Long-term population dynamics of Littorina obtusata: the spatial structure and impact of trematodes

There are few examples of host population regulation by macroparasites in stable communities; however, strong impact of parasites on the host individual is obvious in many cases (for example increased mortality, a reduction in fecundity up to the complete castration). Associations between the host populations (periwinkle Littorina obtusata) and prevalence of trematodes was investigated using long-term data (1982–1997) of two L. obtusata populations in the White Sea, northwest Russia. We hypothesized that high prevalence of trematodes will reduce future host population density, and increase mortality. Using a general linear model, we found a significant negative correlation between host population density and the prevalence of the most abundant parasite, Microphallus piriformes in the previous year. We found no correlation between snail reproduction and the prevalence, but observed a significant reduction in middle-aged mollusk abundance which was associated with high prevalence. This indicates the importance of parasite-induced mortality for the dynamics of the host population. There was an association between trematode infection and L. obtusata populations that influence their distribution within littoral zone. The ‘source’ population, located in the lower section of the macrophyte zone, appears to be self-sustaining, controlling the whole population recruitment and dynamics.     

The habitat and feeding preferences of Littorina obtusata (L.) and L. mariae sacchi et rastelli

The vertical zonation of the flat periwinkles Littorina obtusata (L.) and L. mariae Sacchi et Rastelli is different, but overlaps. At a sheltered location on the Isle of Cumbrae, Scotland, L. obtusata, although most abundant on Fucus vesiculosus L. and Ascophyllum nodosum (L.) Le Jol. on the midshore, ranged over the entire shore from the Pelvetia zone down to the upper limits of Laminaria. Littorina mariae exhibited a much more restricted distribution being associated predominantly with Fucus serratus L. on the lower shore. In laboratory trials with adult plants, flat periwinkles found fucoid algae both more attractive and more palatable than other seaweeds. Littorina obtusata also preferred the reproductive receptacles of fucoid algae to their vegetative tissue. Tiny germlings of a diversity of seaweeds were readily consumed by L. obtusata. In choice experiments germling Ulva lactuca L. was preferred to adult U. lactuca, whereas adult Fucus serratus was preferred to germling F. serratus. Contrasting foraging behaviours between different species of flat periwinkle are discussed in relation to differences in their radular dentition.     

Bottlenecks to coral recovery in the Seychelles

Processes that affect recovery of coral assemblages require investigation because coral reefs are experiencing a diverse array of more frequent disturbances. Potential bottlenecks to coral recovery include limited larval supply, low rates of settlement, and high mortality of new recruits or juvenile corals. We investigated spatial variation in local abundance of scleractinian corals in the Seychelles at three distinct life history stages (recruits, juveniles, and adults) on reefs with differing benthic conditions. Following widespread coral loss due to the 1998 bleaching event, some reefs are recovering (i.e., relatively high scleractinian coral cover: ‘coral-dominated’), some reefs have low cover of living macrobenthos and unconsolidated rubble substrates (‘rubble-dominated’), and some reefs have high cover of macroalgae (‘macroalgal-dominated’). Rates of coral recruitment to artificial settlement tiles were similar across all reef conditions, suggesting that larval supply does not explain differential coral recovery across the three reef types. However, acroporid recruits were absent on macroalgal-dominated reefs (0.0 ± 0.0 recruits tile^?1) in comparison to coral-dominated reefs (5.2 ± 1.6 recruits tile^?1). Juvenile coral colony density was significantly lower on macroalgal-dominated reefs (2.4 ± 1.1 colonies m^?2), compared to coral-dominated reefs (16.8 ± 2.4 m^?2) and rubble-dominated reefs (33.1 ± 7.3 m^?2), suggesting that macroalgal-dominated reefs have either a bottleneck to successful settlement on the natural substrates or a high post-settlement mortality bottleneck. Rubble-dominated reefs had very low cover of adult corals (10.0 ± 1.7 %) compared to coral-dominated reefs (33.4 ± 3.6 %) despite no statistical difference in their juvenile coral densities. A bottleneck caused by low juvenile colony survivorship on unconsolidated rubble-dominated reefs is possible, or alternatively, recruitment to rubble-dominated reefs has only recently begun. This study identified bottlenecks to recovery of coral assemblages that varied depending on post-disturbance habitat condition.     

