Status of coastal dune communities invaded by Chrysanthemoides monilifera

It has often been assumed that the introduced species Chrysanthemoides monilifera is displacing native species in invaded coastal communities in Australia but no direct evidence has been presented that this displacement is occurring and, if so, to what extent. At an invaded site near Moruya, New South Wales, we found strong negative correlations between the presence (measured as cover, frequency or volume of shrub) of C. monilifera and Acacia longifolia, the hitherto dominant native species, and of C. monilifera and Lomandra longifolia. The negative correlations are partly accounted for by differences in the amounts of each species on different zones on the dune, but the negative correlations remain even within these zones. Also a comparison with uninvaded sites indicates that C. monilifera is displacing A. longifolia from the fore-dune and mid-dune but is having less effect in the swale. In addition, there is a trend towards reduced seed production of A. longifolia when growing in proximity to C. monilifera. There is no significant difference in the presence of L. longifolia in the uninvaded and invaded sites. In invaded sites, significantly fewer C. monilifera seedlings occur in the presence of L. longifolia. Thus it appears that the observed negative associations arise from different causes. Chrysanthemoides monilifera is displacing the structurally similar, and previously dominant A. longifolia, while the tussock-forming L. longifolia is locally hindering the invasion.     

Selection of Cassida spp. from southern Africa for the biological control of Chrysanthemoides monilifera in Australia

Four species of tortoise beetle (Chrysomelidae, Cassidinae) (Cassida spatiosa Spaeth and three undescribed Cassida species, labelled 1. 2 and 3) from South Africa, were assessed as potential biological control agents against Chrysanthemoides monilifera monilifera (L.) T. Norl. and C. m. rotundata (DC.) T. Norl. These southern African plants are important weeds of conservation areas in Australia. In South Africa, Cassida spatiosa was found feeding on C monilifera subcanescens (DC.) T. Norl., Cassida sp. 1 on Chrysanthemoides monilifera pisifera (L.) T. Norl. and C m. monilifera, Cassida sp. 2 on C m. pisifera and Chrysanthemoides incana (Burm.f.) T. Norl. and Cassida sp. 3 on C. m. rotundata. The life cycle of each species was completed in about three months on the leaves of the host. Cassida species 1, 2 and 3 were tested against eight species of plant and only fed and completed development on Chrysanthemoides species and the related species. Calendula officinalis L. Cassida species 1 and 3 showed no preference between Calendula officinalis and Chrysanthemoides species for oviposition. Cassida species 3 was selected for further assessment as a biological control agent based on match to the host subspecies and target climate in Australia.     

Preliminary host specificity testing of Endophyllum osteospermi (Uredinales, Pucciniaceae), a biological control agent against Chrysanthemoides monilifera ssp. monilifera

Chrysanthemoides monilifera ssp. monilifera, indigenous to the Western Cape Province of South Africa, is a serious invader of native vegetation in south-eastern Australia. The rust fungus Endophyllum osteospermi causes witches' brooms on C. monilifera ssp. monilifera in South Africa, and is associated with a reduction in growth and seed production of its host under natural conditions, as well as mortality of severely infected bushes. This rust fungus is considered to be a potential biological control agent for use against C. monilifera ssp. monilifera in Australia. Endophyllum osteospermi has a long latent period, typically between 6 and 24 months between infection and the initiation of witches' brooms. This long latent period makes the logistics of doing traditional host specificity testing, in which all test plant species are inoculated and observed for symptom development, unfeasible for this rust fungus. Germination of aecidioid teliospores and penetration by basidiospores were observed on the surface of excised leaves of 32 test plant species at 4 days after inoculation, and compared to that on C. monilifera ssp. monilifera. Germinating aecidioid teliospores aborted on 14 test plant species, whilst no penetration was attempted on a further 12 test plant species. Penetration only occurred on nine of the 32 test plant species, in addition to C. monilifera ssp. monilifera. Inoculating whole plants of nine selected test plant species confirmed the above results. Therefore, only the test plant species in which penetration occurred, or at least was attempted, need to undergo comprehensive host specificity testing. Pending these results, E. osteospermi may be suitable for release in Australia for the biological control of C. monilifera ssp. monilifera.     

