Differences in environmental response between the sexes of the dioecious shrub Baccharis halimifolia (Compositae)

Summary Baccharis halimifolia (Compositae) is a dioecious shrub which grows on the upland fringe of tidal marshes along the Atlantic and Gulf Coasts of North America. We examined the responses of the two sexes to variation in nutrient and moisture availability plant density, and defoliation. By growing plants from seedlings to flowering adults under various combinations of soil type, fertilization rate and plant density, we were able to establish different rates of plant growth and mortality. Plants grown at high density and low nutrient and water supply grew the least, incurrent the most mortality and showed a male-biased sex ratio (73% male). At low density with abundant nutrients and water, plants grew more, survived well, flowered frequently, and were female-biased (75% female). Changes in sex ratio were probably the result of sex-related mortality rather than sexual lability of the seedlings. While changes in sex ratio occurred under experimental conditions in the green-house, no evidence for differences in habitat utilization between the sexes were found in the field and the sex ratio (59% female) did not vary across habitats. In the marsh habitats we sampled where water and nutrients were apparently available, there was no evidence for differential mortality between the sexes. When defoliated (75% of leaf tissue), both sexes showed similar reductions in reproductive effort (number of flower heads/shoot). Our results indicate that differences between the sexes of Baccharis in their response to environmental growing conditions is an important ecological factor associated with the separation of male and female function into separate individuals.     

Replacement of estuarine communities by an exotic shrub: distribution and invasion history of Baccharis halimifolia in Europe

Baccharis halimifolia L. (Asteraceae) is a shrub native to North America which is invading estuarine communities in Europe. We report the invasion history and the distribution limits of B. halimifolia in Europe, with particular emphasis on the frequency of its presence in estuarine communities in Spain. B. halimifolia has been cultivated in Europe since the 17th century to present. It was first recorded as naturalized in the Bay of Biscay in 1906, where it forms currently stable and locally abundant populations in almost all the estuaries of Northern Spain and Western France. The ongoing invasion to the west could reach well conserved estuarine communities in Galicia (Spain). B. halimifolia also forms scattered populations in Northern and Southern France, Belgium, Netherlands, United Kingdom and Italy. In these countries it has experienced a rapid expansion during the last years. In Northern Spain, subhalophilous communities dominated by rush (Juncus maritimus) and/or sea couch (Elytrigia atherica), common reed stands (Phragmites australis) and ungrazed wet meadows are the most vulnerable to invasion. The subhalophilous communities are part of natural habitats of community interest according to the habitats directive 92/43/EEC. In some areas of Northern Spain these communities have been totally replaced by monospecific stands of B. halimifolia. In contrast, halophilous communities of the low marsh are resistant to invasion, suggesting that the survival of B. halimifolia may be limited by threshold values of salinity and waterlogging. With this study we want to raise awareness about the risk of replacement of estuarine subhalophilous communities by the ongoing invasion of B. halimifolia in Europe.     

Patterns of growth,reproduction, defense,and herbivory in the dioecious shrub Baccharis halimifolia (Compositae)

