Altitudinal change in soil and foliar nutrient concentrations and in microclimate across the tree line on the subtropical island mountain Mt. Teide (Canary Islands)

Mt. Teide (Tenerife, Spain) is a high volcanic island mountain with an unusually low tree line elevation (2000–2100 m). While searching for the causes of this tree line depression, we analysed the concentrations of total N, available P, and salt-exchangeable Ca, K and Mg in three soil horizons, and the foliar N, P, Ca, K and Mg concentrations in six abundant plant species (trees, shrubs, forbs) along a transect from 1400 (1600) to 3100 m a.s.l. The objective of the study was to detect altitudinal trends in soil and plant nutrient status below and above the tree line. For characterising elevational changes in microclimate and hydrology, we also conducted measurements of air and soil temperatures (T_a and T_s), atmospheric water vapour saturation deficit (D), potential evaporation (E) and soil moisture (θ) on 3–4 occasions along the transect.T_a and T_s did not linearly decrease with elevation but were highest at or immediately below the tree line and were relatively low in the closed Pinus canariensis forest at 1600 m. T_s reached maxima at about 70 °C near the tree line. The absence of a linear temperature decrease with elevation was caused by a stable temperature inversion at about 2000 m and by canopy shading in the forest canopy below the tree line. In June, the topsoil (10 cm) dried out almost completely between 1800 and 3100 m, but remained moist in the pine forest at 1600 m. This gradient reflects the transition from the montane cloud belt to the dry alpine belt higher upslope. The subsoil (30 cm) contained >30 vol% of soil water at all elevations even in June except for the uppermost site (3100 m). Potential evaporation increased with elevation despite a decrease in D. We assume that this is mainly due to the air pressure-dependent increase with elevation in the diffusion coefficient for water vapour in air.The concentrations of N and ‘available’ P (after Olsen) in the mineral topsoil were by far smaller than in alpine soils of other humid mountains which is thought to be a consequence of a very dry and biologically inactive topsoil in the semi-arid alpine belt of Mt. Teide. In contrast to many other mountains, foliar N, P and cation concentrations in the plants did not increase with elevation but either remained unchanged, or decreased as in the tree line species P. canariensis. Nevertheless, P. canariensis probably is not limited by nutrient deficiency at the tree line despite rather low N and P needle concentrations. Rather, drought and heat stress effects on seedling establishment are thought to be the causes of the tree line depression.     

Secondary successional patterns in a sagebrush (Artemisia tridentata) community as they relate to soil disturbance and soil biological activity

The relationship between secondary succession, soil disturbance, and soil biological activity were studied on a sagebrush community (Artemisia tridentata) in the Piceance Basin of northwestern Colorado, U.S.A. Four levels of disturbance were imposed. I: the vegetation was mechanically removed and as much topsoil as possible was left; 2: the vegetation was mechanically removed and the topsoil scarified to a depth of 30 cm; 3: topsoil and subsoil were removed to a depth of 1 m, mixed and replaced; 4: topsoil and subsoil were removed to a depth of 2 m and replaced in a reverse order. Plant species composition, dehydrogenase and phosphatase enzymatic activity, mycorrhizae infection potentials, and percent organic matter were the variables measured.Treatment 4 drastically altered the pattern of vegetation succession. Treatments 2, 3, and 4 started with Salsola iberica as the dominant species but six years later, 3 and to lesser extent 2 changed in the direction of the species composition of 1, dominated by perennial grasses and perennial forbs. Treatment 4 developed a shrub dominated community. The rate of succession was not decreased by the increased levels of disturbance. Both dehydrogenase enzyme activity and mycorrhizae infection potential (MIP) increased with the change from Salsola iberica to a vegetation dominated by either perennial grasses and forbs or shrubs. The intensity of disturbance in 2, 3, and 4 reduced drastically dehydrogenase activity and MIP, but in six years they recovered to levels comparable to 1. Phosphatase enzyme activity and organic matter were unrelated to species composition but related to treatment and time elapsed. In both cases a significant decrease was observed throughout the six-year period.Nomenclature followsThis study was funded by the United States Department of Energy under Contract No. DE-AS02-76EV04018.     

