMAIN A BOREALPICEA ABIESCANOPY

Usnea longissimawas sampled in SE Norway on six main branches of onePicea abies, comprising its full vertical canopy range. Nearly all specimens (n=781) were unattached and fragmented, in contrast with associated lichens. A combination of weak axes and lack of holdfasts resulted in a population of repeatedly displaced thalli, a similar situation as in ground-dwelling vagant lichens. Comparisons with a litterfall population indicated that thalli, regardless of size, had a similar probability to end as litter on the ground. Specimens were frequently sorediate, but only 12 small thalli with a basal holdfast had been directly recruited through soredial establishment. Its patchy distribution within apparently homogenous forest stands is probably a consequence of a rare successful long-range dispersal through soredia coupled with an abundant local dispersal of coarse thallus fragments. Old forests with ample, diffuse skylight at lower and humid canopy levels are probably essential to maintain a vagant epiphytic life form with a predominantly downward dispersal of thallus fragments.     

Invasion of novel habitats uncouples haplo‐diplontic life cycles

Baker's Law predicts uniparental reproduction will facilitate colonization success in novel habitats. While evidence supports this prediction among colonizing plants and animals, few studies have investigated shifts in reproductive mode in haplo‐diplontic species in which both prolonged haploid and diploid stages separate meiosis and fertilization in time and space. Due to this separation, asexual reproduction can yield the dominance of one of the ploidy stages in colonizing populations. We tested for shifts in ploidy and reproductive mode across native and introduced populations of the red seaweed Gracilaria vermiculophylla. Native populations in the northwest Pacific Ocean were nearly always attached by holdfasts to hard substrata and, as is characteristic of the genus, haploid–diploid ratios were slightly diploid‐biased. In contrast, along North American and European coastlines, introduced populations nearly always floated atop soft‐sediment mudflats and were overwhelmingly dominated by diploid thalli without holdfasts. Introduced populations exhibited population genetic signals consistent with extensive vegetative fragmentation, while native populations did not. Thus, the ecological shift from attached to unattached thalli, ostensibly necessitated by the invasion of soft‐sediment habitats, correlated with shifts from sexual to asexual reproduction and slight to strong diploid bias. We extend Baker's Law by predicting other colonizing haplo‐diplontic species will show similar increases in asexuality that correlate with the dominance of one ploidy stage. Labile mating systems likely facilitate colonization success and subsequent range expansion, but for haplo‐diplontic species, the long‐term eco‐evolutionary impacts will depend on which ploidy stage is lost and the degree to which asexual reproduction is canalized.     

Size-dependence of reproductive allocation of Sargassum thunbergii (Sargassaceae, Phaeophyta) in Bohai Bay, China

Reproductive allocation (the proportion of biomass allocated to reproductive tissue, RA) in Sargassum thunbergii was studied in Yantai, Shandong Peninsula, southern coast of Bohai Bay, China. Annual reproduction initiated in mid-June and peaked in mid-July (90 ± 8% fertile thalli and 75 ± 6% RA mean). Both RA and percentage of fertile thalli exhibited significant temporal variations during reproduction. Sterile thalli were only observed in small length hierarchies at peak reproduction and mean values of RA showed a significantly hierarchical variation, suggesting that the size of thalli played an important role in reproduction and RA were size-dependent. Numerous receptacles were produced along the lateral branches during the reproductive period. A distinct seasonal pattern was observed wherein the presence of lateral branches was followed by the onset of reproduction. RA was positively correlated with the number of lateral branches, as well as the total length of lateral branches. In addition, fertile thalli decayed quickly after peak reproduction. The probability of decay was evidently higher for fertile than for sterile thalli, because all surviving thalli were sterile and short during July–August. So, a trade-off between reproduction and survival may exist at individual levels in S. thunbergii.     

Annual cycle of tintinnids in Admiralty Bay with an emphasis on seasonal variability in Cymatocylis affinis/convallaria lorica morphology

In water samples collected from the middle of Admiralty Bay(King George Island 62°08'S 58°25'W) between February1990 and January 1991, 17 Tintinnina species were noted. TotalTintinnina numbers in summer were very high (up to 5000 cellsm^–3), but species diversity was low, consisting mainlyof Cymatocylis affnislconvallaria, forma convallaria. Duringaustral winter, cell numbers were very low, but species compositionwas diverse. Cymatocylis affnislconvallaria, forma affinis,and C.affmis/convallaria, forma convallaria, the polymorphicforms of one species C.affinis/convallaria, appear to be interchangeableduring the year. The transition from one form to the other occursin the spring and autumn. The typical Tintinnina polymorphismcan be attributed to the prevailing environmental conditions.     

