Impact of alien trees on mammal distributions along an ephemeral river in the Namib Desert

Ephemeral rivers and the vegetation they support have numerous ecological and economic values to the mammals and people who rely on these systems. Yet, these crucial environments are believed to be threatened by exotic plant invasion. In Africa, invasive trees of the genus Prosopis have detrimental effects on native vegetation, bird and dung beetle communities; however, to date, there is no evidence that Prosopis establishment has affected indigenous wild mammalian distribution and ecology in its introduced range. Using a combination of camera traps and vegetation surveys, we tested the hypothesis that Prosopis invasion has a negative impact on the mammals of the ephemeral Swakop River in Namibia by reducing mammal species richness and species occupancies. Prosopis was found to have no negative impact on species richness; however, evidence for species‐specific responses to Prosopis abundance was found. This is the first study to confirm an impact of Prosopis on sub‐Saharan African mammals, providing a foundation for future research and the development of appropriate management policy.     

The present status and potential distribution of relict populations of Aesculus hippocastanum L. in Greece and the diverse infestation by Cameraria ohridella Deschka & Dimić.

Aesculus hippocastanum is a well-known species, which is popular because of its ornamental value. However, data on the demographic structure and potential distribution of A. hippocastanum are limited. The invasion of Cameraria ohridella into Europe has harmed those trees growing in artificial sites, but the presence of this insect in natural stands has been little studied. Here we aimed to investigate the demographic structure infestation level of natural populations of horse-chestnut. Additionally, Maxent modelling was used to predict the potential range of A. hippocastanum, based on the localities available in the literature. Field data analysis indicated that natural populations of A. hippocastanum are mostly found nearby mountain streams. The populations showed a diverse height structure and large numbers of seedlings, which indicate high population dynamics. The level of infestation by C. ohridella varied greatly and correlated with altitude. Secondary infestation might explain this infestation variability in some natural populations. Other hypotheses, such as environmental resistance factors or different genetic variability, are also discussed. By spatial distribution modelling, we found that the precipitation of the coldest and warmest quarters, as well as altitude, are important factors influencing the potential distribution of A. hippocastanum.     

Prediction of Prosopis species invasion in Kenya using geographical information system techniques

Tree species from Prosopis genus were widely planted for rehabilitation of degraded drylands of Kenya. However, they have invaded riverine ecosystems where they cause negative socio‐economic and ecological impacts. GIS was used to estimate the reverine area threatened by Prosopis invasion in Kenya. Landsat satellite images, field surveys and past studies were also used to assess the resulting potential ecological impacts in the Turkwel ecosystem in Kenya. The study revealed that 3.0 to 27.7 million hectares are threatened by invasion, based on documented riverine forests width of 0.5–3 km. Image analysis showed that 34% of the sites under positive change were invaded, with most invasions occurring in natural forests and abandoned farms. Prosopis had overall occurrence of 39% in all the sampled sites in 2007, in contrast to 0% in 1990 that was reported in an earlier study. In these areas, Acacia tortilis occurrence dropped from 81% in 1990 to 43% in 2007, suggesting that Prosopis could be displacing it. Utilization of Prosopis for fodder, fuel wood and pods for animal feeds is recommended as a management tool to reverse the trend. The methods used in this study are also recommended for invasion prediction and management in other similar ecosystems.     

Tree-shrub interactions in a subtropical savanna parkland: competition or facilitation?

