Resource space partitioning by the Bryozoa of a Fucus serratus L. Community

The spatial distributions of five Bryozoa species encrusting Fucus serratus L. plants in Strangford Lough, Northern Ireland, have been examined in two cross-sectional samples yielding 80 plants and 1326 sampling units drawn from 13 environmentally diverse sites. The co-occurences of Flustrellidra hispida (Fabricius), Alcyonidium hirsutum (Fleming), Electra pilosa (L.), Membranipora membranacea (L.), and Celleporella hyalina (L.) on individual Fucus serratus plants were consistent with independent settlement onto each plant, but their joint distributions within the plants were markedly non-random. Measurements of behavioural interactions between species showed that competitive abilities declined through the sequence Flustrellidra, Alcyonidium, Electra, Membranipora, which parallels the sequence of competition (Levins' α) coefficients calculated from spatial distribution data. Competitive ability varied with position on the plant. Flustrellidra and Alcyonidium reversing their relative competitive superiority between basal and distal regions of the fronds. Flustrellidra and Electra were strongly associated under all conditions. Alcyonidium was also frequently present with this pair, though its numerical abundance was inversely related to that of Flustrellidra. Celleporella tended to co-occur with Electra whilst avoiding Flustrellidra and Alcyonidium. Membranipora numbers were inversely related to those of the other species. Analyses of competition coefficients showed that Flustrellidra, Alcyonidium and Electra each peaked in competitive ability against Membranipora at different points along the frond, such that the latterwaas effectively excluded by direct as well as by diffuse competition. Membranipora populations had a greater dependence on the segment area available for colonization than had any other bryozoan species.     

Predator-induced polymorphism in the bryozoan Membranipora membranacea (L.)

Three described species of the bryozoan genus Membranipora, M. membranacea (L.), M. serrilamella Osburn, and M. villosa Hincks, occurring together on brown macroalgae off the west coast of North America are eco-phenotypic variants. These forms should be called M. membranacea on the basis of priority.The presence of spines, an important diagnostic feature separating these Membranipora forms, is induced by two nudibranch predators, Corambe pacifica MacFarland & O'Donoghue and Doridella steinbergae Lance. Spines protect Membranipora zooids against predation by these nudibranchs. The population costs of spine production include lower growth and reproductive rates.     

Invasive species as a new food source: does a nudibranch predator prefer eating an invasive bryozoan?

Membranipora membranacea is an invasive bryozoan that was first found in the Gulf of Maine in 1987 and within two years became the dominant organism living on kelps. Membranipora may have become dominant so quickly because it had little competition in a relatively unoccupied niche; however, lack of predation has also probably played a major role. Where Membranipora is native, there is usually a specialist nudibranch predator that keeps the population in check. For example, in European populations, the nudibranch Polycera quadrilineata prefers Membranipora while Onchidoris muricata is known to prefer another bryozoan, Electra pilosa. Electra, Membranipora, and Onchidoris are all now found in the Gulf of Maine while Polycera is not. We tested whether Onchidoris would (1) eat Membranipora at all, (2) eat Membranipora and Electra at different rates, and (3) show a preference for eating Membranipora or Electra when given a choice. We found that Onchidoris does eat Membranipora, and it generally eats Membranipora faster than Electra. However, when given a choice, Onchidoris prefers Electra. Onchidoris typically reproduces in the spring and grows over the fall and winter, but has recently been found reproducing in the winter in New Hampshire. Although it does not survive the winter as well as Electra, Membranipora is the dominant organism living on many macroalgae in the late summer and fall. Thus, the large Membranipora food source now available in the summer and fall may allow Onchidoris to reproduce earlier.     

Development,metamorphosis, and natural history of the nudibranch Doridella obscura Verrill (Corambidae: Opisthobranchia)

The doridacean nudibranch Doridella obscura Verrill was raised through one complete generation in laboratory culture, and spawning behavior monitored for a year at monthly intervals in Barnegat Bay, New Jersey.The nudibranch deposited egg masses throughout the year in Barnegat Bay, and the larvae remained viable at temperatures ranging from 1.5 to 28 °C. At 25 °C the eggs hatch 4 days after oviposition, and the planktotrophic veliger larvae swim and feed for 9 days before they metamorphose. Settlement occurs specifically on the bryozoan Electro crustulenta (Pallas). The spirally coiled larval shell grows rapidly until the dorsal mantle fold is retracted from the aperture 5–6 days after hatching. Although starved larvae grow only slightly and do not metamorphose, they resume normal development on introduction of suitable food. Newly metamorphosed juveniles consume algae and debris on the surface of the bryozoan until they grow large enough to attack the living zooids of E. crustulenta.The life cycle of Doridella obscura is short (26 days at 25 °C), allowing the nudibranchs to take advantage of short-lived Electra crustulenta colonies in unstable habitats in bays and estuaries.     