Germination and population structure of spear thistle Cirsium vulgare in relation to experimentally controlled sheep grazing

Summary The size and number of rosettes of Cirsium vulgare were censused in a 4 ha sheep grazing trial on lowland calcareous grassland in paddocks receiving controlled winter, spring and summer grazing treatments. Spring grazing significantly increased thistle rosette numbers, and there were always fewer rosettes in size classes 250mm–300 mm under the heavier grazing treatments. The emergence of seeds of Cirsium vulgare sown by hand into each of the grazing treatments was monitored and showed a positive effect of spring grazing. When the percent emergence of sown seeds was used as a covariate in the analysis of rosette numbers in the experimental paddocks, it was found to account for 77% of the variance in thistle numbers. It was concluded that the main effect of grazing on thistle rosette numbers is an indirect one exercised through the effects of grazing on germination conditions. Seeds of Cirsium vulgare were sown into artificial gaps at another grassland site, to determine the effect of gap-size upon seedling emergence. Germination was poor, but significantly more seedlings emerged in gaps 10–20 cm diameter, than in 5 cm gaps or in control plots without a gap. A computer simulation model was used to explore the relationship between gap density and thistle population dynamics. A threshold density of gaps was found to exist, below which thistle populations went extinct, and above which the thistle population grew geometrically. The degree of aggregation of dispersed seeds did not alter the threshold gap-density for plant extinction, but did affect the rate of increase of the thistle population when the threshold gap-density was exceeded. It is concluded that strategies for controlling Cirsium vulgare populations by grazing manangement will be most effective if aimed at reducing suitable sites for establishment in spring. The success of such attempts will depend upon the soil fertility of the site, and sudden outbreaks of Cirsium vulgare infestation can be expected if a sward gradually deteriorates through over-grazing.     

Temperature sensitivity as a general phenomenon in a collection of chlorophyll-deficient mutants of sweetclover (Melilotus alba)

A collection of chlorophyll (Chl)-deficient mutants of sweetclover (Melilotus alba) with defects in eight nuclear loci were grown at 17 or 26° C. Plants grown at either temperature were examined for Chl content, Chla/b ratio, expression of the light-harvesting complex II (LHC-II) apoproteins, and protochlorophyllide (Pchlide) biosynthetic capacity. Except for thech4 mutant, the parental strain and all mutants accumulate more Chl when grown at 26° C than at 17° C. Thech5 mutants, lacking Chl b under any growth condition, and thech12 mutant showed little temperature-dependent phenotypic plasticity, whereas this was a marked phenomenon in the other mutants. Thech10 andch11 mutants demonstrated extreme temperature sensitivity with regard to the production of Chlb and the Chlb-binding LHC-II apoproteins. When excised trifoliolates were supplemented with exogenously supplied -aminolevulinic acid, only thech4 mutant was markedly impaired in the ability to produce Pchlide. These data indicate that temperature-sensitive phenotypic plasticity is a common phenomenon of chlorophyll-deficient mutants and substantiate that only a minority of Chl-deficient mutants is impaired in the biosynthesis of Chl.This research was supported by Grants GM84-CRCR-1-1479 (J.C.O.) and 89-00641 (J.M.) of the United States Department of Agriculture and by National Science Foundation Grant DMB87-03100 (J.M.). This is paper No. 8971, Nebraska Agricultural Research Division.     

Association between feral pig disturbance and the composition of some alien plant assemblages in Hawaii Volcanoes National Park

Association analysis was used to assess relationships among 25 important alien plant species and their association with feral pig rooting in Hawaii Volcanoes National Park, Hawaii, U.S.A. Results of the association analysis were summarized by means of a simplified, rank-based, polar ordination which yielded three subjective species assemblages. One group was characterized by an association with the endpoint species, Ehrharta stipoides. A second group consisted of species associated with the other endpoint species, Andropogon virginicus. The third group comprised a diverse assemblage of 14 species in the middle of the ordination.Comparison of ordination scores with each species's association with pig-induced soil disturbance revealed that members of the Ehrharta group were strongly positively associated with pig activity, whereas members of the Andropogon group were generally negatively associated. The third group showed no association with pig-induced soil disturbance.These results suggest a strong relationship between feral pig activity and the composition of the alien portion of the plant community. Analysis of the ecologies of both plants and pigs suggests that some species may both encourage pig activity and benefit from it. Likewise, other alien plants appear to neither require nor benefit from pig-induced soil disturbance. While pigs appear to play an important role in the organization of these communities, their removal may have a negligible impact on the success of many of the common weeds in the area. Nomenclature: Wagner, W. L., Herbst, D. R., & Sohmer, S. H. 1990. Manual of the Flowering Plants of Hawai'i. University of Hawaii Press/Bishop Museum Press, Honolulu.     