Identification and Phytotoxicity Assessment of Phenolic Compounds in Chrysanthemoides monilifera subsp. monilifera (Boneseed)

Chrysanthemoides monilifera subsp. monilifera (boneseed), a weed of national significance in Australia, threatens indigenous species and crop production through allelopathy. We aimed to identify phenolic compounds produced by boneseed and to assess their phytotoxicity on native species. Phenolic compounds in water and methanol extracts, and in decomposed litter-mediated soil leachate were identified using HPLC, and phytotoxicity of identified phenolics was assessed (repeatedly) through a standard germination bioassay on native Isotoma axillaris. The impact of boneseed litter on native Xerochrysum bracteatum was evaluated using field soil in a greenhouse. Collectively, we found the highest quantity of phenolic compounds in boneseed litter followed by leaf, root and stem. Quantity varied with extraction media. The rank of phenolics concentration in boneseed was in the order of ferulic acid > phloridzin > catechin > p-coumaric acid and they inhibited germination of I. axillaris with the rank of ferulic acid > catechin > phloridzin > p-coumaric acid. Synergistic effects were more severe compared to individual phenolics. The litter-mediated soil leachate (collected after15 days) exhibited strong phytotoxicity to I. axillaris despite the level of phenolic compounds in the decomposed leachate being decreased significantly compared with their initial level. This suggests the presence of other unidentified allelochemicals that individually or synergistically contributed to the phytotoxicity. Further, the dose response phytotoxic impacts exhibited by the boneseed litter-mediated soil to native X. bracteatum in a more naturalistic greenhouse experiment might ensure the potential allelopathy of other chemical compounds in the boneseed invasion. The reduction of leaf relative water content and chlorophyll level in X. bracteatum suggest possible mechanisms underpinning plant growth inhibition caused by boneseed litter allelopathy. The presence of a substantial quantity of free proline in the target species also suggests that the plant was in a stressed condition due to litter allelopathy. These findings are important for better understanding the invasive potential of boneseed and in devising control strategies.     

Impact of the biological control agent Mesoclanis polana (Tephritidae) on bitou bush (Chrysanthemoides monilifera subsp. rotundata) in eastern Australia

The seed fly Mesoclanis polana (Diptera: Tephritidae) was released in Australia in 1996. Its impact on seed production of bitou bush (Chrysanthemoides monilifera subsp. rotundata) was monitored at eight sites along the New South Wales coast from 1996 to 2004. Peak flowerhead production occurred in autumn (March to May); therefore, samples collected in May of each year were used to compare abundance and impact of M. polana across sites and between years. Latitude had a significant effect on abundance and impact of M. polana. By May 2004, 99.6% of flowerheads at the five most northern sites contained at least one egg, while 64% of flowerheads from the two most southern sites contained at least one egg. In May 2004, mean numbers of M. polana eggs per flowerhead were between 13 and 17 at four of the six northern sites and below two at the two southern sites. In May 2004, average seed destruction by M. polana was 58% at the four most northern sites and 11% at the two most southern sites. The highest level of seed destruction recorded was 86% at Kingscliff in May 2003. Percentage seed destruction in May provided a reasonable estimate of seed destruction for the whole year. Parasitoids were found throughout the range of M. polana in Australia. Their attack rate on M. polana was less than 10% at all sites, except at the two most northern sites where parasitism of up to 27% was recorded. Parasitism results are compared with predictions made following an earlier study. The release of Mesoclanis magnipalpis, a species suited to cooler environments, is recommended for the southern range of C. monilifera subsp. rotundata.     

Evaluation and host specificity of two seed flies Mesoclanis polana and M. magnipalpis (Diptera: Tephritidae): biological control agents for Chrysanthemoides monilifera (Asteraceae) in Australia

Larvae of the South African tephritid flies Mesoclanis polana Munro and M. magnipalpis Bezzi feed in the developing seeds of Chrysanthemoides monilifera. Host specificity evaluation using 109 plant species from 25 families indicated that complete development was restricted to their natural host C. monilifera. Minor feeding and limited development was detected on 18 species, but was of no ecological or economic significance. Mesoclanis polana and M. magnipalpis have been released in Australia and M. polana has established and dispersed widely. Mesoclanis magnipalpis has not yet become naturalized. Parasitism of M. polana in Australia by several species of Hymenoptera has been detected, but is not expected to limit the establishment and impact of these flies.     

