Survivorship of juvenile barnacles and mussels: spatial dependence and the origin of alternative communities

Mussels, barnacles, and rockweeds often form a distinct mosaic of patches on rocky intertidal shores, and it has been suggested that these communities may represent alternative community states. One way that alternative community states can arise is if early successional events are scale-dependent, but it is not known if juvenile survivorships of mussels and barnacles are, in fact, scale-dependent. Scale-dependence of barnacles (Semibalanus balanoides (L.)) and mussels (Mytilus edulis L.) was tested in the Gulf of Maine, USA. In winter 1997, clearings of 1, 2, 4 and 8 m in diameter and uncleared controls were made in stands of the rockweed Ascophyllum nodosum (L.) Le Jolis at 12 sites spread evenly across four bays on Swan's Island, ME. Summer and fall-winter survivorship of barnacles, which recruited in spring 1997, were estimated by tracking the 1997 cohort until late winter 1998. Survivorship of mussels was estimated from following the fate of transplanted juveniles over 4 days in late August 1997. Both barnacles and mussels showed better survival in 4 and 8 m clearings than in small clearings and controls. There was also significant variation in survivorship among sites. Densities of gastropods in the clearings did not reflect survivorship patterns of barnacles and mussels. Barnacle survivorship increased in fall and winter, and in large clearings was comparable to survivorship in barnacle-dominated habitats. Mussel survivorship was low in all clearing sizes suggesting that mussel beds develop slowly.     

Experimental confirmation of multiple community states in a marine ecosystem

Small changes in environmental conditions can unexpectedly tip an ecosystem from one community type to another, and these often irreversible shifts have been observed in semi-arid grasslands, freshwater lakes and ponds, coral reefs, and kelp forests. A commonly accepted explanation is that these ecosystems contain multiple stable points, but experimental tests confirming multiple stable states have proven elusive. Here we present a novel approach and show that mussel beds and rockweed stands are multiple stable states on intertidal shores in the Gulf of Maine, USA. Using broad-scale observational data and long-term data from experimental clearings, we show that the removal of rockweed by winter ice scour can tip persistent rockweed stands to mussel beds. The observational data were analyzed with Anderson’s discriminant analysis of principal coordinates, which provided an objective function to separate mussel beds from rockweed stands. The function was then applied to 55 experimental plots, which had been established in rockweed stands in 1996. Based on 2005 data, all uncleared controls and all but one of the small clearings were classified as rockweed stands; 37% of the large clearings were classified as mussel beds. Our results address the establishment of mussels versus rockweeds and complement rather than refute the current paradigm that mussel beds and rockweed stands, once established, are maintained by site-specific differences in strong consumer control. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Using patterns of variability to test for multiple community states on rocky intertidal shores

Predictions based on theory of multiple stable states suggest that larger perturbations should lead to more unpredictable patterns of succession. This prediction was tested in the Gulf of Maine using data from 60 intertidal plots of varying size that were experimentally cleared of the rockweed Ascophyllum nodosum and from 14 benchmark sites from throughout the Gulf. Rockweed was removed from the experimental clearings ranging from 1 to 8 m in diameter in 1996 and data collected in 2004 were used to test effects of clearing size and location on divergence and variability in species composition. Benchmark data were collected in 2005, and the 14 sites were from a dataset on 53 sites throughout the Gulf of Maine. The selected sites were randomly chosen from all sites with > 80% canopy cover by A. nodosum and were expected to be similar to uncleared control plots from the experiment. Experimental removal of A. nodosum resulted in clearings at 12 sites within 4 bays. Abundances of gastropods, barnacles, mussels, and fucoid algae and the percentage cover of barnacles, mussels, fucoid algae, bare space, and other species were sampled. CAP and PERMDISP analyses revealed significant differences in multivariate dispersion and variability with both clearing size and location. Variability generally increased with clearing size and location effects were related to the north-south positioning of the sites. Benchmark sites were similar to the experimental control plots but as variable as the largest clearings. Results suggest that succession in larger clearings has been more unpredictable than in small clearings. The pattern of variability in the experimental clearings is consistent with the predictions of multiple stable states. However, the large amount of variation among the benchmark sites was due to mussels and was unexpected. This unexpected variability underscores the importance of sampling benchmark sites as part of experiments.     