Interannual growth response of Norway spruce to climate along an altitudinal gradient in the Tatra Mountains, Poland

Ring-width chronologies of Picea abies (L.) Karst. from ten sites in the Tatra Mountains, Poland, were developed to explore growth/climate responses in stands along an altitudinal gradient ranging from 839 to 1468 m a.s.l. There were positive relationships between current-year radial growth and mean monthly temperatures in March, April, June and July, but with increasing elevation, the strength of this correlation declined for March–April and increased for June–July temperatures. The mean monthly temperature in October of the previous year positively influenced radial growth of trees at all sites. Lower mean temperatures in January negatively affected growth of trees at the high-elevation sites. Trees at the low-elevation sites responded positively to a warm early spring, whereas trees at the high-elevation sites showed positive growth responses to higher summer temperatures. Growth of trees at the high-elevation sites was better explained by the temperature regime than was growth at the lower-elevation sites. Therefore, it is likely that these sites may be particularly sensitive to potential effects of temperature change.     

Effects of ants on plant diversity in semi-natural grasslands

Ants (L. niger and L. flavus) build conspicuous mounds that are covered with vegetation. The aim of this study was to investigate whether the vegetation on ant mounds in semi-natural grasslands differed from that around the mounds. Another aim was to investigate whether the changes in the vegetation on ant mounds were influenced by grazing management or by habitat characteristics, semi-dry versus moist. Here, the total number of plant species and total plant cover were lower on ant mounds than in patches off-mound. The plant cover of perennials that form rosettes was twice as high on mounds inhabited by L. niger than on those inhabited by L. flavus. Only a few plant species were restricted to either ant mounds or adjacent field and the effects of ants on the plant diversity in semi-natural grasslands seemed to be low. Grazing management did not affect the differences in the vegetation on ant mounds and in equal-sized patches off-mound, whereas habitat characteristics affected ant-induced changes in vegetation cover of some plant species.     

Regional and local variation in seedling emergence,mortality and recruitment of a perennial herb in Mediterranean mountain habitats

Spatial patterns of seedling regeneration of woody Mediterranean plants are widely documented, with seedlings and juveniles growing almost exclusively under shrub and tree canopy. Neighbourhood habitat amelioration and consequent facilitative effects on recruitment have been extensively suggested as the major determinant of such spatial pattern for Mediterranean vegetation. Much less is known on recruitment patterns of perennial herbs. As herbs differ from woody plants in many relevant ecological aspects and in their life cycles, we would expect particularities in their recruitment dynamics. We analysed the spatial (regional, local and fine-scale) variation in environment (herb cover, litter depth, air temperature and irradiance) and its relationship with seedling emergence, mortality, and recruitment of the perennial herb Helleborus foetidus (Ranunculaceae) during 2 years. We conducted the study in three distant localities in south-eastern Iberian Peninsula (defining the regional scale). Local scale was defined in terms of microhabitat type based on three categories of vegetation cover (under evergreen shrubs or trees, under deciduous spiny-shrubs, and in open interspaces). Finally, fine-scale variation was defined in terms of 1 m² sampling plots. Our results showed regional, local and fine-scale variations in recruitment, with the major source of variation changing across scales. Seed input was important in determining differences in recruitment both at the largest (regional) and the finest scale. Environment had minor importance in shaping differences in recruitment at these two scales. At the local scale, variation between microhabitats was mainly related to differences in seedling survival through facilitative effects (alleviation of water and irradiance stresses) of shrub and tree cover, although such facilitative effect was not consistent in all localities or years. Our study thus points to a similarity in the spatial patterns of recruitment (and in the environmental factors shaping such patterns) between Mediterranean woody plants and perennial herbs, but much more effort is needed to assess the generality of such similarity.     

Alien and native plants show contrasting responses to climate and land use in Europe