The effects of infection by Tetracapsuloides bryosalmonae (Myxozoa) and temperature on Fredericella sultana (Bryozoa)

The myxozoan, Tetracapsuloides bryosalmonae, exploits freshwater bryozoans as definitive hosts, occurring as cryptic stages in bryozoan colonies during covert infections and as spore-forming sacs during overt infections. Spores released from sacs are infective to salmonid fish, causing the devastating Proliferative Kidney Disease (PKD). We undertook laboratory studies using mesocosm systems running at 10, 14 and 20 °C to determine how infection by T. bryosalmonae and water temperature influence fitness of one of its most important bryozoan hosts, Fredericella sultana, over a period of 4 weeks. The effects of infection were context-dependent and often undetectable. Covert infections appear to pose very low energetic costs. Thus, we found that growth of covertly infected F. sultana colonies was similar to that of uninfected colonies regardless of temperature, as was the propensity to produce dormant resting stages (statoblasts). Production of statoblasts, however, was associated with decreased growth. Overt infections imposed greater effects on correlates of host fitness by: (i) reducing growth rates at the two higher temperatures; (ii) increasing mortality rates at the highest temperature; (iii) inhibiting statoblast production. Our results indicate that parasitism should have a relatively small effect on host fitness in the field as the negative effects of infection were mainly expressed in environmentally extreme conditions (20 °C for 4 weeks). The generally low virulence of T. bryosalmonae is similar to that recently demonstrated for another myxozoan endoparasite of freshwater bryozoans. The unique opportunity for extensive vertical transmission in these colonial invertebrate hosts couples the reproductive interests of host and parasite and may well give rise to the low virulence that characterises these systems. Our study implies that climate change can be expected to exacerbate PKD outbreaks and increase the geographic range of PKD as a result of the combined responses of T. bryosalmonae and its bryozoan hosts to higher temperatures.     

EFFECTS OF THE ENCRUSTING BRYOZOAN,MEMBRANIPORA MEMBRANACEA,ON THE LOSS OF BLADES AND FRONDS BY THE GIANT KELP,MACROCYSTIS PYRIFERA (LAMINARIALES)1

Juvenile sporophytes of the giant kelp, Macrocystis pyrifera (L.) C. A. Agardh, were transplanted from local kelp beds to stations located various distances from the outfall from an electrical generating station that was known to cause an increase in the settlement of fouling organisms. Plants near the outfall became heavily fouled by the encrusting bryozoan, Membranipora membranacea (L.), and lost about one-third of their blades during the course of the experiment. Blade loss was significantly correlated with amount of fouling. To test the hypothesis that fouling causes blade loss, we paired fouled and unfouled plants of about the same age, overall length, and number of fronds and placed them at stations in nearby kelp beds and near the outfall. At the stations in the kelp beds, the fouled plants lost blades more rapidly than the unfouled controls. However, at the station near the outfall the “control” plants quickly became fouled so there was little difference in treatments and there was no significant difference in blade loss. Plants fouled by Membranipora suffered greater blade loss than clean plants probably because fouled blades are fragile and break off easily and because fish bite off chunks of blade while foraging on the attached bryozoans.     

Orientated growth of Membranipora membranacea (L.) on the thallus of Saccorhiza polyschides (Lightf.) Batt.

Orientated growth is described in colonies of Membranipora membranacea (L.) growing on the large brown seaweed Saccorhiza polyschides (Lightf.) Batt. On average, 60% of the growth of the colonies was directed proximally along the long axis of the plant, irrespective of the region of the plant on which the colonies were growing or their proximity to its primary meristem. These results seem to rule out the possibility that orientated growth in Membranipora is a response to an age dependent polarity in the host plant, since the tissue ages in opposite directions in the lamina and stipe yet the growth of Membranipora is orientated in the same direction on both. The possibility that directional growth in this species may be a rheopositive response is discussed.     