Summary Patterns of growth, reproduction, defense (leaf resin) and herbivory were compared between the sexes of the dioecious shrub Baccharis halimifolia (Compositae). Male plants possessed longer shoots and more tender leaves, grew faster, and flowered and senesced earlier than female plants. Levels of leaf nitrogen, water content, and acetone-soluble resin (shown to deter feeding by polyphagous insect herbivores) did not differ between male and female plants. When offered a choice between leaves from male and female plants, adults of two leaf beetles (Chrysomelidae), the monophagous Trirhabda bacharidis and the polyphagous Paria thoracica, both preferred to feed on male leaves. Similarly, the daily fecundity of older females of T. bacharidis was higher when they were fed leaves from male compared to female plants. However, adult survivorship and total fecundity of T. baccharidis did not differ between male and female leaf treatments. We attribute the feeding preference for and slight increase in fecundity on male plants to the tenderness of male leaves. Larvae of the fly Tephritis subpura (Tephritidae) fed exclusively in the sterile receptacle of male flower heads (85% infested), but the phenology was such that pollen production was not adversely affected. Larvae of two other flies Dasineura sp. and Contarinia sp. (Cecidomyiidae) occupied >95% of only female flower heads where they fed among and on the developing seeds. We conclude that foliage-feeding herbivores are unlikely candidates to explain the female-biased sex ratio (59% female) of B. halimifolia plants in the field, and that their preference for male plants is a result of plant characteristics (e.g. rapid growth) that have been selected by some other factor. However, our data on selective floral herbivory in B. halimifolia are in accord with the argument that dioecy reduces the inadvertent loss of flower parts of one sex when herbivores feed on flower parts of the opposite sex.     

Feeding responses of adapted and non-adapted insects to the defensive properties of Baccharis halimifolia L. (Compositae)

Summary Trirhabda bacharidis (Weber) (Chrysomelidae), a univoltine, monophagous beetle is the dominant herbivore on Baccharis halimifolia (Compositae), a woody, perennial shrub that leafs out in early spring and retains its leaves into November. Available plant biomass increases during the season but T. bacharidis feeds only during spring and early summer. During the remainder of the growing season, there are no major herbivores feeding on B. halimifolia. B. halimifolia leaves increase in toughness and thickness and decrease in moisture and nutrients as the season progresses. In feeding preference tests, T. bacharidis larvae preferred young leaves over leaves of older age classes. When reared on leaves of different ages, larvae fed on young leaves, weighed more, pupated earlier and had greater survivorship. T. bacharidis larvae showed no significant feeding preference for similarly tough B. halimifolia leaves painted with differing concentrations of late season acetone leaf extract. Elm leaves painted with the same leaf extracts were avoided by larvae of the non-adapted specialist Pyrrhalta luteola (Mueller) (Chrysomelidae).These results suggest that the adapted specialist, T. bacharidis, is not deterred by the B. halimifolia acetone soluble secondary chemical which increases in amount over the season. Decreasing leaf nitrogen (perhaps in concert with increasing leaf toughness) seems to be the primary factor that dissuades its feeding. However, acetone soluble secondary chemicals in the leaves of B. halimifolia may be effective in preventing herbivory from non-adapted insects.     

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

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

Ecology matters: Atlantic-Mediterranean disjunction in the sand-dune shrub Armeria pungens (Plumbaginaceae)

Inferring the evolutionary history of Mediterranean plant lineages from current genetic, distributional and taxonomic patterns is complex because of a number of palaeoclimatic and geological interconnected factors together with landscape heterogeneity and human influence. Therefore, choosing spatially simplified systems as study groups is a suitable approach. An amplified fragment length polymorphism (AFLP) study using two restriction enzyme combinations (EcoRI/MseI and KpnI/MseI) was carried out to estimate the structure of genetic variation throughout the range of Armeria pungens. This species has a West Iberian-Corso/Sardinian disjunct distribution on coastal sand-dune ecosystems. Bayesian, amova and genetic distance analyses of the AFLP data revealed the same distinguishable genetic groups, which do not match the main geographical disjunction. Corso-Sardinian populations were found to be genetically closer to southwest Portuguese than to those from the Gulf of Cadiz (the closest geographically). Eastwards long-distance dispersal is therefore invoked to explain this geographical disjunction. A GIS analysis based on bioclimatic envelope modelling aiming to characterize the current locations of A. pungens found strong similarities between the Portugal and Corsica-Sardinia sites and less so between these areas and the Gulf of Cadiz. This coincident pattern between AFLP and climatic data suggests that the geographical disjunction is better explained by climatic factors than by the likeliness of a stochastic dispersal event. Such a combined phylogeographical-GIS modelling approach proves to be enlightening in reconstructing the evolutionary history of plant species.     