Age-dependent zonation of the periwinkle Littorina littorea (L.) in the Wadden Sea

On sedimentary tidal flats near the island of Sylt (German Bight, North Sea) abundance and size distribution of periwinkles, Littorina littorea L., were studied in low intertidal and in shallow and deep subtidal mussel beds (Mytilus edulis L.). In low intertidal mussel beds, surveys revealed that high densities (1,369±571 m^–2) of juvenile snails (≤13 mm) were positively correlated with strong barnacle epigrowth (Semibalanus balanoides L. and Balanus crenatus Bruguière) on mussels. A subsequent field experiment showed that recruitment of L. littorea was restricted to the intertidal zone. Abundances of periwinkles (213±114 m^–2) and barnacles abruptly decreased in the adjacent shallow subtidal zone, which served as a habitat for older snails (>13 mm). L. littorea was completely absent from disjunct deep (5 m) subtidal mussel beds. Snail abundance varied seasonally with maxima of >4,000 m^–2 in low intertidal mussel beds in October and minima in July, just before the onset of new recruitment. I suggest that the presence of cracks and crevices among the dense barnacle overgrowth in intertidal mussel beds favoured recruitment and survival of juvenile snails. Larger (older) specimens are assumed to actively migrate to the less favourable adjacent subtidal. Therefore, intertidal mussel beds are considered as nurseries for the population of L. littorea in the Wadden Sea. Received in revised form: 25 September 2000 Electronic Publication     

Rates of spontaneous hybridization and hybrid recruitment in co-existing exotic and native mature larch populations

Exotic larch (Larix decidua Mill., Larix kaempferi (Lamb.) Carr. and Larix sibirica Ledeb.) and their hybrids have been introduced into Canada to test adaptation and growth for reforestation purposes. For tree breeding operations, provenance trials and seed orchards of exotic larches and their hybrids have been established in arboreta adjacent to natural forest stands of the indigenous species Larix laricina (Du Roi) K. Koch. This proximity offers an opportunity to measure spontaneous hybridization and the recruitment of hybrids. Using a combination of two chloroplast and three mitochondrial DNA markers, we estimated the rate of spontaneous hybridization taking place between native (L. laricina) and exotic (L. decidua, L. kaempferi, L. sibirica) species of larch in two arboreta. More than 5,000 seeds and 1,000 recruits from the two experimental sites were sampled and genetically identified. The occurrence of spontaneous hybridization between L. laricina and the exotics was found both in seeds and from recruits freely established near the plantations. A low hybridization rate (3 % or less) was observed among the seeds of L. laricina maternal trees. Spontaneous native-exotic hybrids were also observed to establish and survive in the environment under the current climatic conditions at a similarly low rate. However, a much higher and variable rate of establishment was observed for recruits with exotic components (up to 34.6 % at one site). These results indicate that the spontaneous hybridization and the recruitment of hybrid and exotic recruits do occur and should be taken into consideration in the management and establishment of plantations of fast-growing species with exotic components.     

Inheritance of the background shell color in the snails Littorina obtusata (Gastropoda,Littorinidae)

We investigated in a gastropod mollusk Littorina obtusata (L. obtusata) the inheritance of back-ground shell coloration of the shell, which arises on the basis of three pigments: purple, orange, and yellow. We found that the hypothesis on polyallelic inheritance, as in the genus Cepaea, cannot explain the inheritance of shell color in periwinkles. We demonstrated that a separate genetic system is responsible for incorporation of each pigment into the shell. The composition of these genetic systems includes at least two genes each in the case of the yellow and purple pigments. Our analysis shows that caution should be applied when extending the results obtained in the studies of the Cepaea genus to the other species of gastropods.     