The physiological ecology of growth and reproduction in Onchidoris aspera (Alder & Hancock) (Gastropoda:Nudibranchia)

The growth and reproduction of the dorid nudibranch Onchidoris aspera (Alder & Hancock 1842) was examined at East Point, Nahant, Massachusetts (September 1981 to April 1982) and field observations were made over 4yr (June 1978 to June 1982). Juvenile recruitment into the shallow subtidal (6–10 m) occurred in early October. Maximum specific growth rates (biomass) of 0.473 ± 0.759 mg·mg^?1·month^?1 were attained during the period of October through December. Experiments with pairs of individuals showed that reproductive effort could be high, amounting to 41–108% of nonrespired assimilation, over 10 wk of spawning (28 December to 12 March). Adults and juveniles are temporally separated at this site; parental populations persist until approximately June, and juvenile recruits are not observed until the following October. Occurrence of this species at this site has an upper thermal limit of ≈15°C; the temperature at which spawning takes place lies between 0–4°C. Feeding on bryozoan prey is suctorial by means of a well developed buccal pump; observations and experiments showed that the most common encrusting bryozoan, Electra pilosa (L.) was the preferred prey and that upon which maximum growth rates were achieved in Onchidoris aspera.     

Davallia (Polypodiales: Davalliaceae) macrofossils from Early Miocene Otago (New Zealand) with in situ spores

Fossil fern fronds referable to the extant fern genus Davallia (Polypodiales: Davalliaceae) bearing sporangia with in situ spores are described from the Early Miocene Foulden Maar diatomite deposit, Otago, New Zealand. The fronds are the first published Southern Hemisphere macrofossil record for the family and provide valuable palaeoclimate data supporting warm conditions in Early Miocene New Zealand. The matching of Davallia fronds to the form spore taxon Polypodiisporites radiatus shows that the genus has had a long, apparently continuous history throughout late Cenozoic New Zealand.     

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.     

Effects of wave exposure on population and reproductive phenology of an algal turf, Gelidium pusillum (Gelidales,Rhodophyta), Songkhla,Thailand

The effect of wave exposure on the population and reproductive phenology of the common red alga, Gelidium pusillum (Stackhouse) Le Jolis, was investigated between July 2003 and June 2004, at Suan Song Tha Le, Songkla Province, Thailand. Lengths of thalli, percentage cover, percentage of reproductive fronds and the number of reproductive structures were examined monthly in relation to different degrees of wave exposure (sheltered vs. exposed), temperature, rainfall, day length and tidal cycles. Frond length and percentage cover of G. pusillum were different among sites and seasons. Shorter fronds were found on the exposed shore which had a greater percentage cover. Fronds bleached and died off during the summer months (April and May), which resulted in shorter fronds and reduced percentage cover in June. The thalli reproduced throughout the year, with a predominance of tetrasporophytes. The highest percentage of tetrasporic fronds was 33% in February 2004 and 13% of cystocarpic fronds in April 2004, but male gametophytic fronds were never observed. Rainfall showed a strongly negative influence on reproduction since no reproductive fronds were observed during the rainy season (R² = 0.49, P = 0.01). The dominance of G. pusillum at this study site and throughout elsewhere in turf habitats might be a function of persistent vegetative growth, densely clumped, and the ability to reproduce almost throughout the entire year.     

Vigna unguiculata is nodulated in Spain by endosymbionts of Genisteae legumes and by a new symbiovar (vignae) of the genus Bradyrhizobium

Vigna unguiculata was introduced into Europe from its distribution centre in Africa, and it is currently being cultivated in Mediterranean regions with adequate edapho-climatic conditions where the slow growing rhizobia nodulating this legume have not yet been studied. Previous studies based on rrs gene and ITS region analyses have shown that Bradyrhizobium yuanmingense and B. elkanii nodulated V. unguiculata in Africa, but these two species were not found in this study. Using the same phylogenetic markers it was shown that V. unguiculata, a legume from the tribe Phaseolae, was nodulated in Spain by two species of group I, B. cytisi and B. canariense, which are common endosymbionts of Genisteae in both Europe and Africa. These species have not been found to date in V. unguiculata nodules in its African distribution centres. All strains from Bradyrhizobium group I isolated in Spain belonged to the symbiovar genistearum, which is found at present only in Genisteae legumes in both Africa and Europe. V. unguiculata was also nodulated in Spain by a strain from Bradyrhizobium group II that belonged to a novel symbiovar (vignae). Some African V. unguiculata-nodulating strains also belonged to this proposed new symbiovar.     