Assessment of the spatial variability of intertidal benthic communities by axial tomodensitometry: importance of fine-scale heterogeneity

At the water-sediment interface of aquatic ecosystems, the presence of biogenic structures produced by benthic invertebrates strongly affects biogeochemical processes. The quantification of these structures and the assessment of the vertical distribution of fauna are essential for determining the impact of communities in sediments. In the present study, computer axial tomodensitometry (CAT-scan) was used to measure the space occupied by an intertidal community of the St. Lawrence estuary. Three cores were sampled at a site that was considered homogeneous according to surface sediments. The vertical distribution of biogenic structures and gravel were measured in the three cores using CAT-scan; the vertical distribution of fauna was also analysed for each core. The biogenic structures were highest at the water-sediment interface and decreased with depth in the three cores. The number of invertebrates also decreased with depth. We observed similar distributions of biogenic structures in cores 1 and 2. However, fewer biogenic structures were observed below 90 mm in core 3. This result was correlated with a high quantity of gravel from 90 to 140 mm in core 3 whereas the other cores had lower quantities of coarse material. We found relationships among the distributions of biogenic structures, fauna, and sediment characteristics (gravel quantity) that can affect species distribution. The vertical distributions of Macoma balthica, Mya arenaria, Nereis virens, and small-sized gallery-producing species (nematodes and oligochaetes) could also be recorded with the CAT-scan method. Thus, CAT-scan is an excellent tool to determine the fine-scale heterogeneity in the space occupied by benthic invertebrates in sediments.     

Sexual differences in reproductive phenology and their consequences for the demography of Baccharis dracunculifolia (Asteraceae), a dioecious tropical shrub

Patterns of phenological variation and reproductive investment were studied in the dioecious shrub Baccharis dracunculifolia DC (Asteraceae), and possible consequences on survivorship were evaluated. The sex ratio was determined in a natural field population (n = 921) of B. dracunculifolia in Belo Horizonte, Brazil. Fifty-two males and 56 females were sampled at random from this population. During the reproductive season of 1999, inflorescence production, shoot growth and mortality, and xylem water potential were recorded for each individual. The population sex ratio was male-biased (1.27 : 1, P < 0.05), and was associated with a higher mortality of female shoots (38.4 vs. 23.1 %, P < 0.05), and individuals (17.8 vs. 11.5 %, P < 0.1), despite lower water stress in female plants. Flowering phenology also differed between the sexes, with males producing more inflorescences, and earlier, than females. Owing to fruit maturation, the number of inflorescences supported by females was higher than that supported by males later in the reproductive season. This occurred during the dry season, and drought stress may have been responsible for the greater female mortality. Thus, the male-biased sex ratio in this population of B. dracunculifolia is probably due to different reproductive functions of males and females. Intersexual differences in reproductive phenology had consequences for plant demography.     

Zonation of ground beetles (Coleoptera: Carabidae) and spiders (Araneida) in salt marshes at the North and the Baltic Sea and the impact of the predicted sea level increase

The ground beetles and spiders of two salt marshes at the German Northand Baltic Sea coast were investigated by pitfall traps in 1997 and 1998. Whilethe sites at the North Sea coast are tidal salt marshes, the salt marshes at theBaltic Sea are not influenced by tides. Pitfall traps were installed in agradient from 20 to 150 cm above MHT (mean high tide: 157cm + NN, NN: 500 cm above 0 at Amsterdam gauge) atthe North Sea coast or NN at the Baltic Sea coast at six or sevensampling elevations, each with five replicates. Conductivity, water content, organic substance,frequency or duration of floodings, sand content and pH of the soil weredetermined. The flooding regime is the major factor controlling the zonation ofinvertebrates. Two and three invertebrate assemblages at the North and BalticSea, respectively, were distinguished. These corresponded well with thevegetational zones. The border between the two zones was at 60–80cm above MHT at the North Sea. The three zones at the Baltic Seaextended between 20 and 30 cm, 40 to 80 cm and 100 to150 cm above NN. The elevation of the mean abundance of speciesabove MHT or NN was calculated. A tide simulation experiment resulted in ashifting population and in an increasing activity under a tidal regime aspredicted for the global climate change conditions in 2050. From the actualelevation of the mean abundance, the habitat size of salt marsh species wascalculated for a moderate and worse scenario of global climate change. Habitatreduction becomes highest for species of the lower salt marsh zone. Under worseconditions the gradiental length of habitat will only amount to a maximum of 20m at the slopes of the dikes.     