Aim We tested whether the distribution and cover of alien plant species in Europe was related to human disturbance and microclimate. Location Surveys were conducted at 13 sites across Europe, each containing a pair of landscapes with different land‐use intensities. Methods Sampling locations were chosen based on land use and microclimate at two scales: land use was characterized at the patch and landscape scale; climate was expressed as regional and local temperature. The slope of each sample location was derived from a digital elevation model. Cover of plant species was measured using point counts and analysed using mixed effect models. Species were classified as native, archaeophytes and neophytes (pre‐ versus post‐ad 1500 immigrants). Due to the zero inflation observed in the alien groups, their cover was analysed conditional on their presence. Results Anthropogenic disturbance was a significant explanatory variable, increasing the presence and cover of alien species and decreasing the cover of native species. Alien presence was increased in sites under agricultural management, while their cover responded to land use at both local and landscape scales (and to their interaction), such that only natural habitats in semi‐natural landscapes had low alien cover. Microclimate was important for neophytes, with presence concentrated around mesic conditions. Slope was relevant for archaeophytes and native species, suppressing the former group and promoting the latter one. Main conclusions We found that, at the European scale, the distribution of alien plants is related to anthropogenic disturbance more than to microclimatic differences. The presence of neophytes, however, was influenced by climate at local and regional scales, with the highest incidence under mesic conditions. The different patterns observed for the presence and cover of alien species suggest different mechanisms acting during their establishment and spread. They also suggest that to counteract the expansion of alien species natural habitats may need to be maintained at landscape scales.     

Coupling effects of altitude and human disturbance on landscape and plant diversity in the vicinity of mountain villages of Beijing, China

Villages represent the source of human disturbance in mountain regions, but how they alter surrounding landscapes and further affect plant diversity distribution along altitudinal gradient is still not well documented. Although the unimodal pattern along latitudinal gradient and the hypothesis of immediate disturbance (IDH) have been supported in many studies, their coupling effects on plant diversity distribution have been given less attention. In this paper, the coupling effects surrounding mountain villages were detected: (1) altitude determined the mountain landscape at regional scale, while human disturbance altered landscape fragmentation and diversity at local scale surrounding villages. (2) With the reducing human disturbance away from villages, plant diversity decreased, then increased, and finally decreased in different land uses. The plant diversity of shrubs reached the lowest. With the increasing altitude, plant diversity of the forest represented the unimodal trend, but other land use types had different properties. (3) The mechanism of the coupling effects is that the combination of topographic and soil factors determines plant diversity distribution at both landscape and plant community levels surrounding villages.     

Pastoralism, plant conservation, and conflicts on proliferation of Himalayan knotweed in high altitude protected areas of the Western Himalaya, India

The conservation policy of banning pastoral grazing, and subsequently emerging conflicts between conservationists and pastoral communities regarding the proliferation of Himalayan knotweed (Polygonum polystachyum), was studied in the Valley of Flowers National Park, a high altitude protected area of the Western Himalaya, India. A total of 10 habitat types identified in the study area were sampled using quadrats along an altitudinal gradient between 3000 and 4500 m. Plant species richness decreases with altitude and also varies across habitat types. The highest density of P. polystachyum and its associated species, Impatiens sulcata, was found between 3300 and 3500 m in the disturbed habitat types, viz., bouldery areas, fragmented treeline zone, avalanche-prone areas and eroded slopes. Eradication of P. polystachyum from the national park by managers is not going to serve any meaningful purpose to the long-term conservation; rather it initiates soil erosion and instability, hindering the establishment of natural plant communities.     

Drosophilid assemblages as a bioindicator system of human disturbance in the Brazilian Savanna

According to the Convention on Biological Diversity (CBD) the development of bioindicators is extremely necessary to achieve the conservation targets by 2010, and insects are considered an effective group for this goal. Drosophilids are regarded as potential indicators, although this idea remains untested. Therefore, we followed up a protocol to test the drosophilid potential indicator for human disturbance in the Brazilian Savanna, one of the richest and most threatened tropical biomes in the world. Sampling was undertaken in one urban environment and two biological reserves, representing four habitat types (undisturbed gallery forest, disturbed gallery forest, undisturbed savanna, and urban environment). We examined differences in the drosophilid assemblages among habitat types and used the Indicator Value (IndVal) method to point out the indicator species. We also tested the two-stage indicator validation, a protocol recently proposed in the literature, to validate the indicator species for undisturbed gallery forest and savannas, in independent samples. The assemblage variables varied mainly in undisturbed gallery forests, and reflected changes from an undisturbed to a disturbed stage. The IndVal associated with the two-stage protocol showed reliable characteristic species, which are very helpful for diagnostic surveys. Likewise, species that can detect changes in the habitats were also found. We found a set of indicators, which together may be very efficient for both assessing and reflecting a variety of conditions, improving the confidence of the bioindication system, expanding the taxonomic options for bioindicators, and therefore, contributing to the conservation of this region.     