First year demography of the foundation species, Ascophyllum nodosum, and its community implications

Long-lived species of large size can play critical roles as ecosystem engineers but the demography of early life stages is poorly known for many such species. For example, the fucoid alga, Ascophyllum nodosum , serves as the foundation species for many sheltered rocky shore communities of the North Atlantic Ocean, yet its early demography is incompletely characterized. Here we provide data on age-specific survivorship throughout the first year. We outplanted zygotes generated from artificial fertilizations and followed their survivorship under different conditions that tested the following hypotheses: (1) survivorship is inversely related to clearing size, (2) germlings survive better in clearings in conspecific stands than in clearings in mussel beds, (3) survivorship in microhabitat refugia (grooves of artificial surfaces) is greater than that on flat surfaces, and (4) protection from grazing and physical factors, like canopy sweeping and desiccation, enhances germling survival. Germlings were censused on days 2, 9, 13, 17, 64, 102, 127 and 399 following outplanting. Germling mortality was consistently higher on flats than in grooves of artificial surfaces, and was greatest within the first two days of life. Germlings protected by cages survived better during the day 0–2 interval, but after 2 days caging did not matter. Interval-specific mortalities after day 2 showed that survivorship was inversely related to clearing size. The type of clearing, in conspecific stands or mussel beds, did not affect survivorship. Ascophyllum nodosum shows a strong type III survivorship curve with mortality during year 1 exceeding 99.9%. The poor survivorship of first year germlings accounts for the paucity of newly recruited juveniles often observed in mature stands of A. nodosum . The lack of juveniles and the scale dependent survivorship of germlings renders populations of A. nodosum , and the associated community that depends on it, vulnerable to large scale perturbations.     

Changes in composition of rockweed (Ascophyllum nodosum) beds due to possible recent increase in sea temperature in Eastern Canada

Ascophyllum nodosum (rockweed) is the main economic resource of the seaweed industry in the Atlantic Provinces of Canada. The annual harvest steadily increased since 1995, reaching a historic peak of 37,000 tonnes in 2007. Due to a high demand for fertilizers and animal feed supplements derived from rockweed, this trend seems likely to continue. The current management plan for the sustainable harvest of the A. nodosum resource is considered conservative. The resource has been managed with a precautionary approach since 1995 to protect the integrity of the habitat. Acadian Seaplants Limited (ASL) has been granted approximately 90% of the government-issued licenses to harvest A. nodosum resources in the Maritimes. The Canadian approach is based on an annual harvest from a given bed and not strip-and-return after several years. Since 1995, ASL has proactively undertaken extensive annual surveys and research on biomass productivity of this renewable resource to establish acceptable annual exploitation rates. Historically, the rockweed beds of southwestern Nova Scotia (NS) have been almost 99% pure A. nodosum, with a minor component of Fucus vesiculosus. However, since 2004, a steady increase in F. vesiculosus, with a peak of 4.6% of the total biomass in 2008, was recorded. This coincided with one of the mildest winters on record for the Maritimes. This increase in temperature seemed to be also responsible for an unusual recruitment of the blue mussel Mytilus edulis in rockweed beds in some areas of southern New Brunswick (NB) in 2006, causing the detachment of up to 30% of the seaweed biomass in some harvesting sectors. Another phenomenon observed in southwestern NS during 2003 and 2004 was extensive ice damage on rockweed beds produced by an early melting of the ice, with losses of up to 90% of the rockweed biomass in some areas.     

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.     

Barnacle larval supply to sheltered rocky shores: a limiting factor?

In northwest Europe, sheltered rocky shores are dominated by fucoid canopy algae and barnacles are rare, although the latter are extremely abundant on exposed shores. The supply of the intertidal barnacle Semibalanus balanoides (L.) to sheltered, fucoid dominated rocky shores was investigated to determine the importance of larval supply in limiting the abundance of adults in shelter. Larval supply was measured at two spatial scales, at the scale of shore (100s of metres), by comparing larval concentrations at exposed and sheltered sites, and at a smaller spatial scale (m), by examining the role of fucoid canopies in limiting supply to the substratum. Replicate plankton trawls were carried out above the intertidal zone at high water at two sheltered sites and nearby exposed headlands. The concentration of S. balanoides cyprid larvae was significantly higher at the sheltered sites on two out of three sampling occasions with up to 14 times greater larvae on one occasion than the nearby exposed site. The effect of the macroalgal canopy on supply to the substratum was assessed in two ways: directly, by pumping water from the substratum in areas with and without Ascophyllum nodosum (L.) Le Jolis, and indirectly by measuring cyprid settlement in a canopy-manipulation experiment. Pumped plankton samples from mid tide level showed that the A. nodosum canopy did not form a barrier to larval supply and may have had a positive effect on larval concentrations at the substratum. Cyprid settlement was assessed in the mid shore A. nodosum and low shore Fucus serratus L. zones to areas with canopy algae (but protected from the sweeping effects of macroalgal fronds) and without canopy. Settlement over three consecutive 24-h periods showed a consistent pattern; settlement was consistently lower beneath the F. serratus canopy than in cleared areas, suggesting that this algal species forms a barrier, limiting supply of cyprid larvae to the substratum.     