Abstract. Prosopis glandulosa var. glandulosa has played a central role in the encroachment of woody plants in southern Texas, grasslands and savannas by acting as a nurse plant for various shrubs that establish in its understory. To test for continued facilitation of established understory shrubs by Prosopis and to determine if established shrubs compete with the Prosopis nucleus, selective removal experiments were conducted and monitored over a 2–5 yr period. Short-term (1–3 days) and long-term (2 yr) growth and physiological activities (midday net photosynthesis and leaf/shoot water potential) of two common understory shrubs, Zanthoxylum fagara and Berberis trifoliolata, growing with Prosopis, were generally comparable to those of individuals occurring in clusters where Prosopis was removed. Shrubs growing with an intact Prosopis occasionally showed significantly higher leaf-[N] and pre-dawn water potentials than those in clusters lacking a live Prosopis, especially under drought conditions; however, these differences did not translate into greater midday leaf gas exchange or shoot growth. By comparison, removal of understory shrubs elicited large increases in Prosopis net photosynthesis, annual trunk growth in each of the 5 yr monitored, and seed pod production in three of the four years monitored. Seven of 26 Prosopis plants in experimental clusters with an intact understory died over a 5-yr period, compared to only two of the 26 plants in clusters with the cleared understory. Results indicate that (1) the founding overstory Prosopis plant may continue to facilitate understory shrubs following their establishment, but these beneficial effects appear to be small and transitory, and (2) the understory shrubs have a pronounced negative effect on Prosopis, such that competition between overstory and understory woody plants is strongly asymmetrical. These findings suggest that understory shrubs will likely persist despite changes in microclimate and soils (potentially) that occur after the Prosopis plant, which facilitated their ingress or establishment, has died. Soil resource depletion by shallow-rooted understory shrubs appears to be a primary factor contributing to the demise of the deeply rooted, overstory Prosopis plants, especially on upland sites with duplex soils where below-ground competition is accentuated.     

Direct measurement of denitrification in a Prosopis (Mesquite) dominated Sonoran Desert ecosystem

Summary Denitrification was directly measured using the acetylene inhibition technique in a Sonoran Desert ecosystem dominated by Prosopis glandulosa. Soil under Prosopis and from the unvegetated area between Prosopis was wetted with 50 mm of water and denitrification measured for 48 hours. The mean denitrification rate under Prosopis was 11.6 g N ha^-1h^-1 compared to only 0.2 g N ha^-1h^-1 away from Prosopis. The denitrification response to wetting was rapid and rates peaked about 24 h after water application.The much higher denitrification under Prosopis probably results from high available organic C under Prosopis, but other soil chemical and physical changes effected by Prosopis may influence denitrification rates. About 0.5 kg N ha^-1 of Prosopis cover may be lost from this ecosystem by denitrification after infrequent major rainfalls.     

Influence of an overstorey tree (Prosopis glandulosa) on associated shrubs in a savanna parkland: implications for patch dynamics

The arborescent legume, honey mesquite (Prosopis glandulosa), appears to play a central role in patch dynamics of southern Texas savannas by modifying soils and microclimate and by facilitating the ingress, establishment and/or growth of shrubs in its understorey. As an indirect test for the occurrence and persistence of facilitation in mature shrub clusters (patches), we examined the gas exchange, water relations and production of associated shrubs growing in patches where a Prosopis overstorey was present and in patches where Prosopis had succumbed to natural mortality. Surface (0–10 cm) soils associated with shrub patches were enriched in total [N] and [C] compared to soils of neighboring herbaceous zones. However, there were no detectable differences in soil [N] or [C] in patches with and without Prosopis. Foliar [N] and biomass of various shrub species were also statistically comparable for patches with and without Prosopis. These results are in accordance with other studies that indicate the nutrient legacy associated with Prosopis occupation of a patch may persist for decades after its demise. In comparison to plants growing in the absence of Prosopis, leaf water potentials (predawn and midday), and net photosynthesis and water vapor conductance (morning and midday) of outer-canopy sunlit leaves over an annual growth cycle were comparable for two common evergreen shrubs, Zanthoxylum fagara and Berberis trifoliolata, growing in patches with a live Prosopis. These findings indicate that the presence of Prosopis was not enhancing the growth or activity of mature understorey shrubs; facilitation may, therefore, be important only during early stages of cluster development. In addition, we found no indication that the loss of Prosopis has initiated a downward phase in a cyclic succession of patch initiation, growth and death. Rather, the understorey shrubs appear to be able to maintain growth and productivity in the absence of a Prosopis overstorey, and may, therefore, represent persistent components of woody patches on these savanna landscapes.     