Competition strength of two significant invasive species in coastal dunes

To investigate the effect of increased nutrient availability on competition amongst invasive and native plants, I measured changes in above and below ground biomass of Chrysanthemoides monilifera spp. rotundata (bitou bush) and Asparagus aethiopicus (asparagus fern) competing with two native species, Banksia integrifolia and Ficinia nodosa, under high- and low-nutrient regimes. Bitou bush, as a primary invader, was competitive under all conditions lowering the growth of native species in both high and low nutrients. Asparagus fern as a secondary invader, did not influence growth of native species but responded, like bitou bush, to high nutrients. Native species were generally negatively affected by increases in nutrients. With bitou bush soils often providing higher nutrients, the chance of secondary invasion by asparagus fern is more likely, although asparagus fern is unlikely to invade low nutrient soils quickly. The invasive species, therefore, differed in their competitive ability in these coastal dune communities.     

Evidence for allelopathy as a mechanism of community composition change by an invasive exotic shrub, Chrysanthemoides monilifera spp. rotundata

Chemical interference is increasingly suggested as a mechanism facilitating exotic plant invasion and plant community composition. In order to explore this further, we employed a comprehensive extract-bioassay technique that facilitated detection and demarcation of phytotoxicity, direct allelopathy and indirect allelopathy of bitou bush (Chrysanthemoides monilifera spp. rotundata) compared to an indigenous dominant of the invaded system, acacia (Acacia longifolia var. sophorae). Extracts of the leaves and roots of both species exhibited phytotoxic effects against five indigenous plant species. Evidence for allelopathy between co-evolved indigenous plants was detected between acacia and Isolepis nodosa. Allelopathy between bitou bush and four indigenous plant species was also detected. Therefore we propose that both the acacia and bitou bush have the potential to chemically inhibit the establishment of indigenous plants. Eventual dominance of bitou bush is predicted, however, based on more ubiquitous effects on seedling growth.     

The Impact of the Weed Chrysanthemoides monilifera ssp. rotundata on Coastal Leaf Litter Invertebrates

In coastal areas of Australia, there are extensive infestations of the environmental weed Chrysanthemoides monilifera ssp. rotundata (bitou bush). This study looked at the impact of long-term infestations on the abundance and assemblage composition of leaf litter invertebrates. Assemblages were compared in weed infested and native shrublands along the New South Wales coastline over 12 months. The total abundance was not significantly reduced in the weedy habitat but the abundance of mites, thrips, spiders, ants, and centipedes was reduced at many sites. The invertebrate assemblages also differed between habitats, with the C. monilifera supporting a lower diversity of beetles. However, the millipedes, amphipods, earthworms, pseudoscorpions and isopods appeared to respond positively to the invasion, occurring in higher abundance and detected more frequently in the weedy areas. This has been partially attributed to a change in microclimate within the C. monilifera infestations. It is generally moister and darker, which these invertebrates tend to prefer. Secondly, C. monilifera produces less leaf litter of higher quality, and possibly higher palatability than the native sclerophyllous vegetation, which may encourage species that consume litter.     

Ecological niche modelling of an invasive alien plant and its potential biological control agents

Invasive alien plants are of concern in South Africa. Pompom weed (Campuloclinium macrocephalum) is currently invading the Grassland and Savannah biomes of South Africa and is likely to continue spreading in the southern African sub- region. Two possible biological control agents (Liothrips tractabilis and Cochylis campuloclinium) have been identified for control of pompom weed. We used ecological niche modelling to predict which areas in southern Africa are likely to be suitable for pompom weed and the two potential biological control agents. The overlap between areas predicted to be highly suitable for pompom weed and areas suitable for the biological control agents was assessed. Methods of reducing sampling bias in a data set used for calibrating models were also compared. Finally, the performance of models calibrated using only native range data, only invaded range data and both were also compared. Models indicate that pompom weed is likely to spread across a greater region of southern Africa than it currently occupies, with the Savannah and Grassland biomes being at greatest risk of invasion. Poor overlap was found between the areas predicted to be highly suitable for pompom weed and those areas predicted to be suitable for the biological control agents. However, models of the potential distribution of the biological control agents are interpreted with caution due to the very small sample size of the data set used to calibrate the models. Models calibrated using both native range and invaded range data were found to perform best whilst models calibrated using only native range data performed the worst. There was little difference found between models that were calibrated using spatially reduced (selecting only one record per 30 min grid cell) and randomly reduced (randomly selecting 50% of available records) biased data sets.     