Responses to fertility and disturbance in a low-diversity grassland

We examined variation in species composition in a low-diversity, anthropogenic grassland in response to 11 years of nitrogen (N) manipulation and disturbance. The species-poor grassland (2–3 species/0.5 m²) represents a wide spread vegetation type (>10 million ha in North America) dominated by the introduced perennial grasses Bromus inermis and Agropyron cristatum. Four levels of N and three of soil disturbance were applied in all combinations to plots (5 × 15 m, N = 120) in a completely randomized design each year. Seeds or transplants of 47 species were added to ensure that dispersal was not a barrier to changes in species composition. After 11 years of treatment, all but the most disturbed plots continued to be dominated by B. inermis. The cover of the second-most abundant species, A. cristatum, decreased with disturbance but did not vary significantly with N. Despite the lack of changes in the identity of the dominant species, our environmental manipulations strongly influenced ecosystem characteristics. Added N increased soil available N, and decreased the cover of bare ground and light availability. Soil disturbance decreased aboveground biomass, and increased the cover of bare ground and light availability. Sawdust application, designed to decrease N availability, significantly reduced community biomass, and increased light availability and the cover of bare ground, but did not alter nutrient availability or species composition. The results highlight the difficulty of restoring diversity in species-poor, anthropogenic communities dominated by introduced species, and thus the importance of conserving remnants of diverse natural grasslands.     

Understory vegetation response to thinning disturbance of varying complexity in coniferous stands

Question: Can augmented forest stand complexity increase understory vegetation richness and cover and accelerate the development of late‐successional features? Does within‐stand understory vegetation variability increase after imposing treatments that increase stand structural complexity of the overstory? What is the relative contribution of individual stand structural components (i.e. forest matrix, gaps, and leave island reserves) to changes in understory vegetation richness? Location: Seven study sites in the Coastal Range and Cascades regions of Oregon, USA. Methods: We examined the effects of thinning six years after harvest on understory plant vascular richness and cover in 40‐ to 60‐year‐old forest stands dominated by Douglas‐fir (Pseudotsuga menziesii). At each site, one unthinned control was preserved and three thinning treatments were implemented: low complexity (LC, 300 trees ha^?1), moderate complexity (MC, 200 trees ha^?1), and high complexity (HC, variable densities from 100 to 300 trees ha^?1). Gaps openings and leave island reserves were established in MC and HC. Results: Richness of all herbs, forest herbs, early seral herbs and shrubs, and introduced species increased in all thinning treatments, although early seral herbs and introduced species remained a small component. Only cover of early seral herbs and shrubs increased in all thinning treatments whereas forest shrub cover increased in MC and HC. In the understory, we found 284 vascular plant species. After accounting for site‐level differences, the richness of understory communities in thinned stands differed from those in control stands. Within‐treatment variability of herb and shrub richness was reduced by thinning. Matrix areas and gap openings in thinned treatments appeared to contribute to the recruitment of early seral herbs and shrubs. Conclusions: Understory vegetation richness increased 6 years after imposing treatments, with increasing stand complexity mainly because of the recruitment of early seral and forest herbs, and both low and tall shrubs. Changes in stand density did not likely lead to competitive species exclusion. The abundance of potentially invasive introduced species was much lower compared to other plant groups. Post‐thinning reductions in within‐treatment variability was caused by greater abundance of early seral herbs and shrubs in thinned stands compared with the control. Gaps and low‐density forest matrix areas created as part of spatially variably thinning had greater overall species richness. Increased overstory variability encouraged development of multiple layers of understory vegetation.     

Habitat modification inhibits conspecific seedling recruitment in populations of an invasive ecosystem engineer

Invasive species that strongly modify their physical habitat are a particular management concern. Theoretical models predict that habitat modification could speed spread rates or allow invasion of sites that would otherwise resist invasion. There are few empirical tests of this hypothesis, however. We tested whether habitat modification by invading Spartina alterniflora populations facilitates conspecific seedling recruitment and spatial spread in Willapa Bay, WA, USA. Established S. alterniflora individuals strongly modified their local physical environment. Hydrologic flow, porewater salinity, and light availability were decreased while sediment NH₄ ⁺ increased with increasing S. alterniflora stem density. The S. alterniflora seed bank was greater and spring seedlings were denser within meadows of S. alterniflora than on unvegetated tideflats. However, almost all seedling recruitment after 1 year occurred on tideflats or on meadow edge plots where the above ground S. alterniflora biomass had been removed. Instead of facilitating invasive spread, ecosystem engineering in this system appears to create conditions that inhibit local seedling recruitment. These results suggest that the influence of ecosystem engineering on invasive spread is highly contingent on the relative spatial scales of habitat modification, environmental heterogeneity, and propagule availability. Control activities could change these spatial relationships, however, inadvertently promoting invasive recruitment.     