Resistance to freshwater exposure in White Sea Littorina spp. I: Anaerobic metabolism and energetics

 Anaerobic metabolism and changes in the osmotic concentration of extravisceral fluid were studied in the White Sea periwinkles (Littorina littorea, Littorina saxatilis and Littorina obtusata) during freshwater exposure. Resistance to hypoosmotic stress increased in the order: L. obtusata < L. saxatilis < L. littorea. Our data suggest that osmotic shock is not a primary reason for mortality of the periwinkles under these conditions. During environmental anaerobiosis, considerable succinate accumulation (up to 10–19 μmol g⁻¹ wet weight), and depletion of phosphagen and ATP pools were found in the studied species. Other metabolic end products (alanopine, strombine, lactate, acetate or propionate) were not detected. Succinate accumulation and net ATP breakdown were the fastest in the least resistant species, L. obtusata, and slowest in the most resistant, L. littorea. Rate of ATP turnover decreased during freshwater exposure in L. littorea and L. saxatilis, but not in L. obtusata. Our data suggest that differential resistance of three studied Littorina spp. to extreme hypoosmotic stress may be related to their different abilities to reduce metabolic rate and ATP turnover during sustained anoxia. Species-specific variations in anaerobic capacity of Littorina spp. are discussed in relation to their vertical distribution, size and ecology. Accepted: 4 October 1999     

Recruitment dynamics and first year growth of the coral reef surgeonfish Ctenochaetus striatus,with implications for acanthurid growth models

Newly recruited Ctenochaetus striatus were monitored over a 16-month period in American Samoa, 2002–2003. During this period, a mass recruitment of age-0 C. striatus occurred in March 2002 with numbers reaching 22.9 recruits m^?2. This program provided an invaluable opportunity to (1) analyze the dynamics of a mass recruitment episode and to assess its significance with respect to more typical patterns of recruitment and (2) establish the pattern of recruit growth during their first year of life. Age-based analysis indicated that the mass recruitment generated about 90 % of annual recruitment, but recruit mortality was high; thus, most recruitment was provided by continuous settlement throughout the year. The mass event appeared to be a short-lived pulse with recruits residing on the reef an average of 14.1 d compared with 161.1 d for other recruits. Recruits grew rapidly, achieving 90 % of their adult size during their first year, and they formed their first otolith annulus after 1 yr, thereby providing a firm basis for otolith interpretation of fish ages during the early life history phase of this species. The extensive age-based documentation of their first year growth in this study validates the distinctive “square” growth pattern exhibited by acanthurids as described in the literature (i.e., long life span with rapid initial growth that quickly reaches an asymptotic size), and it demonstrates the impact that the presence of age-0 fish has when generating growth parameters for populations exhibiting square growth. We found that the parameters from the re-parameterized von Bertalanffy growth function have preferred characteristics when modeling square growth in fish and that fixing age-at-length zero to pelagic larval duration is a preferable method to constrain growth models when lacking age-0 fish.     

Dynamics of a <Emphasis Type="Italic">Limecola</Emphasis> (<Emphasis Type="Italic">Macoma</Emphasis>) <Emphasis Type="Italic">balthica</Emphasis> population in a tidal flat area in the western Wadden Sea: effects of declining survival and recruitment

We followed the dynamics of the population of the bivalve Limecola (Macoma) balthica in the westernmost part of the Wadden Sea by monitoring for 44 years (1973–2016) its numbers and age composition at 15 sites in a 50-km² tidal-flat area. During the first half of this period, the annual recruitment and adult survival were at a relatively constant level, resulting in rather constant numbers. During the second half of the observation period, annual recruitment and adult survival showed declining trends, resulting in seriously reduced adult abundance. Sudden substantial reductions in adult survival started around 1996 at a few sites to spread over the entire area within 5 years, like an infectious disease. The resulting small adult stocks produced small numbers of recruits. The stock–recruitment curve showed an increasing part up to about 30 adults m^?2, followed by invariably successful recruitments at > 40 adults m^?2. Both recruitment and adult survival were negatively related to water temperatures. However, elevated temperatures after 1997 could not explain the very low survival rates observed after 1996 and the very low recruitment success after 2003. So far, recovery of the population has hardly taken place, with both recruitment and adult survival at lower levels than observed for the initial stable period.     