The importance of phenotypic differentiation and plasticity in the range expansion of the annual Atriplex tatarica (Amaranthaceae)

The ability of a species to adapt to sub-optimal conditions at the margin of its distribution range and to cope with environmental stress is considered to be important for its successful geographic expansion. To ascertain the roles of phenotypic differentiation and plasticity in the expansion of the annual Atriplex tatarica, we compared plants from populations found in Marginal and Central areas of the species’ range. We grew these plants under marginal climatic conditions in pots with different types of substrate. We assessed the population genetic structure at five putatively neutral allozyme loci to evaluate whether there was any evidence of reduced genetic diversity in Marginal populations compared to Central ones. We used the Q_ST vs. F_ST approach (while F_ST gives a standardised measure of the genetic differentiation among populations for a genetic locus, Q_ST measures the amount of genetic variance among populations relative to the total genetic variance) to ascertain the roles of adaptive vs. non-adaptive processes on phenotypic differentiation. Plants native to the Marginal area of the species’ range flowered earlier and had a lower shoot mass and a higher reproductive allocation than plants native to the Central part of the species’ range. The Marginal populations of Atriplex tatarica showed lower genetic diversity at allozyme loci and higher phenotypic differentiation than the Central populations. We recorded similar plastic responses to substrates in plants native to both regions. Our results indicate that Marginal populations of expanding A. tatarica maintain the ability to adapt locally and to elicit a plastic response to environmental stress, despite loss of genetic diversity.     

Morphology and growth habit of Dicksonites pluckenetii from the Upper Carboniferous of Graissessac (France)

Morphological variability of the vegetative fronds of Dicksonites pluckenetii (SCHLOTHEIM ex STERNBERG) STERZEL is described from new material collected in the Stephanian Basin of Graissessac (Southern Massif Central France). Complete bipartite fronds sometimes still attached to their parent stem allow us to reconstruct the plant as a scrambler/climber. This interpretation is supported by the relatively small slender stems with long internodes and densely covered by short spines, which are also present along the entire length of petioles. A fertile pinna with slightly reduced pinnules bearing up to six radially symmetrical synangia is interpreted as a portion of male frond of the same species; its organization confirms the relationships of Dicksonites with the Callistophytales as was already suggested by features of the female fronds.     

A SHORT-TERM DEMOGRAPHIC STUDY OF CYSTOSEIRA OSMUNDACEA (FUCALES: CYSTOSEIRACEAE) IN CENTRAL CALIFORNIA1

Several demographic features were examined in the field over a period of eleven months for the subtidal alga Cystoseira osmundacea (Turner) C. Agardh at two localities near Monterey, California. This species showed a restricted distribution with depth, with peak average numbers of 3–4 plants per m² occurring at 6–8 m and forming a dense canopy on the surface of the sea. Below 10 m depth, there was a decline in the abundance of plants and also in the proportion of plants which bore seasonal fronds and reproductive tissue. Macrocystis pyrifera (L.) C. Agardh plants were also common at both localities. There was a negative correlation between the presence of this species and C. osmundacea at a scale of 1 m², but no correlation at 4, 9, and 25 m². Cohorts of C. osmundacea were mapped and tagged at one locality. The plants had markedly seasonal growth, with the greatest elongation of fronds occurring between March and June. Dense surface canopies and reproductive tissue were present during the summer months of June–August. These seasonal fronds were shed during September, leaving small perennial holdfasts and fronds. Plants were strictly dioecious, and there was a one-to-one ratio of male to female plants at both localities. Length-dry weight relationships showed that the largest plants had over 80% of their weight invested in seasonal fronds (vegetative plus reproductive tissue). Recruits of C. osmundacea at one locality appeared during September–November only in areas cleared to bare substratum. There was 15% survival of recruits between November and the following May, while 98% of the original mapped cohort survived for the 11 months of the study.     

Effects of the bryozoan Membranipora tuberculata (Bosc.) on the photosynthesis and growth of Gelidium rex Santelices et Abbott

Several species of Membranipora commonly occur on brown and red algae. It is currently believed that these bryozoans could affect growth of the host plant by modifying incident light. In the present study, we report on the changes that occur in both intensity and quality of light passing through colonies of M. tuberculata (Bosc.) and on the effects of bryozoans on growth of the host plant. M. tuberculata decreases photon-flux density by up to 56% of the incident light and slightly affects the quality of the incident light. Although the bryozoan reduces the photosynthetic rate of Gelidium rex Santelices et Abbott, it does not have a net effect on algal growth. It is concluded that the effects of epifauna on algal growth depend on the nature of the algal structure colonized. Therefore, it cannot be predicted a priori that encrusted thalli should grow at a slower rate than nonencrusted thalli.     