Commercially Important Trees as Invasive Aliens – Towards Spatially Explicit Risk Assessment at a National Scale

Alien species that are desirable and commercially important in parts of the landscape, but damaging invaders in other parts, present a special challenge for managers, planners, and policy-makers. Objective methods are needed for identifying areas where control measures should be focussed. We analysed the distribution of forestry plantations and invasive (self-sown) stands of Acacia mearnsii and Pinus spp. in South Africa; these two taxa account for 60% of the area under commercial plantations and 54% of the area invaded by alien trees and shrubs. The distribution of commercial forestry plantations and invasive stands of these taxa were mapped and the data was digitised and stored as Geographic Information System (GIS) (Arc/Info) layers. A series of environmental parameters were derived from GIS layers of climate, topography, geology, land use, and natural vegetation. The current distribution of the two taxa was subdivided into three groups according to the degree of invasion, planting history and the precision of the data collection. We used regression-tree analysis to relate, for each taxon, the distribution of invasive stands with environmental variables, and to derive habitat suitability maps for future invasion. The current distribution of invasive stands in South Africa was largely influenced by climatic factors. At a national scale, the distribution of large commercial plantations was a poor predictor of areas invaded by both taxa. Using environmental factors identified by the regression trees, we found that 6.6% and 9.8% of natural habitats currently not invaded and untransformed by urbanisation or agriculture are suitable for invasion by Pinus spp. and A. mearnsii, respectively. We then derived guidelines for policy on alien plant management based on vegetation type, degree of transformation, extent of invasion, and the risk of future alien spread. These factors were used to identify demarcated areas where these alien species can be grown with little risk of invasions, and areas where special measures are needed to manage spread from plantations.     

Coastline ecosystems on Oahu,Hawaii

Summary Coatline ecosystems on Oahu, Hawaii were described in terms of vegetation composition and structure, and environmental components such as climate, physiography, exposure, substratum, and ground water. Vegetation patterns were related to these environmental components. Relevés were established in 22 study areas around Oahu to study the broader patterns of vegetation and environment, and 2 m wide belt-transects were laid out in 16 areas to study structural and floristic zonation from the ocean edge to the limit of the strand flora inland. Physiographic position, substrate type, wind exposure and climate (through climate-diagrams) were determined for each site. Soil parameters determined along the transects were salt concentration, pH, organic carbon, moisture equivalent, soil moisture at 15 atmospheres pressure, and available moisture.Three levels of pattern in vegetation distribution were inferred. First, there were similarities and differences between study areas in terms of vegetation composition and structure, and environmental components. Thirteen ecosystem-classes were recognized on the bases of dominant species and landform: Hibiscus ecosystem on beach flat, Scaevola on dunes, Scaevola on talus and alluvium, Scaevola on raised coral rock, Scaevola on rocky basalt coast, Chloris-Sida on talus, Prosopis on dunes, Prosopis on talus and alluvium, Prosopis on tuffaceous headland, Chloris-Prosopis on dune and clay flat complex, Batis on mud flats, Rhizophora on tidal flats, and Scirpus-Eichornia in fresh-water pond.A second level of pattern was that of zonation into physiognomic zones, and a third that of floristic zonation or change in species composition with distance from sea. Only seven ecosystems, representing five ecosystem-classes (Scaevola on dunes, Scaevola on coral rock, Prosopis on dunes, Prosopis on talus and alluvium, and Prosopis on tuffaccous headland), showed distinct physiognomic zones. But all emerged ecosystems showed floristic zonation.The distribution of the ecosystems coastwise is most broadly related to rainfall and drought patterns, to exposure to wind and surf, and to salinity of soil and water. Hibiscus ecosystems occur in wet, protected beach flats of terrigenous sand, found in windward coasts; Scaevola ecosystems in areas exposed to intense wind and salt spray; Prosopis ecosystems and their variant, Chloris-Prosopis ecosystem, in droughtly areas of the leeward coast; Chloris-Sida ecosystem on rocky soils in the same droughty climate; Batis ecosystems on mud flats of high soil- and ground-water salinity; Rhizophora ecosystems in protected shorelines subject to tides; and Scirpus-Eichornia ecosystem in still fresh water.Salt concentration decreased markedly with distance inland in fine-textured soils, but showed only slight increases or decreases in coarse-textured dune soils. Organic matter showed increases, with three transects registering increases of 15% organic C at peak portions. pH was very variable; alkaline values arising largely from a high proportion of calcium carbonate parent material or high salt content and more acid values arising from greater organic matter incorporation. The range of many of the strand species has been increasingly limited by direct disturbance and indirectly by the seaward encroachment of introduced Prosopis. But they are still persisting (in larger or smaller numbers) in Scaevola ecosystems.     