Development of a Pesticide Exclusion Technique for Assessing the Impact of Biological Control Agents for Chrysanthemoides monilifera

Seven biological control agents have been released on the environmental weed Chrysanthemoides monilifera in Australia and two are widely established on infestations of C. monilifera subsp. rotundata in New South Wales. Five pesticides were screened for their impact on seed production of C. monilifera and two on the survival of the agent Comostolopsis germana, a shoot tip-feeding lepidopteran. The insecticides carbaryl, carbofuran, dimethoate, fluvalinate and the fungicide benomyl had no significant effect on seed production of C. monilifera when applied over a period of four months. Numbers of C. germana larvae were readily suppressed after three months by applications of fluvalinate or a mixture of carbaryl, carbofuran and dimethoate. Shoot growth was not affected by applications of carbaryl, dimethoate and benomyl. It is concluded that exclusion by pesticides of biological control agents is a valid method of measuring the impact of agents on C. monilifera.     

Soil pH and species diversity in coastal dunes

Soil pH was measured at two different spatial scales in coastal dunes on Norderney, North Sea, and in Mecklenburg-Vorpommern, Baltic Sea, Germany. Relationships between the variability in soil pH, species richness and species diversity are presented. Species richness and diversity were highest in grey dunes, where soil pH was at intermediate levels; both variables were lower in yellow and brown dunes. The variability in pH increased with increasing species diversity and also with scale. Overall, soil pH variability decreased with increasing vegetation cover. The lowest pH heterogeneity was found in heath dominated by Empetrum nigrum L. and grey dunes dominated by Campylopus introflexus (Hedw.) Brid. Increasing abundance of dominant species and decreasing species diversity of vegetation apparently reduces soil heterogeneity. Decreasing species diversity of vegetation is likely to explain decreasing variability in soil pH.     

Resource allocation patterns in Spinifex sericeus R.Br.: A dioecious perennial grass of coastal sand dunes

The resource allocation patterns within populations of Spinifex sericeus R.Br, were studied on the mid north coast of NSW. Sand burial and nutrient availability were two major factors influencing the allocation of resources to roots, culms, tillers and inflorescences, resulting in different growth forms in different sections of the dune. The data suggest that pollen availability may affect allocation to seeds late in the flowering season. Spinifex sericeus was found to have a greater below- than above-ground biomass, especially on the crest of the incipient foredune where sand accretion was high. The ability of culms of S. sericeus to grow upwards at a rate that generally prevents complete burial of the tillers is one mechanism which allows S. sericeus to vary resource allocation patterns in response to a varying environment. Differences in the inflorescence characteristics of males and females indicate their different functions. The much greater floret number of males indicated the importance of pollen dispersal. Female inflorescences consisted mainly of structures other than florets and suggested the importance of the female inflorescence in seed dispersal. Addition of nitrogen and phosphorus increased the number of spikelets and seeds but not the proportion of spikelets which produced seed.     

Identification of volatile compounds released by roots of an invasive plant, bitou bush (Chrysanthemoides monilifera spp. rotundata), and their inhibition of native seedling growth

Allelopathy has been suggested as a mechanism promoting the monoculture formation of some invasive exotic plants. Previous studies have shown that hydrophobic extracts of the roots and soil of exotic bitou bush (Chrysanthemoides monilifera spp. rotundata (DC.) T. Norl.) inhibited the seedling growth of five Australian native plants, including the dominant acacia (Acacia longifolia var. sophorae (Labill.) F. Muell.). Based on this finding, we compared the hydrophobic root and soil chemical profiles of bitou bush and acacia to determine whether bitou bush roots release allelopathic compounds that are novel to the invaded system. We detected three compounds that were exclusive to the bitou bush root and soil, and seven compounds that were common to the bitou bush and acacia roots but only present in the bitou bush soil. The compounds unique to the bitou bush invaded soil were all sesqui- and diterpenes. Several of these compounds were found to inhibit the seedling growth of a native sedge, Isolepis nodosa (Rott.) R. Br. Of particular interest are the sesquiterpenes: β-maaliene, α-isocomene, β-isocomene, δ-cadinene, 5-hydroxycalamenene and 5-methoxycalamenene which were found in high concentrations in the bitou bush root and soil extracts and exhibited phytotoxic activity. Therefore, we present evidence to suggest that bitou bush exudes low molecular weight volatile compounds into the soil which inhibit native plant seedling growth. The reduced establishment of native plants via allelopathy is likely to create space and contribute to the invasion of bitou bush on the eastern Australian coast.     