QUANTITATIVE VARIATION IN VULPINIC AND PINASTRIC ACIDS PRODUCED BY TUCKERMANNOPSIS PINASTRI (LICHEN-FORMING ASCOMYCOTINA,PARMELIACEAE)

Tuckermannopsis pinastri is a lichen species found commonly on rocks and tree branches in boreal and alpine habitats in the northern United States. Members of this species produce three yellow-pigmented phenolic compounds: usnic, pinastric, and vulpinic acids. The objective of our study was to quantify the variation in concentration of the latter two of these compounds in relation to substrate factors, chlorophyll content, and thallus size. Using high-performance liquid chromatography, we analyzed 120 thalli of T. pinastri collected randomly from a single large population located at Spruce Knob, West Virginia. Although individuals were sampled from both tree and rock substrates that differed markedly in light intensity, these environmental factors were not correlated with observed variations in vulpinic or pinastric acid concentrations. Instead, compound concentrations were correlated most closely with thallus size, with small rather than large thalli having the highest concentrations of the two compounds. Small thalli did not have higher concentrations of chlorophylls than large thalli, however, which suggests that the rate of production of secondary compounds by the fungus in T. pinastri is independent of algal biomass. Inasmuch as lichen secondary compounds serve a defensive role against microorganisms and herbivores, our results suggest that small, juvenile thalli are better defended than more mature thalli.     

The attached form of the endangered freshwater alga Aegagropila linnaei Kützing (Chlorophyta) is found in the Zuideindigerwiede, The Netherlands

The freshwater green macroalga Aegagropila linnaei used to occur in several locations in The Netherlands in unattached growths forms (including the enigmatic ‘lake balls’) more than 60 years ago. A recent survey of most of the historical collection sites of the species reported the attached growth form for the first time for The Netherlands, but only in miniscule quantities and from one location only. Here we report on the finding of the attached form from the Zuideindigerwiede, one of the two historical collection sites that had not been surveyed previously. Unattached growth forms do not seem to occur anymore in The Netherlands, and the species as a whole must be regarded as rare and endangered.     

The life history of Porphyra endiviifolium from the South Shetland Islands, Antarctica

This paper describes the reproduction and life history of an intertidal species, Porphyra endiviifolium, from Antarctica. Field specimens were examined microscopically, prepared for electron microscopy and used to establish cultures. Wild populations comprised two kinds of leafy thalli, morphologically similar but distinguished by their mode of reproduction, either sexual or asexual. Carpospores from monoecious leafy gametophytes developed into conchocelis filaments in culture, and under “winter-spring” conditions these formed conchospores that germinated to produce leafy thalli. Monospores from asexual leafy thalli developed directly into two different forms of leafy thalli. Only one of the cultured morphotypes became fertile, reproducing asexually by monospores. We conclude that the phases of the life history of P. endiviifolium show different ecological strategies, the conchocelis phase reproducing in response to short days unlike the leafy thalli in which growth and reproduction respond primarily to irradiance. Received: 2 May 1997 / Accepted: 11 October 1997     

The role of macroalgae in epiphytism of the toxic dinoflagellate Gambierdiscus toxicus (Dinophyceae)

Populations of the toxic, epiphytic dinoflagellate Gambierdiscus toxicus Adachi et Fukuyo are asSociated closely with Jania sp. on Hitiaa and Papara fringing reefs in Tahiti. Small populations were also observed to be asSociated with Amphiroa sp. and Halimeda opuntia (L.) Lamouroux. The cells attached themselves to the thallus by means of a short thread. When the thalli were irradiated, the cells began to detach from them and swim around the branches. The swimming cells stopped and attached to substrata when a disturbance occurred. The attached cells began to swim within a short time under light conditions when the thallus of Jania sp. were placed near the attached cells. Amphiroa sp. and H. opuntia also induced this re-commencement of swimming of the attached cells. These observations suggest that G. toxicus usually swims around macroalgal thalli on coral reefs. When sudden disturbance or strong water motion occurs, they attach to the surface of macroalgae and are not dispersed. Soon after water motion becomes slow, the cells begin to swim into the water around the thalli. The epiphytism of G. toxicus is different from epiphytic pennate diatoms, most of which adhere to the thallus all the time. The population of G. toxicus is maintained as an asSociation to a limited number of species of macroalgae which support the re-commencement of swimming after disturbance.     