Woody plant invasion of grasslands: establishment of honey mesquite (Prosopis glandulosa var.glandulosa) on sites differing in herbaceous biomass and grazing history

Summary Emergence and survival of honey mesquite (Prosopis glandulosa var.glandulosa Torr.) seedlings was quantified on sites with contrasting grazing histories: long-term continuous grazing (LTG) and long-term protection (LTP) from grazing by cattle. On each site, different levels of heroaceous defoliation were imposed at monthly intervals (no defoliation=ND, moderate=MD and heavy=HD). The two weeks following seed dissemination appeared to be the most critical toProsopis establishment on LTP-ND plots. Openings in the herbaceous layer created by moderate defoliation of grasses on the LTP site increased germination and/or survival 7-to 8-fold during this period. However, increasing the degree of defoliation from moderate to heavy did not stimulate additional emergence on either the LTP or LTG site. Emergence from scarified seed placed in cattle dung (17 to 30%) was lower than that of bare seed placements in various microhabitats (43–60%). However, deposition of scarifiedProsopis seed in dung in conjunction with graminoid defoliation may be the most likely combination of events when livestock are present. Emergence from seeds transported into grasslands by other fauna likely would be low, unless seeds were deposited in areas where grasses had been defoliated.Prosopis survival was comparably high in dung and bare seed placements after one growing season. survival of seedlings present two weeks after seed dissemination ranged from 74 to 97% at the end of the second growing season. Seedling survival and shoot development (biomass, leaf area and height) were similar on LTP and LTG sites, regardless of the level of herbaceous defoliation or seed placement. In addition, the magnitude and patterns of net photosynthesis, stomatal conductance and xylem water potential were comparable among one-year-old seedtings on ND, MD and HD plots, even though differences in herbaceous species composition and above- and below-ground biomass between these treatments were substantial. Such data suggest competition for soil resources between grasses andProsopis may be minimal early in the life cycle ofProsopis. High rates ofProsopis emergence and establishment on LTP-MD plots are counter to the widespread assumption that long-term and/or heavy grazing is requisite forProsopis encroachment into grasslands. Results are discussed with regard to factors contributing to the recent, widespread invasion of this woody legume into grasslands of southwestern North America.Abbreviations LTG long-term grazed - LTP long-term protected from grazing - ND non-defoliated - MD moderate defoliation - HD heavy defoliation     

Community‐level changes in Australian subalpine vegetation following invasion by the non‐native shrub Cytisus scoparius

Question: What are the changes associated with the recent invasion by the non‐native legume, Cytisus scoparius? Location: Subalpine vegetation (1500 m a.s.l.) in Australia. Methods: We used multivariate techniques and regression analyses to assess vegetation and environmental changes across six study sites. Vegetation and environmental variables were investigated at three different stages of invasion: (1) recent invasion (8–10 yr), (2) mature invasion (15–16 yr) and (3) long‐term invasion (25 yr). Results: Substantial changes in floristic composition and species richness were evident after 15 yr and these changes became more pronounced after 25 yr. Changes due to invasion were associated with a dramatic loss of native species or a reduction in their abundance. No ‘new species’ were evident under invaded stands. Forbs were most affected by the establishment of C. scoparius, although all growth forms responded negatively. Dense canopy shading and an increasingly dense, homogeneous litter layer in the understorey as a result of C. scoparius were strong environmental drivers of vegetation change. Greenhouse studies confirmed the importance of these processes on the germination and growth of two native species. Conclusions: This study highlights the potential for C. scoparius to alter both vegetation and environmental processes in the subalpine region.     