Soil‐mediated effects on germination and seedling growth of coastal wattle (Acacia sophorae) by the environmental weed,bitou bush (Chrysanthemoides monilifera ssp. rotundata)

Bitou bush (Chrysanthemoides monilifera ssp. rotundata; Asteraceae) is a major woody weed that competes with the native legume Acacia sophorae in coastal ecosystems of eastern Australia. Three glasshouse experiments examined whether litter or soil from beneath bitou bush or Acacia plants could influence seed germination and seedling growth of A. sophorae. The presence of litter decreased seed germinability and this effect was greater for bitou bush litter than for Acacia litter. Shoot growth was increased by the addition of Rhizobium after 40 days, irrespective of soil type. After 78 days, shoot and root biomass were significantly lower for seedlings grown in bitou bush soil than for those grown in Acacia soil. There was a non‐significant trend towards a lower median population of Rhizobium in the soil beneath bitou bush than in that beneath Acacia. The results demonstrated a slight effect of bitou bush on the growth of A. sophorae, which could, however, be overshadowed by the judicious use of herbicides or fire for weed control and revegetation.     

The vegetation of road verges in the coastal dunes of the Netherlands

The vegetation of road verges in the coastal dune area of the Netherlands was surveyed phytosociologically. Nine plant communities have been distinguished. They belong to theAmmophiletea, Koelerio-Corynephoretea, Molinio-Arrhenatheretea, Plantaginetea majoris andArtemisietea vulgaris. Most plant communities appear to be, fragmentarily developed and have thus been described by the so-called deductive method. Vegetation in road verges often appears to be heterogeneous. Species composition is influenced by man, distance from the sea, fertility and moisture content of the soil. It is clearly correlated with total nitrogen, magnesium and organic matter. Therophytes are preponderant; these and sclerophytes indicate the dry environment. The majority of the species are apophytes.     

Seasonality of mycorrhizae in coastal sand dunes of Baja California

 Populations of arbuscular mycorrhizal fungi were estimated from spores associated with seven plant species in coastal dunes at El Socorro, near Ensenada, Baja California, during six months in 1992. The seasonal patterns of percent root colonization were also described in the same species during the wet season (January–March) and the dry season (April–July). Comparisons were made between the pioneer species (Abronia maritima) in the mobile dunes and six species (Abronia umbellata, Atriplex julacea, Camissonia californica, Haplopappus venetus, Helianthus niveus and Lotus spp.) in the fixed dunes. Mycorrhizal colonization in Abronia maritima was slight (<1%) and we observed few spores (<1/g soil). All of the species in the fixed dune formed mycorrhizae with up to 80% colonization in early summer, and no more than 4 spores/g soil by late summer. The highest percentages of total colonization and abundance of spores did not coincide temporally for any of the seven species, but the percentages were higher in summer than in spring. Arbuscules were more abundant when the soil was moist, and vesicles more abundant when it was dry. Accepted: 16 August 1995     

Analysis of the specificity of three root-feeders towards grasses in coastal dunes

Among the root-feeding nematodes that accumulate in the rhizosphere of grasses in European dunes, the genus Pratylenchus is of special relevance given its diversity and distribution. Although different species of Pratylenchus have been reported in dune grasses, the specificity towards dune plants, a fundamental aspect of the biology of the species, has hitherto not been studied. Two inoculation experiments using different combinations of grasses and nematodes were performed. The multiplication and the effect on plant growth of P. dunensis and P. brzeskii, two species which only occur in dune areas was compared with that of P. penetrans, a broad host-range species. The three Pratylenchus spp. could multiply under all hosts; however, there was a clear host-dependent response. The species-specific response observed might account for the shift of Pratylenchus spp. detected in the field. Although, a negative effect on the growth of A. arenaria was demonstrated for the three nematode species, different densities were needed to observe the same effects in plant biomass which point at nematode-specific tolerance. While the typical dune species needed very high densities to produce damage, P. penetrans needed very few specimens. The results obtained indicate that species with similar feeding adaptations show very different multiplication abilities on co-occurring hosts, an aspect that is usually overlooked for belowground herbivores in natural systems. The obtained results might suggest a coevolutionary relationship between specific nematode species and Ammophila arenaria.