Factors organizing rocky intertidal communities of New England: Herbivory and predation in sheltered bays

In New England, U.S.A., shores exposed to severe wave action are dominated by the common blue mussel Mytilus edulis L. while moderately protected areas are covered with perennial algae. It is thought that algae are limited by mussels which are a superior competitor. Because the effectiveness of predators is inhibited by wave activity, it is assumed that the rate of predation, which varies across this environmental gradient, accounts for the observed distribution of mussels and algae.Shores along sheltered bays appear to be an exception to this pattern and this study addresses some of the possible causes. In New England bays, mussels and barnacles Semibalanus balanoides (L.) are the most common organisms on the solid surfaces in the lower intertidal zone. Perennial macroalgae, such as Chondrus crispas Stackhouse and Fucus vesiculosus L., are rare. The distribution and abundance of species differs from that on moderately protected shores and is similar to very exposed shores which are dominated by mussels and barnacles.Herbivory by the common periwinkle Littorina littorea (L.) limits the abundance of F. vesiculosus and indirectly affects the success of mussels. During 4 years of experimental manipulations, F. vesiculosus rarely recruited in the presence of periwinkles but dominated experimental surfaces if periwinkles were excluded. When experimental surfaces with F. vesiculosus, which had been protected from herbivory for > 1 year, were exposed to natural conditions, herbivores cleared most of the surfaces within several months. Recruitment by barnacles and mussels was higher when periwinkles were excluded. However, the effect of periwinkles on mussels was indirect; the snails reduced barnacle success and thus reduced mussel recruitment which was enhanced by the surface irregularities provided by barnacles.The occurrence of mussels in sheltered bays is not due to a lack of predators. Predators were commonly seen at all sites. Most mussels on experimental surfaces were removed <4 wk when surfaces were exposed to natural levels of predation. Experiments do not provide an explanation for the occurrence of mussels, although the enhancement of mussel recruitment by barnacles suggests that the availability of settlement sites may be important.     

Density-associated recruitment mediates coral population dynamics on a coral reef

Theory suggests that density-associated processes can modulate community resilience following declines in population size. Here, we demonstrate density-associated processes in two scleractinian populations on the outer reef of Moorea, French Polynesia, that are rapidly increasing in size following the effects of two catastrophic disturbances. Between 2006 and 2010, predation by the corallivorous crown-of-thorns sea star reduced coral cover by 93 %; in 2010, the dead coral skeletons were removed by a cyclone, and in 2011 and 2012, high coral recruitment initiated population recovery. Coral recruitment was associated with coral cover, but the relationship differed between two coral genera that are almost exclusively broadcast spawners in Moorea. Acroporids recruited at low densities, and the density of recruits was positively associated with cover of Acropora, whereas pocilloporids recruited at high densities, and densities of their recruits were negatively associated with cover of Pocillopora. Together, our results suggest that associations between adult cover and density of both juveniles and recruits can mediate rapid coral community recovery after large disturbances. The difference between taxa in sign of the relationships between recruit density and coral cover indicate that they reflect contrasting mechanisms with the potential to mediate temporal shifts in taxonomic composition of coral communities.     