Forest dynamics following eastern hemlock mortality in the southern Appalachians

Understanding changes in community composition caused by invasive species is critical for predicting effects on ecosystem function, particularly when the invasive threatens a foundation species. Here we focus on dynamics of forest structure, composition and microclimate, and how these interact in southern Appalachian riparian forests following invasion by hemlock woolly adelgid, HWA, Adelges tsugae. We measured and quantified changes in microclimate; canopy mortality; canopy and shrub growth; understory species composition; and the cover and diversity in riparian forests dominated by eastern hemlock Tsuga canadensis over a period of seven years. Treatments manipulated hemlock mortality either through invasion (HWA infested stands) or girdling (GDL) hemlock trees. Mortality was rapid, with 50% hemlock tree mortality occurring after six years of invasion, in contrast to more than 50% mortality in two years following girdling. Although 50% of hemlock trees were still alive five years after infestation, leaf area lost was similar to that of girdled trees. As such, overall responses over time (changes in light transmittance, growth, soil moisture) were identical to girdled stands with 100% mortality. Our results showed different growth responses of the canopy species, shrubs and ground layer, with the latter being substantially influenced by presence of the evergreen shrub, rhododendron Rhododendron maximum. Although ground layer richness in the infested and girdled stands increased by threefold, they did not approach levels recorded in hardwood forests without rhododendron. Increased growth of co‐occurring canopy trees occurred in the first few years following hemlock decline, with similar responses in both treatments. In contrast, growth of rhododendron continued to increase over time. By the end of the study it had a 2.6‐fold higher growth rate than expected, likely taking advantage of increased light available during leaf‐off periods of the deciduous species. Increased growth and dominance of rhododendron may be a major determinant of future responses in southern Appalachian ecosystems; however, our results suggest hemlock will be replaced by a mix of Acer, Betula, Fagus and Quercus canopy genera where establishment is not limited by rhododendron.     

Host‐specificity constrains evolutionary host change in the psyllid Prosopidopsylla flava

1. At the higher taxonomic levels Psylloidea have largely co‐evolved with their host plants, and the colonisations of new plant lineages have been relatively few. The mechanisms that have constrained the evolution of host relationships throughout the history of this superfamily are not understood. The host relationships of Prosopidopsylla flava were studied in order to identify possible genetic or ecological constraints to macroevolutionary change in host range, using methodology developed for the host specificity testing of potential biological control agents. 2. The five Prosopis taxa (Leguminosae) tested appeared to be indistinguishable as hosts. Adult feeding required for survival, and for the initiation and continuation of egg production, was specific to Prosopis species. Oviposition occurred on 57 of the 58 non‐Prosopis plant species tested within the Leguminosae and Rosaceae, and was highest on plant species that belonged to the same subfamily. Eggs were inserted into plant tissue by their peduncle but hatched independently of host species. Complete development was restricted to Prosopis, although some early nymphal development was observed on species within the same subfamily as Prosopis (Mimosoideae). 3. Multiple phylogenetic constraints restrict host selection and utilisation by P. flava to Prosopis species, implying a long association between insect and host. Specificity of adult feeding was of special significance, being required for survival, oogenesis, and probably indirectly determining the oviposition host. This supports the hypothesis that genetically set limits in particular aspects of life history are responsible for the inability of some psyllids to readily colonise new plant lineages, rather than stabilising selection.     

Structural characteristics and canopy dynamics of Tsuga canadensis in forests of the southern Appalachian Mountains, USA

Tsuga canadensis (L.) Carr. forests of the southern Appalachian Mountains are currently facing imminent decline induced by a nonnative insect pest, the hemlock woolly adelgid (Adelges tsugae Annand). To effectively manage these forest systems now and in the future, land managers need baseline data on forest structure and dynamics prior to large-scale Tsuga canadensis mortality. Most of our knowledge concerning the dynamics of Tsuga canadensis forests comes from more northern locations such as the Great Lakes region and New England and, therefore, may not pertain to the ecological systems found within the southern Appalachian Mountains. We examined the structure and canopy dynamics of four Tsuga canadensis forest stands within the Cataloochee watershed, in the far eastern part of Great Smoky Mountains National Park (GSMNP). We characterized the environmental settings and vertical forest layers, as well as the diameter and age-structures of each Tsuga canadensis forest stand. These environmental and structural data showed that there were indeed differences between forest stands with and without successful Tsuga canadensis regeneration. The two forest stands exhibiting successful Tsuga canadensis regeneration were located above 1,000 m in elevation on well-drained, moderately steep slopes and had the greatest canopy openness. Structural data from these two forest stands indicated a history of more continuous Tsuga canadensis regeneration. We also constructed disturbance chronologies detailing the history of canopy response to disturbance events and related these to Tsuga canadensis regeneration within each forest stand. Student t-tests adjusted for unequal variances indicated significant differences in the number of release events per tree between forest stands with and without successful Tsuga canadensis regeneration. While forest stands with successful Tsuga canadensis regeneration were more frequently disturbed by minor to major canopy disturbances, events of moderate intensity were found to be most significant in terms of regeneration. These data will be of value to land managers maintaining stands of Tsuga canadensis where treatment for hemlock woolly adelgid infestation has been successful. In areas where treatment is impractical or unsuccessful, land managers will be able to use these data to restore Tsuga canadensis forests after the wave of hemlock woolly adelgid induced mortality has passed. As of August 2008, Joshua A. Kincaid will be a member of the Environmental Studies program at Shenandoah University in Winchester, Virginia, USA     