Management and production of the brown algae Ascophyllum nodosum in the Canadian maritimes

Commercial exploitation of Ascophyllum nodosum (rockweed) along the coast of Nova Scotia began in the late 1950s when it was used as a raw material for manufacturing alginate and “kelp” meal. Today, this resource is used as a biostimulant extract for crops and animal feed supplements and is the main economic resource of the seaweed industry in the Maritime Provinces and Canada. The management of rockweed, by dividing the resource into many sectors, permitted the assessment of yield per unit area of bed. In Nova Scotia, mechanical harvesting of A. nodosum operated on a pulse of 2- to 3-year schedule providing yields of 21.9 to 47.7?wet?t ha^?1 averaging 35.3?±?7.6?wet?t ha^?1. Acadian Seaplants Limited (ASL) has become the dominant player in the region since 1995 with more than 75% of the total biomass under its leases and more than 90% of the total landing of rockweed in the last 13?years. Sustained harvests at ASL leases and under an annual harvest schedule using hand cutting methods have averaged 17.4?±?2.6?t ha^?1. Exploitation rates above 35% of the harvestable crop lead to a pulse harvest strategy and the need to move infrastructure year after year. In New Brunswick, a new approach to management began in 1995 with a regulated 17% exploitation rate. Thus, the yield in that province is 14.3?wet?t ha^?1. The current summer standing crop for this region has been calculated at 352,723?wet?t, covering an area of 4,960?ha, with an average biomass of 71.3?t ha^?1. The harvest in the region reached peak landings in 2010 with just over 40,100?t. The consistent yield per hectare of A. nodosum beds is proof of good management practices and an ecologically sustainable harvest in the Canadian Maritimes.     

Hierarchical spatial structure in soft-bottom mussel beds

Mussels (Mytilus edulis L.) are unusual because they thrive in both rocky shore and soft-bottom habitats. Despite their ecological and economic importance, little is known about their spatial structure. Mussels do not generally recruit to bare soft substrate because larvae and postlarvae cannot attach to a bottom of small sediment particles. They attach to hard objects on the sediment surface (especially other mussels), so soft-bottom mussel beds may be spatially organized in ways that are fundamentally different from those on rocky shores. The purpose of our study was to characterize the scales of spatial variability for several mussel abundance parameters in soft-bottom, intertidal M. edulis beds in coastal Maine. We used a random factor nested-ANOVA design of 200 cm² Cores within 1 m² Quadrats within 6 m Transects within Positions within bed Sites along 70 km (euclidean distance) of the Maine coast. Based on the literature and our field observations, we hypothesized that Sites and Positions account for most of the spatial variance in soft-botttom mussel beds. We rejected this hypothesis. Sites and Positions were not important in explaining variation in total mussel density, density of new recruits, or density of larger mussels. Although most of the variance in surface silt-clay fraction did occur at these levels, most mussel variation occurred at smaller spatial scales, specifically at the Quadrat scale for new recruits and total mussels and at the Transect scale for larger mussels. Variance in mussel parameters was not closely linked to the silt-clay fraction of surface sediment or to Site rankings of wind exposure and tidal flow. Variance in total mussel density was due primarily to variance in recruitment. No single scale explained more than about half the mussel variance, and no single scale was best at explaining all the mussel parameters. Greater knowledge about mussel bed spatial variability would be useful because it can help direct scale-dependent sampling regimes, field experiments, and coastal management practices.     

The effects of seaweed harvesting on fishes: a critique

Synopsis Black & Miller (1991) concluded that there was no large impact of an experimental harvest of rockweed,Ascophyllum nodosum, on fishes. A critique of their study demonstrates that this conclusion cannot be substantiated by their data because of sampling biases, errors in experimental design and low statistical power. Further, evidence is presented which supports the hypothesis that rockweed may provide an important nursery habitat for juvenile fishes.     