Population biology and long-term mariculture studies in the brown alga <Emphasis Type="BoldItalic">Lessonia trabeculata</Emphasis> in Atacama,Chile

Lessonia trabeculata is one of the most valuable seaweeds in Chile, especially in the northern zone where its harvest has been going on for decades. We carried out population dynamics studies in the Atacama Desert coast (Bahia Chasco), in order to assess its productivity under natural and harvesting scenarios. We found very slow but consistent growth (1.98 cm month^?1) and density (3–4 individuals m^?2 with no monthly variation) during 18 months of observations in an undisturbed subpopulation. However, after total harvesting, L. trabeculata exhibited different responses. Its recruitment was season-specific, with exceedingly high values in autumn (ca. 80 individuals m^?2 in 5 months) and a dramatic reduction of recruits in summer (1–5 individuals m^?2 in 7 months, with many areas with no recruitment). Gradually, density values tended to stabilize to growth rates under un-altered conditions. In parallel, pruning systems at three different thallus levels (frond meristem base cuts, removal of half and total canopy) were all inefficient and harmful: (i) Biomass takes longer to be harvested; (ii) pruned individuals die off; and (iii) do not detach easily from the substrata, delaying the recovery by potentially emerging L. trabeculata juveniles. Some of these results agreed with our culture experiments, where 26 months were needed to obtain up to 100 cm long thalli with shrub-like morphology. We conclude that management of L. trabeculata beds must be improved in order to guarantee survival of the industry, and we propose some practices that at some stage should involve the complete removal of older/senescent individuals.     

Crystalline calcium in littorinid mucus trails

Previous work has shown that the feet of terrestrial and freshwater snails are important in calcium regulation, often secreting granules of CaCO₃. This phenomenon has not, until now, been observed in marine snails. Here we report the presence of CaCO₃ granules in the trail mucus of Littorina littorea (L.), L. saxatilis (Olivi) and L. obtusata (L.) Fixed mucus trails on plastic coverslips were examined by X-ray microanalysis under the SEM. Of the single-metal granules observed in the mucus trails the most abundant were of calcium (means: L. littorea, 440 mm⁻²; L. saxatilis, 401 mm⁻²; L. obtusata, 348 mm⁻²) followed for each species by silicon (maximum mean density: L. saxatilis, 120 mm⁻²) and iron (maximum mean density: L. saxatilis, 65 mm⁻²) granules. Single-metal granules of Al, Ti, Mg and P were also found but only in the mucus trails of L. obtusata, perhaps reflecting its different collection site from the other two species. The mean size of the calcium granules showed significant interspecific variation (L. littorea, 1.32 μm diameter±0 08 μm, n = 143; L. saxatilis, 1.80 μm±0.12, n = 113; L. obtusata, 2.14 μm±0.09, n = 167). Most calcium granules (L. littorea, 80%, n = 35; L. saxatilis, 57%, n = 113; L. obtusata, 69%, n = 167) were attached to, or embedded within, microthreads of mucus which tended to run parallel to the direction of locomotion. The significance of this is unknown although it may imply that the CaCO₃ granules are secreted with the mucus. It is concluded that calcium losses via this route are too small for pedal mucus to function significantly in ionoregulation of calcium. The calcium in the trail may therefore perform other functions, for example indicating trail polarity.     

Spatio-temporal distribution of recruits (0 group) of Merluccius merluccius and Phycis blennoides (Pisces, Gadiformes) in the Strait of Sicily (Central Mediterranean)

Aspects of the recruitment of hake (Merluccius merluccius L., 1758) and greater fork beard (Phycis blennoides Brunnich, 1768) in the Strait of Sicily (Central Mediterranean) are presented. Data were collected from 1994 to 1999 during the international bottom trawl survey program MEDITS. In view of the available literature on juvenile growth of these two species in the Mediterranean area, a length-based criterion was adopted to separate fish belonging to the 0 group (recruits). Recruit density indices (R/km²) by haul were calculated based upon the likely variability of recruit growth among years and used to study abundance variability and spatial pattern of recruitment and to identify the main nursery areas. Although there was inter-annual variability, two stable areas for M. merluccius were identified on the eastern side of the Adventure Bank and the Malta Bank at depths ranging between 100 and 200 m. The main nursery areas for P. blennoides were deeper (from 200 to 400 m) and two stable nursery areas were identified on the western and eastern side of the Adventure Bank; other nurseries were found in the easternmost part of the Strait in 1998 and 1999. Recruitments of the two species were significantly correlated, with the strongest recruitment occurring in 1998 and 1999 for each species.     