Biofuels and invasive species from an African perspective – a review

A large number of proposed biofuel crops share the same traits as known invasive plant species, many of which are already present in Africa and include species such as Prosopis glandulosa Torrey (Mimosaceae), P. juliflora (Sw.) DC, Leucaena leucocephala (Lam.) de Wit (Mimosaceae), Azadirachta indica A. Juss. (Meliaceae), and others. In this paper, we mainly assess the impacts of invasive Prosopis species in Africa, particularly in Kenya and South Africa. Introduced Prosopis species have invaded over 4 million hectares in Africa, threatening crop and pasture production, reducing underground water reserves, and displacing native plant and animal species. This has major implications for millions of people who depend on natural resources for their survival. It is therefore suggested that known invasive or potentially invasive plant species not be introduced to countries or regions for biofuel production. If (after a stringent cost–benefit analysis) the introduction of a potentially invasive species is deemed critical for economic development and the benefits clearly outweigh the potential costs, countries should endeavour to abide by the Principles and Criteria for Sustainable Biofuel Production developed by the Roundtable on Sustainable Biofuels.     

Soil moisture redistribution as a mechanism of facilitation in savanna tree–shrub clusters

Plant–soil water relations were examined in the context of a selective removal study conducted in tree–shrub communities occupying different but contiguous soil types (small discrete clusters on shallow, duplex soils versus larger, extensive groves on deep, sandy soils) in a subtropical savanna parkland. We (1) tested for the occurrence of soil moisture redistribution by hydraulic lift (HL), (2) determined the influence of edaphic factors on HL, and (3) evaluated the significance of HL for overstory tree–understory shrub interactions. Diel cycling and nocturnal increases in soil water potential (Ψ_soil), characteristic signatures of HL, occurred intermittently throughout an annual growth cycle in both communities over a range of moisture levels (Ψ_soil=−0.5 to −6.0 MPa) but only when soils were distinctly stratified with depth (dry surface/wet deep soil layers). The magnitude of mean (±SE) diel fluctuations in Ψ_soil (0.19±0.01 MPa) did not differ on the two community types, though HL occurred more frequently in groves (deep soils) than clusters (shallow soils). Selective removal of either Prosopis glandulosa overstory or mixed-species shrub understory reduced the frequency of HL, indicating that Prosopis and at least one other woody species was conducting HL. For Zanthoxylum fagara, a shallow-rooted understory shrub, Prosopis removal from clusters decreased leaf water potential (Ψ_leaf) and net CO₂ exchange (A) during periods of HL. In contrast, overstory removal had neutral to positive effects on more deeply-rooted shrub species (Berberis trifoliolata and Condalia hookeri). Removal of the shrub understory in groves increased A in the overstory Prosopis. Results indicate the following: (a) HL is common but temporally dynamic in these savanna tree–shrub communities; (b) edaphic factors influencing the degree of overstory/understory development, rooting patterns and soil moisture distribution influence HL; (c) net interactions between overstory and understory elements in these woody patches can be positive, negative and neutral over an annual cycle, and (d) Prosopis-mediated HL is an important mechanism of faciliation for some, but not all, understory shrubs.     