Endogenous isoflavone methylation correlates with the in vitro rooting phases of Spartium junceum L. (Leguminosae)

Spartium junceum L. (Leguminosae) is a perennial shrub, native to the Mediterranean region in southern Europe, widespread in all the Italian regions and, as a leguminous species, it has a high isoflavone content. An in vitro culture protocol was developed for this species starting from stem nodal sections of in vivo plants, and isoflavone components of the in vitro cultured tissues were studied by means of High Performance Liquid Chromatography (HPLC) analytical techniques. Two main isoflavones were detected in the S. junceum tissues during the in vitro propagation phases: Genistein (4′,5,7-Trihydroxyisoflavone), already reported in this species, and its methylated form 4′,5,7-Trimethoxyisoflavone, detected for the first time in this plant species (0.750 ± 0.02 mg g⁻¹ dry tissue). The presence of both of these compounds in S. junceum tissues was consistently detected during the in vitro multiplication phase. The absence of the methylated form within plant tissues in the early phases of the in vitro adventitious root formation was correlated with its negative effect displayed on root induction and initiation phases, while its presence in the final “root manifestation” phase influenced positively the rooting process. The unmethylated form, although detectable in tissues in the precocious rooting phases, was no longer present in the final rooting phase. Its effect on rooting, however, proved always to be beneficial.     

Translocation of C in Macrocystis pyrifera (Giant Kelp)

The pattern of import and export of ¹⁴C-labeled assimilates in Macrocystis pyrifera (L.) C. A. Agardh in southern California was studied by labeling single blades on fronds, in situ, with [¹⁴C]NaHCO₃ for 24 hours. The pattern was found to be similar to that known in dicotyledons: actively growing tissue imported and did not export. As a blade reached maturity it began to export, at first only acropetally to the apex which formed it, later also down the frond to sporophylls and frond initials at the base of the frond, and into the apical regions of juvenile fronds; finally there was a phase of declining export, late in the life of the blade, when transport was only downward. Young fronds imported from older fronds until they were approximately 3 meters long, by which time they had developed mature, upward exporting blades. No translocation was found from a younger frond to an older frond, nor was there transport upward from a blade on a frond lacking the apical region.     

The demography of fronds of Chondrus crispus Stackhouse

A demographic analysis was done of an intertidal stand of Chondrus crispus Stackhouse at Pubnico Point, Nova Scotia. The distribution of gametophytic and tetrasporophytic fronds was assessed. Fronds were placed in one of four size-classes. Size-specific measurements for fronds are presented. Gametophytic fronds dominated the stand. Fronds in Size-Class 2 exhibited high survivorship. Survivorship fell sharply when fronds entered Size-Class 3 and Size-Class 4. Fronds became fertile and highly branched when in the third and fourth size-classes. It is suggested that these two factors contribute to the higher mortality rates of such individuals.Monthly spore release rates·m^?2 for gametophytes and tetrasporophytes were measured. The annual release rate of carpospores exceeded the annual release rate of tetraspores by a ratio of 4.7:1. Rates did not fall below 10⁶ spores released·m^?2·month^?1 for either phase.     

Effect of abscisic acid on photosynthetic products of Lemna minor

Lemna minor fronds were grown on nutrient only, or nutrient plus 10^?6M abscisic acid (ABA) for 2 or 8 days. After various ¹⁴CO₂ pulse-chase time periods, the fronds were harvested and the photosynthetic products separated into acidic, lipid, residue, sugar and amino acid fractions. Compared with the control fronds, total ¹⁴C-fixation was 15% higher in the 2 day ABA-treatment and 6% lower in the 8 day ABA-treatment. This pattern was reflected in all the fractions examined, and it appeared that ABA did not alter the distribution of ¹⁴C between the photosynthetic products during the ¹⁴CO₂ pulse. During the chase, less ¹⁴C was lost from the carbohydrate fractions in the ABA-treated fronds than in the control fronds. The results indicate that the previously reported ABA-mediated increase in carbohydrate levels was a consequence of decreased degradation rather than an increase in synthesis from assimilated carbon.