Congener comparison of native (Zostera marina) and introduced (Z. japonica) eelgrass at multiple scales within a Pacific Northwest estuary

A congener comparison of native (Zostera marina) and introduced (Zostera japonica) eelgrasses was conducted in Willapa Bay, Washington, USA. Along intertidal transects, Z. japonica (0.1–1.5 m above mean lower low water [MLLW]) occurred above Z. marina (<0.6 m MLLW). Both species declined in shoot length at higher elevation, but Z. japonica was always shorter (20 vs. 100 cm) and occurred at higher shoot density (>3,800 vs. <360 m^?2 in Z. marina). Z. japonica exhibited greater seasonal variation in biomass, with increases supported by both sustained asexual reproduction (rhizome branching) and recruitment from seeds (30 vs. <5% in Z. marina). Z. japonica’s successful invasion appears related to small size and high reproductive output, allowing it to spread quickly in a variable and stressful intertidal environment where competitive effects are low. Based on interannual changes in abundance, the native eelgrass has also recently increased in Willapa Bay, and one hypothesis involves “engineering” of suitable habitat at higher tidal elevations by Z. japonica.     

Above- and belowground competition between two submersed macrophytes

Several studies have shown that submerged macrophytes provide a refuge for zooplankton against fish predation, whereas the role of emergent and floating-leaved species, which are often dominant in eutrophic turbid lakes, is far less investigated. Zooplankton density in open water and amongst emergent and floating-leaved vegetation was monitored in a small, eutrophic lake (Frederiksborg Slotssø) in Denmark during July–October 2006. Emergent and floating-leaved macrophytes harboured significantly higher densities of pelagic as well as plant-associated zooplankton species, compared to the open water, even during periods where the predation pressure was presumably high (during the recruitment of 0+ fish fry). Zooplankton abundance in open water and among vegetation exhibited low values in July and peaked in August. Bosmina and Ceriodaphnia dominated the zooplankton community in the littoral vegetated areas (up to 4,400 ind l^?1 among Phragmites australis and 11,000 ind l^?1 between Polygonum amphibium stands), whereas the dominant species in the pelagic were Daphnia (up to 67 ind l^?1) and Cyclops (41 ind l^?1). The zooplankton density pattern observed was probably a consequence of concomitant modifications in the predation pressure, refuge availability and concentration of cyanobacteria in the lake. It is suggested that emergent and floating-leaved macrophytes may play an important role in enhancing water clarity due to increased grazing pressure by zooplankton migrating into the plant stands. As a consequence, especially in turbid lakes, the ecological role of these functional types of vegetation, and not merely that of submerged macrophyte species, should be taken into consideration.     

Specific dynamic action demonstrated in the herbivorous marine periwinkles,LIttorina littorea L. and Littorina obtusata L. (Mollusca,Gastropoda)

• 1.1. Respiration rates of starved periwinkles, Liltorina littorea and Littorina obtusata, increased by 40–60% when fed their preferred algal food for 1 hr, or after having been exposed for the same period to an aqueous extract of the alga.
• 2.2. The stimulus causing the rise in metabolic rate by algal extracts is therefore thought to be chemosensory in nature, and possibly composed primarily of dissolved free amino acids.
• 3.3. The respiration rates of L. littorea responded only to the green alga Enteromorpha intestinalis while L. obtusata demonstrated an increase in respiration rates when fed the ubiquitous brown alga, Ascophyllum nodosum, and to a lesser degree to Enteromorpha.
• 4.4. These results compare very well to postprandial increases in oxygen consumption demonstrated in vertebrates and marine bivalves where the components of specific dynamic action (SDA) are thought to be chiefly biosynthetic costs of digestion and assimilation.