Mussels matter: postlarval dispersal dynamics altered by a spatially complex ecosystem engineer

This study investigated postlarval dispersal of soft-bottom macrofauna at a spatially complex intertidal mudflat comprising patches of bare sediment and an ecosystem engineer, the mussel Mytilus edulis. At each of four sites in Guard Point Cove, Maine, USA, we took core samples and deployed bedload traps in bare sediment and mussel bed habitats to estimate ambient densities, rates of sediment flux, and several measures of postlarval dispersal. Univariate and multivariate nonmetric multidimensional scaling (nMDS) results showed few significant site effects and no habitat×site interactions. In contrast, there were numerous significant habitat effects. Compared to the bare sediment, the mussel bed habitat had: fewer species; higher ambient density and proportional abundance of the oligochaete Tubificoides benedeni (the dominant species in both habitats); lower ambient densities and proportional abundances of major taxa and the nonoligochaetes as a group; and higher sediment flux and relative (i.e., per capita) dispersal of nonoligochaetes. Macrofauna species dispersed in relative proportions that were different from those in the ambient assemblage. Per capita T. benedeni transport rates were low in mussel beds compared to those for nonoligochaetes, consistent with the view that beds represent favorable habitat for oligochaetes. The number of total macrofauna individuals trap⁻¹ day⁻¹ was negatively correlated with ambient density and positively correlated with sediment flux in both habitats, but these relationships were significant only in the mussel bed. The results indicate that altered transport rates of sediment and postlarvae are important mechanisms by which mussels act as ecosystem engineers to modify soft-bottom habitats. Differential transport rates caused by aggregations of mussels and other foundation species must be considered in explanations of spatial pattern in soft-bottom communities.     

Effects of canopy gaps on the growth of tree seedlings from subtropical broad-leaved evergreen forests of southern China

Seedling performance, morphology, and leaf characteristics of evergreen tree species from different successional stages of Chinese broadleaved evergreen forests were studied in four simulated gap environments: in 100, 55, 33, and 18% of full sunlight. The first two represent the light regimes of clearings (gaps secondarily become more open due to human activity and natural erosion). The hypothesis was tested that early-successional species achieve their greatest seedling size in clearings and contrast in this respect with late-successional species. Late-successional species were expected to show a stronger morphological response (leaf/root ratio) to clearings than early-successional ones. The results provided some evidence in support of these hypotheses with one exception, namely that late-successional Castanopsis fargesii appeared to be a highly sun-tolerant species. It is suggested that Castanopsis seedlings are competitive in large gaps and clearings.     

Effects of small-scale disturbances of canopy and grazing on intertidal assemblages on the Swedish west coast

The effects of small-scale disturbances (80×30-cm plots) of canopy and grazers on intertidal assemblages were investigated in this 4-year experiment on sheltered rocky shores on the Swedish west coast. Canopy disturbances due to ice scouring were mimicked by removal of adult plants of the seaweed Ascophyllum nodosum (L.) Le Joli. Density of the main epilithic grazing gastropods, Littorina spp., was lowered by exclosure and handpicking. Based on earlier experiments in other areas, the general hypothesis was that canopy removal and grazer exclosure, alone or in combination, should increase the recruitment of A. nodosum or other fucoid juveniles, and change the structure of the understorey assemblage.There was an effect of canopy removal on the development of this assemblage, lasting for more than 31 months. Both increased and decreased abundances of species were found as short-term effects, but there was also a longer-term effect with increased abundance. Grazer exclosure was only effective in combination with canopy removal, causing a short-term increase in ephemeral green algae. Short-term effects of canopy removal were also the increase in recruitment of Semibalanus balanoides (Linnaeus) and the decrease of the red alga Hildenbrandia rubra (Sommerfelt) Meneghini. Fast recruitment and growth of fucoid species (Fucus serratus L. and F. vesiculosus L.) restored the canopy and conditions of the understorey within 18 months. Thus, the canopy removal changed the physical conditions for the understorey, making it possible for other species to coexist in this community. Surprisingly, no effect of canopy removal or grazer exclusion was found on the recruitment of juvenile A. nodosum, neither by canopy removal nor grazer exclosure. The lack of such effects might be due to the early mortality caused by other grazers (small, mobile crustaceans), or to the low density of periwinkles on these shores. However, despite the patchy and generally low recruitment of A. nodosum juveniles, observations suggested that the cover of A. nodosum in manipulated patches would return to initial levels, either by recruitment or regrowth of small holdfasts and from growth of edge plants.     