Use of Plant Strategy Ordination,DCA and ANOVA to elucidate relationships among habitats of Salix planifolia and Salix monticola

Abstract. Detrended Correspondence Analysis (DCA) was used on a data set of 112 species from 14 stands dominated by Salix planifolia and/or S. monticola. Environmental variables were determined which might be important to explain differences in stand vegetation composition, and willow stature (height), in willow carrs in the Rocky Mountain National Park. Correlation of environmental variables with DCA stand scores indicate that peat depth and soil redox potential are highly related to the primary DCA axis. Soils in stands of S. planifolia are more reduced in the early summer and have greater peat depths, than soils in stands of S. monticola. The second DCA axis is related to a water chemistry gradient of hydrogen ion, sulphate, and ammonium concentration. Four environmental variables selected by correlation analysis were regressed on stand scores from the first two DCA axes. Regression coefficients of August soil redox potential and soil water pH were significantly non-zero on the primary DCA axis. In addition to overall vegetation relationships elucidated by DCA, the relationships between willow stature and environmental variables were tested by simultaneous ANOVAs. ANOVA results and DCA ordinations indicate that S. planifolia and S. monticola plants achieve the greatest stature in the least reduced soils, and in shallow peat. Competitive, stress tolerant, and ruderal plant strategy theory applied to the vegetation data produced an arrangement of stands highly correlated to the DCA Axis 1 stand arrangement. These correlations indicate that the primary axis of vegetation structure can be extracted either by species composition, or species morphology/physiology relationships. Concurrent use of DCA and plant strategy theory is suggested as a tool for the prediction of community composition.     

Early response of herbaceous vegetation to Rhododendron ponticum subsp. baeticum invasion in European Atlantic forests

Questions

Rhododendron ponticum subsp. baeticum is an invasive shrub of growing concern in continental Europe, but little is known about its impact on native plant communities. Here we ask: do environmental conditions differ between forest stands invaded by it and uninvaded stands? Do these differences correlate with R. ponticum's cover? Are these differences associated with differences in taxonomic and functional diversity of vascular plant species of the herb layer? Can these vegetation changes be explained by the sorting of certain life-history traits by R. ponticum-induced environmental changes?

Location

Several forests invaded by R. ponticum in the French Atlantic domain.

Methods

We recorded vegetation composition and a number of environmental variables in 400-m² plots that were established in 64 paired forest stands (32 invaded vs 32 uninvaded). We compiled traits from existing databases. We computed several metrics of taxonomic and functional diversity. We compared environmental variables and diversity metrics between invaded and uninvaded stands. We used correlation and regression analyses to relate them with R. ponticum's cover. We ran RLQ and fourth-corner analyses to explore the relationships between R. ponticum invasion, environmental variables, species traits, and vegetation composition.

Results

Independent of its abundance, R. ponticum invasion was associated with lower light arrival at the forest floor and increased litter thickness. Concomitantly, species richness and diversity and trait diversity were reduced. The major driver of species assemblages was soil pH, which strongly interacted with the invasion gradient. R. ponticum did not sort species according to traits associated with shade tolerance and thick-litter tolerance. However, tree and shrub saplings were more abundant in invaded than uninvaded stands, at the expense of graminoid and fern species.

Conclusions

As R. ponticum becomes the dominant shrub, it exerts new selection forces on life-history traits of extant species, mostly via reduced light availability, increased litter thickness, and physical competition, thereby reducing taxonomic and functional diversity of the herb layer, without impeding tree and shrub self-regeneration, at least in the short term.     

Post-fire management targeting invasive annual grasses may have inadvertently released the exotic perennial forb Chondrilla juncea and suppressed its biocontrol agent

Top-down and bottom-up factors affecting invasive populations are rarely considered simultaneously, yet their interactive responses to disturbances and management interventions can be essential to understanding invasion patterns. We evaluated post-fire responses of the exotic perennial forb Chondrilla juncea (rush skeletonweed) and its biocontrol agents to landscape factors and a post-fire combined herbicide (imazapic) and bacteria (Pseudomonas fluorescens strain MB906) treatment that targeted invasive annual grasses in a sagebrush steppe ecosystem. Biocontrol agents released against C. juncea in previous decades included Cystiphora schmidti (gall midge), Aceria chondrillae (gall mite), and Puccinia chondrillina (rust fungus). C. juncea abundance was greater in sprayed than unsprayed plots, and where soils were coarser, slopes faced southwest, solar heat loads and topographic water accumulation were greater, and cover of deep-rooted native perennials was lower. Mite infestation was greater in unsprayed plots, midge infestation was greater at higher elevations on steeper slopes, and midges were more abundant while rust was less abundant on gravelly soils. Biocontrol infestation levels varied considerably between years and could not be predicted in 2019 from 2018 infestation levels. Multiple biocontrol species were often present at the same plots but were rarely present on the same C. juncea individuals. These results suggest that spatial patterns of invasion by C. juncea are related to deep-soil water availability, warmer conditions, and alleviation of competition. Treatments designed to reduce invasive annual grasses may inadvertently release C. juncea by both reducing plant competition for soil resources and affecting biocontrol agent (mite) abundance.