The effects of egg position, egg mass size, substrate and biofouling on embryo mortality in the squid Sepioteuthis australis

Using a combination of laboratory and field investigations, this study examined embryo mortality in the southern calamary Sepioteuthis australis as a function of egg mass size, the substrate upon which the mass is attached, the position of the embryo within the mass, and the degree of biofouling. Egg mass size ranged from 2 to 1,241 egg strands, however most masses consisted of 200–299 strands. Small egg masses (<300 strands) were generally attached to soft-sediment vegetation (Amphibolis antarctica, Heterozostera tasmanica, Caulerpa sp.), whereas larger masses (>300 strands) were either securely attached to robust macroalgae holdfasts (Ecklonia sp., Marcocystis pyrifera, Sargassum sp.) or unattached. Rates of embryo mortality were highly variable ranging from 2 to 25%. Both laboratory and field results indicated a positive relationship between egg mass size and embryo mortality. Larger, unattached egg masses contained twice as many dead embryos than those securely attached to a substrate. Mortality rates were significantly affected by the embryos’ relative position within the mass and were highest in embryos located near the attachment point of the egg strand, within the interior of the mass, and in close contact with the substrate. This was attributed to the inability of the embryos to respire adequately and eliminate metabolic wastes. Biofouling did not strongly influence embryo mortality, but colonisation occurred in areas conducive to growth, photosynthesis, and respiration indicating ‘healthy’ regions within the mass.     

‘Seed’ production of Porphyra spp. by tissue culture

Differentiation of archeospores was observed from excised tissue of young thalli of various monoecious Porphyra species ( P. tenera, P. yezoensis, P. suborbiculata, P. okamurae) after 4–8 days in culture at temperatures of 20 and 25 °C. Excised tissue from adult thalli did not differentiate into archeospores, but rather regenerated directly into blades and rhizoids of foliose thalli. Tissues from young thalli of two dioecious Porphyra species ( P. dentata and P. pseudolinearis) also regenerated into blades and rhizoids after manipulation of the culture conditions. In addition, 1–2 celled tissue pieces of both monoecious and dioecious species were also seen to develop directly into blades. Polarity of regeneration of blades and rhizoids was observed in these species. These results suggest that ‘seed’ can be obtained through tissue culture instead of using conventional conchocelis culture for commercial nori aquaculture in Japan. This revised version was published online in June 2006 with corrections to the Cover Date.     

INTERPOPULATIONAL VARIATION IN THE GAMETOPHYTES OF PELLAEA ANDROMEDAEFOLIA

The development and mature morphology of the gametophytes from both sexual and apogamous populations of the fern Pellaea andromedaefolia were investigated. While most sexual examples were indistinguishable, some differences were noted. An insular collection was distinctive in its variability and irregularity of form. Although the latter was a representative of var. pubescens, other collections of the variety could not be distinguished from var. andromedaefolia on the basis of gametophytic characteristics. The apogamous gametophytes were decidedly more variable in development and often very different from sexual thalli. The mature asexual thalli tended to be more irregular in form and usually sharply divergent from the typical cordate type characteristic of the sexual populations. Each of the five apogamous samples was unique with respect to gametophyte development. The differences among the gametophytes of the various populations do not correlate with the sporophytic characteristics which differentiate the two varieties of the 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.     

Crossing test among floating Ulva thalli forming `green tide' in Japan

Crossing tests were made to determine the relationship between the identified Ulva pertusa, which commonly grows in Japan as an attached form on exposed rocks, and the floating Ulva forming "green tide" inside calm bays. The floating Ulva thalli were collected from five major green tide sites in Japan (Yokohama, Mikawa, Miyajima, Kochi and Hakata). Reproductive maturation was induced in U. pertusa and the floating thalli from each site. Mating between induced gametes was observed. It is therefore believed that the floating thalli from Yokohama, Mikawa and Miyajima were mainly U. pertusa, while those from Kochi and Hakata were of a different species (Ulva sp.1). Furthermore, the Ulva species found in Mikawa is also a species (Ulva sp.2) different from both U. pertusa and Ulva sp.1.