Effect of residual stress and heterogeneity on circumferential stress in the arterial wall

Quantifying the stress distribution through the arterial wall is essential to studies of arterial growth and disease. Previous studies have shown that both residual stress, as measured by opening angle, and differing material properties for the media-intima and the adventitial layers affect the transmural circumferential stress (sigma theta) distribution. Because a lack of comprehensive data on a single species and artery has led to combinations from multiple sources, this study determined the sensitivity of sigma theta to published variations in both opening angle and layer thickness data. We fit material properties to previously published experimental data for pressure-diameter relations and opening angles of rabbit carotid artery, and predicted sigma theta through the arterial wall at physiologic conditions. Using a one-layer model, the ratio of sigma theta at the internal wall to the mean sigma theta decreased from 2.34 to 0.98 as the opening angle increased from 60 to 130 deg. In a two-layer model using a 95 deg opening angle, mean sigma theta in the adventitia increased (112 percent for 25 percent adventitia) and mean sigma theta in the media decreased (47 percent for 25 percent adventitia). These results suggest that both residual stress and wall layers have important effects on transmural stress distribution. Thus, experimental measurements of loading curves, opening angles, and wall composition from the same species and artery are needed to accurately predict the transmural stress distribution in the arterial wall.     

Tracing and interpreting fine-scale human impact in northern Fennoscandia with the aid of modern pollen analogues

A set of moss samples (n=58) was collected and analysed to obtain modern pollen analogues for both natural and human induced vegetation types in northern Fennoscandia. Vegetation types with settlement, trampling and grazing were selected from the different latitudinal forest zones (birch, pine, and mixed forest). The moss species varied between samples but the size of the sample was kept constant. Numerical analyses such as Redundancy Analysis (RDA) indicate that in northern areas human presence is mainly visible as increased values of Gramineae, Rumex acetosa/acetosella, R. obtusifolius, Solidago-type and Achillea-type pollen. Partial RDA further reveals that settlement is strongly correlated with high values of Rumex acetosa/acetosella, whereas trampled sites are significantly characterised by Gramineae and, to a smaller extent, Cyperaceae pollen. It is therefore possible to distinguish the impact of different types of human activities on vegetation on the basis of the pollen spectra. Later, these data will be used to interpret the presence, nature, and duration of human impact from fossil records from the same area. Received June 6, 2001 / Accepted December 12, 2001     

Floristic heterogeneity between forested sites in Kibale National Park, Uganda: insights into the fine-scale determinants of density in a large-bodied frugivorous primate

1.  Despite a long history of research on the influence of fruit availability on the population density of large-bodied vertebrate frugivores, operational understanding of the factors regulating density in these taxa remains elusive. We propose that fruit resources can be distinguished from one another on the basis of their functional role for the animals in question, and that such a classification system can aid in identifying the most influential determinants of frugivore density.
2.  We compared the availability of several resource classes between two sites in Kibale National Park, Uganda separated by only 12 km yet differing threefold in density of chimpanzees ( Pan troglodytes ).
3.  We categorized plant species used for fruit by chimpanzees according to their availability relative to habitat-wide fruit productivity, and by their tendency towards inter-individual fruiting synchrony. We predicted that the site of high chimpanzee density would support a higher density of food plant species tending to produce crops during periods of high habitat-wide productivity [high fruit abundance (HFA foods)] and of those tending to fruit synchronously among individuals during times of low habitat-wide availability (sLFA foods). The first food class should provide chimpanzees with a high nutrient density (and thus promote population growth), whereas the second should provide stable subsistence during lean periods and thus a temporally consistent resource base.
4.  Counter to our prediction, only sLFA resources were more abundant at the site of high chimpanzee density than at the site of low density. We suggest that sLFA resources are most important in influencing density of large-bodied frugivores.     