Cheating the Locals: Invasive Mussels Steal and Benefit from the Cooling Effect of Indigenous Mussels

The indigenous South African mussel Perna perna gapes during periods of aerial exposure to maintain aerobic respiration. This behaviour has no effect on the body temperatures of isolated individuals, but when surrounded by conspecifics, beneficial cooling effects of gaping emerge. It is uncertain, however, whether the presence of the invasive mussel Mytilus galloprovincialis limits the ability of P. perna for collective thermoregulation. We investigated whether varying densities of P. perna and M. galloprovincialis influences the thermal properties of both natural and artificial mussel beds during periods of emersion. Using infrared thermography, body temperatures of P. perna within mixed artificial beds were shown to increase faster and reach higher temperatures than individuals in conspecific beds, indicating that the presence of M. galloprovincialis limits the group cooling effects of gaping. In contrast, body temperatures of M. galloprovincialis within mixed artificial mussel beds increased slower and exhibited lower temperatures than for individuals in beds comprised entirely of M. galloprovincialis. Interestingly, differences in bed temperatures and heating rates were largely dependent on the size of mussels, with beds comprised of larger individuals experiencing less thermal stress irrespective of species composition. The small-scale patterns of thermal stress detected within manipulated beds were not observed within naturally occurring mixed mussel beds. We propose that small-scale differences in topography, size-structure, mussel bed size and the presence of organisms encrusting the mussel shells mask the effects of gaping behaviour within natural mussel beds. Nevertheless, the results from our manipulative experiment indicate that the invasive species M. galloprovincialis steals thermal properties as well as resources from the indigenous mussel P. perna. This may have significant implications for predicting how the co-existence of these two species may change as global temperatures continue to rise.     

Effects ofFucus vesiculosus covering intertidal mussel beds in the Wadden Sea

The brown algaFucus vesiculosus formamytili (Nienburg) Nienhuis covered about 70% of mussel bed (Mytilus edulis) surface area in the lower intertidal zone of Königshafen, a sheltered sandy bay near the island of Sylt in the North Sea. Mean biomass in dense patches was 584 g ash-free dry weight m^?2 in summer. On experimental mussel beds, fucoid cover enhanced mud accumulation and decreased mussel density. The position of mussels underneath algal canopy was mainly endobenthic (87% of mussels with >1/3 of shell sunk into mud). In the absence of fucoids, mussels generated epibenthic garlands (81% of mussels with <1/3 of shell buried in mud). Mussel density underneath fucoid cover was 40 to 73% of mussel density without algae. On natural beds, barnacles (Balanidae), periwinkles (Littorina littorea) and crabs (particularly juveniles ofCarcinus maenas) were significantly less abundant in the presence of fucoids, presumably because most of the mussels were covered with sediment, whereas in the absence of fucoids, epibenthic mussel clumps provided substratum as well as interstitial hiding places. The endobenthic macrofauna showed little difference between covered and uncovered mussel beds. On the other hand, grazing herbivores — the flat periwinkleLittorina mariae, the isopodJaera albifrons and the amphipodsGammarus spp. — were more abundant at equivalent sites with fucoid cover. The patchy growth ofFucus vesiculosus on mussel beds in the intertidal Wadden Sea affects mussels and their epibionts negatively, but supports various herbivores and increases overall benthic diversity.     

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.     

Stable musselMytilus edulis beds in the Wadden Sea—They’re just for the birds

Predation by eiders, oystercatchers and herring gulls on natural mussel bedsMytilus edulis was studied in the Königshafen, a sheltered bay in the Wadden Sea. About 15 ha (2.5%) of the Königshafen were covered with mussel patches of a biomass of about 1300 g AFDW m^?2. The biomass on the mussel beds was dominated by old mussels and found to be constant over several years. Birds annually removed 30% of the standing stock. Eiders were by far the most important predators and consumed 346 g AFDW m^?2, followed by oystercatchers with 28 g AFDW m^?2 and herring gulls with 3.6 g AFDW m^?2. Birds consumed a substantial part of the annual production of the mussel beds which was estimated from literature data to be approx. 500 to 600 g AFDW m^?2. As other predators were absent, the production of the mussels was sufficient to sustain the high predation rate by birds. Stable mussel beds form a short and efficient link between primary production and bird predation which is unusual for the Wadden Sea, where the main part of primary food supply is thought to be unavailable for higher trophic levels.     