     

Patterns of recruitment and microhabitat associations for three predatory coral reef fishes on the southern Great Barrier Reef, Australia

This study examined recruitment patterns and microhabitat associations for three carnivorous fishes, Plectropomus maculatus, Lutjanus carponotatus and Epinephelus quoyanus, at the Keppel Islands, southern Great Barrier Reef, Australia. Habitat selectivity was highest for recruits that were found mostly with corymbose Acropora, predominantly on patches of live coral located over loose substrates (sand). Adults were more commonly associated with tabular Acropora. The proportion of P. maculatus (72 %) found with live corals was higher than for L. carponotatus (68 %) and E. quoyanus (44 %). Densities of recruits were highly variable among locations, but this was only partly related to availability of preferred microhabitats. Our findings demonstrate that at least some carnivorous reef fishes, especially during early life-history stages, strongly associate with live corals. Such species will be highly sensitive to increasing degradation of coral reef habitats.     

Reproductive allocation in rhizomatous,seminiferous, and pseudoviviparous Leiothrix (Eriocaulaceae) species

Pseudoviviparous species are considered to rely almost entirely on vegetative propagation, because flowers are replaced by sprouts, and seedling recruitment is rare. We compared reproductive allocation (inflorescence and seed production) in three propagation modes presented by sympatric species of Leiothrix (Eriocaulaceae): rhizomatous and seminiferous (RS—L. crassifolia); rhizomatous, seminiferous, and pseudoviviparous (RSP—L. spiralis); and seminiferous and pseudoviviparous, in which seed production is considered rare due to reduced inflorescence size and small number of pistillate flowers (SP—L. arrecta and L. propinqua). We hypothesize that such propagation modes constitute a continuum of reproductive investment, that is: RS > RSP > SP. We harvested 154 Leiothrix spp. individuals at the Serra do Cipó, SE Brazil, from which we recorded the number of capitula and seeds, mean seed weight, and dry biomass allocated to plant structures. We then compared distinct propagation modes in relation to reproductive allocation, inflorescence-based and seed-based reproductive effort, and biomass partitioning. Contrary to expectations, the reproductive investment hierarchy found was SP ≥ RS > RSP. Our results do not fully concur with the current view that pseudoviviparous plants allocate few resources to reproduction. We suggest that the larger reproductive investment observed in SP helps to wait for better recruitment conditions in crowded and highly competitive mature populations (temporal escape), and to cope with destructive disturbances such as fire, since SP species lacks rhizomes.     

Synchronous annual recruitment variation in barnacles and ascidians in the White Sea shallow subtidal 1999–2010

Multiple foundation species in a community may exhibit alternative ecological strategies. Barnacles Balanus crenatus Bruguiere and solitary ascidians Styela spp. often co-dominate on mixed sediments in the White Sea shallow subtidal, supporting numerous dependent organisms. Larvae of B. crenatus stay in plankton for several weeks, while ascidian tadpoles float for 1–2 days. Given this difference in spreading potential, we expected recruitment in barnacles and ascidians to be controlled by the factors operating at different spatial scales. In 1999–2010, we annually sampled the community dominated by barnacles and ascidians to relate their recruitment rates to the substrate space availability, abundance of adults, and climatic variables. Most barnacles recruited to the surfaces of shells, stones, and conspecific adults. Ascidian recruits were chiefly found on barnacles. Annual recruitment rates of barnacles and ascidians were strictly correlated and strongly depended on average temperatures of the preceding fall (positively), winter (negatively), and current summer (negatively). Variation of mean annual recruitment rates was 26-fold for barnacles and 30-fold for ascidians. We found no limitation of recruitment by hard substrate availability. Inconsistent with our original hypothesis, large-scale environmental factors similarly accounted for most annual recruitment variation in both foundation species studied.