Osmotic relations of the drought-tolerant shrub Artemisia tridentata in response to water stress

Turgor maintenance, solute content and recovery from water stress were examined in the drought-tolerant shrub Artemisia tridentata. Predawn water potentials of shrubs receiving supplemental water remained above ?2 MPa throughout summer, while predawn water potentials of untreated shrubs decreased to ?5 MPa. Osmotic potentials decreased in conjunction with water potentials maintaining turgor pressures above 0 MPa. The decreases in osmotic potentials were not the result of osmotic adjustment (i.e. solute accumulation). Leaf solute contents decreased during drought, but leaf water volumes decreased more than 75% from spring to summer, thereby passively concentrating solutes within the leaves. The maintenance of positive turgor pressures despite decreases in leaf water volumes is consistent with other studies of species with elastic cell walls. Inorganic ion, organic acid, and carbohydrate contents of leaves declined during drought. The only solutes accumulating in leaves of A. tridentata with water stress were proline and a cyclitol, both considered compatible solutes. Total and osmotic potentials recovered rapidly following rewatering of shrubs; solute contents did not change except for a decrease in proline. Maintaining turgor through the passive concentration of solutes may be advantageous compared to synthesis of new solutes for osmotic adjustment in arid environments.     

Man and the last great wilderness: human impact on the deep sea

The deep sea, the largest ecosystem on Earth and one of the least studied, harbours high biodiversity and provides a wealth of resources. Although humans have used the oceans for millennia, technological developments now allow exploitation of fisheries resources, hydrocarbons and minerals below 2000 m depth. The remoteness of the deep seafloor has promoted the disposal of residues and litter. Ocean acidification and climate change now bring a new dimension of global effects. Thus the challenges facing the deep sea are large and accelerating, providing a new imperative for the science community, industry and national and international organizations to work together to develop successful exploitation management and conservation of the deep-sea ecosystem. This paper provides scientific expert judgement and a semi-quantitative analysis of past, present and future impacts of human-related activities on global deep-sea habitats within three categories: disposal, exploitation and climate change. The analysis is the result of a Census of Marine Life--SYNDEEP workshop (September 2008). A detailed review of known impacts and their effects is provided. The analysis shows how, in recent decades, the most significant anthropogenic activities that affect the deep sea have evolved from mainly disposal (past) to exploitation (present). We predict that from now and into the future, increases in atmospheric CO(2) and facets and consequences of climate change will have the most impact on deep-sea habitats and their fauna. Synergies between different anthropogenic pressures and associated effects are discussed, indicating that most synergies are related to increased atmospheric CO(2) and climate change effects. We identify deep-sea ecosystems we believe are at higher risk from human impacts in the near future: benthic communities on sedimentary upper slopes, cold-water corals, canyon benthic communities and seamount pelagic and benthic communities. We finalise this review with a short discussion on protection and management methods.     