Tree selection by the endangered beetle <Emphasis Type="Italic">Rosalia alpina</Emphasis> in a lapsed pollard beech forest

Abandoned pollard beech forests are particular habitats that may require special conservation efforts to preserve the endangered beetle Rosalia alpina, a model species whose protection may perpetuate the habitat of many other saproxylic species. Forest use can determine the tree selection and population size of R. alpina, variables previously not researched in pollard forests. Selected tree traits and population size, indicated by presence of adult specimens and recent exit holes, were determined in a pollard beech forest before the implementation of habitat manipulations targeted to preserve the species. Multivariate analysis showed tree condition (living, snag, fallen) and clearing size and aspect as influential variables, with the former explaining more variance than the latter. R. alpina positively selected pollard snags and trees located in big and dry clearings, avoiding those in small clearings or shaded areas. Snags showed most adults (90 %) and recent emergence holes (84 %). Distance to the nearest occupied tree, trunk diameter, bark thickness and presence of sap leaks had no effect on the occurrence of R. alpina. Population size was estimated in 0.1 individuals day^?1 ha^?1, and 38 % of available habitat trees showed exit holes, values lower than those observed in other European countries. These results show the need to start re-pollardings in the forest to extend pollard standing life and enhance exposure to sunlight. Pinpointing pollard snags and clearing the surrounding vegetation are recommended as the first measures to be taken in order to favor R. alpina populations on pollard stands with closed canopies.     

Survival of cold-adapted species in isolated mountains: the population genetics of the Sudeten ringlet,Erebia sudetica sudetica,in the Jeseník Mts., Czech Republic

Relic populations of cold-adapted species, trapped in isolated mountain pockets within the temperate zone, are predicted to suffer considerably due to ongoing climate warming. The butterfly Erebia sudetica sudetica is an example restricted to the Eastern Sudety Mts. Here, the butterfly forms permanent populations on subalpine tall-herb grasslands, but also occupies woodland clearings and hay meadows at lower altitudes. We assume differences among the genetic diversities of the populations due to differences in the temporal continuity of these habitats. Therefore, 17 allozyme loci were analysed for 276 individuals from 13 different localities (six tall-herb stands, two meadows, five forest clearings) in the Jeseník Mts. with a maximum distance of 20 km among them. We obtained a significantly higher genetic diversity for the subalpine populations than for the forest clearing populations. The genetic differentiation among the forest clearing populations was higher than among the subalpine ones. They also showed a significant isolation-by-distance system. These findings support the idea that the lower-elevation populations might have been founded by more than one dispersal event from the subalpine sites, but also secondary colonisations and gene flow in the forest belt. Due to founder effects and possibly further subsequent bottlenecks, these forest clearing populations did not harbour the entire genetic diversity of the taxon. Therefore, conservation actions should focus on the subalpine tall-herb formation.     

Arrested succession in logging gaps: is tree seedling growth and survival limiting?

Thirty years after selective timber harvest in the Kibale National Park, Uganda, many abandoned logging gaps are dominated by Acanthus pubescens, and show little forest recovery. To examine if this arrested successional state was caused by limited tree seedling growth and survival, we planted seedlings of four forest tree species (Albizia grandibracteata, Mimusops bagshawei, Prunus africana and Uvariopsis congensis) in A. pubescens‐dominated logging gaps and in control areas of adjacent forest. To assess if clearing A. pubescens facilitates forest regeneration, we planted seedlings of two species (A. grandibracteata and U. congensis) in small clearings cut within the logging gaps. We examined mortality, growth, herbivory and site characteristics among the treatments. Finally, we described the physical attributes of the A. pubescens‐dominated gaps. Seedlings of all the four species survived and grew equally well in A. pubescens and forest treatments, and most site characteristics were also similar. Seedlings planted in clearings grew more than in either forest or A. pubescens sites. Very few established trees were found in A. pubescens sites, and most of these were near the forest edges. We also discussed the role of elephants (Loxodonta africana) and collapsing A. pubescens canopies in the maintenance of an arrested successional state in these logging gaps.