     

Discovery of a life history shift: precocious flowering in an introduced population of Prosopis

The genus Prosopis contains many valuable, long-lived, multi-purpose legume trees, some of which are also invasive species. The time of first flowering is important for increasing production of sweet, protein-rich pods in plantations, especially under short rotations, and affects the rate of spread as a weed. Trees generally begin flowering at 3–5 years of age. However, seedlings from a small seedlot collected from an introduced population in southern Mauritania were observed to begin flowering at 3–4 months after germination. This is unknown in the native range of any Prosopis species, and appears to represent a major evolutionary event triggered after naturalization. This paper reports a detailed investigation with seed collected from Aleg, Brakna region, Mauritania, in 1998 (Prosopis sp. ‘Aleg’). Two experiments were established, in glasshouses at Cirad, Montpellier, France, and Coventry University, UK. Flowering began at the two sites 104 and 169 days after sowing, respectively, and 97.5% of plants had flowered at Coventry after 306 days. Flowers produced abundant pollen with 50–60% viability indicated by FDA staining and 40–50% germination on an artificial medium. No pods were formed. All plants analysed were diploid. Morphologically, these juvenile plants were similar to American species of section Algarobia, and exhibited leaf characters typical of P. pallida. However, due to the large variation in morphology within this section of the genus, further studies are required to confirm the actual species. This is the first record of precocious flowering in Prosopis, and has been found only in a single, introduced population. Implications for genetic improvement of tropical Prosopis, and particularly the spread of these species as invasive weeds, are discussed. Such a life history shift in the increased reproductive ability of a species following introduction, with potentially significant environmental effects, may not be restricted to Prosopis and merits further detailed investigation.     

Changes in aboveground primary production and carbon and nitrogen pools accompanying woody plant encroachment in a temperate savanna

When woody plant abundance increases in grasslands and savannas, a phenomenon widely observed worldwide, there is considerable uncertainty as to whether aboveground net primary productivity (ANPP) and ecosystem carbon (C) and nitrogen (N) pools increase, decrease, or remain the same. We estimated ANPP and C and N pools in aboveground vegetation and surface soils on shallow clay and clay loam soils undergoing encroachment by Prosopis glandulosa in the Southern Great Plains of the United States. Aboveground Prosopis C and N mass increased linearly, and ANPP increased logarithmically, with stand age on clay loam soils; on shallow clays, Prosopis C and N mass and ANPP all increased linearly with stand age. We found no evidence of an asymptote in trajectories of C and N accumulation or ANPP on either soil type even following 68 years of stand development. Production and accumulation rates were lower on shallow clay sites relative to clay loam sites, suggesting strong edaphic control of C and N accumulation associated with woody plant encroachment. Response of herbaceous C mass to Prosopis stand development also differed between soil types. Herbaceous C declined with increasing aboveground Prosopis C on clay loams, but increased with increasing Prosopis C on shallow clays. Total ANPP (Prosopis+herbaceous) of sites with the highest Prosopis basal area were 1.2 × and 4.0 × greater than those with the lowest Prosopis basal area on clay loam and shallow clay soils, respectively. Prosopis ANPP more than offset declines in herbaceous ANPP on clay loams and added to increased herbaceous ANPP on shallow clays. Although aboveground C and N pools increased substantially with Prosopis stand development, we found no corresponding change in surface soil C and N pools (0–10 cm). Overall, our findings indicate that Prosopis stand development significantly increases ecosystem C and N storage/cycling, and the magnitude of these impacts varied with stand age, soil type and functional plant traits