Vertical structure of seed banks and the impact of depth of burial on recruitment in two temporary marshes

The structure of the seed bank (including Chara oospores), in relation to depth within the sediment and disturbance, was studied in two Rhône delta temporary marshes for two years. The seeds of all species were concentrated in the top 2 cm of sediment with very low numbers beeing found below 4 cm. When an exclosure eliminated disturbances of the sediment by animals, the vertical repartition of seeds at site 2 was more pronounced than outside the exclosure.In experiment 1, the emergence capacity of seeds from different depths and buried under layers of sterile equivalent to those in the field was measured. Depending of the species, 22 to 98% of the seeds germinated from unburied seeds in the top 2 cm. Only 1% of the oospores of Chara (from site 2) at 2 to 4 cm depth in the sediment emerged.In experiment 2, surface seed bank samples were placed under 0, 2 or 4 cm sterile sediment depth. The samples contained numerous recent seeds and the emergence percentage reached 41% (for Ruppia maritima). Only the seeds of Zannichellia spp failed to germinate from a depth of 2 cm or more. The emergence percentage from 2 cm depth or more was always lower than at the surface. These experiments showed that both burial and ageing of seeds decrease germination capacity.The majority of the active seeds located at the surface germinate when the marsh is flooded. Seeds located between 2 and 4 cm can be brought back to the surface by disturbances and play the role of a reserve involved in maintenance of populations that go without seed production for one or some years.     

Immunocytochemical evidence suggesting heterogeneity in the population of sea urchin egg cortical granules

Unfertilized eggs of many species of animals contain cortical granules, which are specialized secretory granules that upon fertilization release their contents from the egg. The unfertilized eggs of the sea urchin, Strongylocentrotus purpuratus, contain cortical granules that all display an identical and elaborate internal morphology. It has been assumed that they all contain identical components. In this report we present immunocytochemical data which indicate that the cortical granule population of S. purpuratus eggs is heterogeneous. Two monoclonal antibodies are shown to react to the spiral lamellae region of approximately 20% of the cortical granules, implying that the contents of the reactive granules differ from the contents of the majority of the population. An egg protein of greater than 320 kDa is recognized by the antibody. These antibodies also stain a 130-kDa protein expressed on the surface of primary mesenchyme cells in later development. Both antibodies recognize a post-translational modification of this protein. This suggests that an antigenically similar epitope is present both on the 130-kDa primary mesenchyme cell-specific protein and in the cortical granules. To determine if the primary mesenchyme and cortical granule proteins are related, a fusion protein antibody specific for a region of the 130-kDa protein was used to stain unfertilized eggs. This antibody did not stain cortical granules. Thus, 20% of the cortical granules contain a molecule that has an epitope antigenically similar to the post-translational modification recognized in primary mesenchyme cells by the monoclonal antibodies.     

Non-structural carbohydrate accumulation and use in an understorey rain-forest shrub and relevance for the impact of leaf herbivory

1. Piper arieianum, an evergreen, understorey shrub of lowland moist forests of Central and South America, exhibits marked seasonal variation in reproductive activity even though climatic variation is low at the study site. Despite a lack of climatic seasonality, previous experimental leaf removal suggested that carbohydrate accumulation is seasonal, occurring prior to flowering.
2. We first tested the hypothesis that carbohydrates necessary for reproduction are accumulated prior to flowering, rather than during or after. By measuring non-structural carbohydrate production in the form of glucose and starch we found that the concentration of these reserves is greatest 1–3months before flowering, decreasing by 50% during peak fruit maturation.
3. The hypothesis that reproduction was the cause of this decrease in carbohydrate reserves was then tested by comparing reserves in plants that were prevented from flowering with those that flowered and produced fruit naturally. As predicted, reserves declined more in flowering than in non-flowering plants. A smaller decline in reserves of non-flowering plants was accompanied by greater stem and leaf production, suggesting that stored carbohydrates are also required for growth.
4. Because concentrations of non-structural carbohydrates were similar in roots, stems and leaves, and because the greatest amount of plant biomass is in stems for plants of a range of sizes, stems appear to be the main storage site of carbohydrate reserves in this plant species.
5. These results, together with previous studies, demonstrate that the impact of leaf herbivory on seed production in P. arieianum depends on the timing of that herbivory relative to the accumulation and use of non-structural carbohydrates.