Edaphic patchiness influences grassland regeneration from the soil seed‐bank in mountain grasslands of central Argentina

The soil seed‐banks in the main natural vegetation patches that make up mountain grasslands on granite substrates in central Argentina were studied. The main natural vegetation types are moist swards, tall‐tussock grasslands and stony grasslands. Ten compound soil samples from each community at two soil depths (0–5 and 5–10 cm) were taken. The density of soil seed‐banks was highest in moist swards, intermediate in stony grasslands and lowest in tall‐tussock grasslands. Low levels of similarity were found between the established vegetation and total soil seed‐bank in tall‐tussock grasslands and stony grasslands, but the similarity was higher in swards. In all three communities the seed‐bank was most frequently transient in nature. Short‐term persistent and long‐term persistent seed‐banks were less frequent. Regeneration from the seed‐banks after disturbance is expected to differ among communities on different edaphic patches. On the basis of the density and longevity of the soil seed‐banks and the similarity to the established vegetation, potential for in situ regeneration should be lower in tall‐tussock grasslands, intermediate in stony grasslands and higher in moist swards.     

Drought response of a native and introduced Hawaiian grass

The alien grass, Pennisetum setaceum, dominates many of the lowland arid regions that once supported native Heteropogon contortus grassland on the island of Hawaii. Response to drought in a glasshouse was compared between these C₄ grasses to test if success as an invader is related to drought tolerance or plasticity for traits that confer drought tolerance. Pennisetum produced 51% more total biomass, allocated 49% more biomass to leaves, and had higher net photosynthetic rates (P _n) on a leaf area basis than Heteropogon. Plants of both species under drought produced less total biomass and increased their allocation to roots compared to well-watered plants, but there was no difference between the two species in the magnitude of these responses. The decline in P _n with decreasing leaf water potential (₁) was greater for Pennisetum compared to Heteropogon. Plasticity in the response of P _n to ₁, osmotic potentials, and the water potentials at turgor loss in response to drought were not different between the two species. Stomata were more responsive to w in Heteropogon than in Pennisetum and for well-watered plants compared to droughted plants. Plasticity for the stomatal response to w, however, was not different between the species. There was no evidence that the alien, Pennisetum, had greater plasticity for traits related to drought tolerance compared to the native, Heteropogon. Higher P _n and greater biomass allocation to leaves resulted in greater growth for Pennisetum compared to Heteropogon and may explain the success of Pennisetum as an invader of lowland arid zones on Hawaii.     

Induced defences in an endangered amphibian in response to an introduced snake predator

Introduced species have contributed significantly to the extinction of endemic species on islands. They also create new selection pressures on their prey that may result in modified life history strategies. Introduced viperine snakes (Natrix maura) have been implicated in the decline of the endemic midwife toad of Mallorca (Alytes muletensis). A comparison of A. muletensis tadpoles in natural pools with and without snakes showed that those populations subject to snake predation possessed longer tails with narrower tail fins but deeper tail muscles. Field and laboratory experiments showed that these changes in tail morphology could be induced by chemical and tactile cues from snakes. Populations of tadpoles that were subject to snake predation also displayed clear bimodal size-frequency distributions, with intermediate-sized tadpoles missing from the pools completely. Tadpoles in pools frequented by snakes developed faster in relation to their body size than those in pools without snakes. Variation in morphology between toad populations may therefore be caused by a combination of size-selective predation and tadpole plasticity. The results of this study indicate that the introduction of alien species can result in selection for induced defences, which may facilitate coexistence between predator and prey under certain conditions.     

Habitat Invasibility and Dominance by Alien Annual Plants in the Western Mojave Desert

Patterns of habitat invasibility and alien dominance, respectively measured as species richness and biomass of alien annual plants, were evaluated in association with four habitat factors at the Desert Tortoise Research Natural Area (DTNA) in the western Mojave Desert, USA. Habitat factors varied in levels of disturbance outside (high) and inside (low) the DTNA, and in levels of soil nutrients in washlet (high) and hummock (low) topographic positions, in Larrea-north (high), Larrea-south (medium), and interspace (low) microhabitats near creosote bushes (Larrea tridentata), and during 1995 when rainfall was 207% (high) and 1994 when rainfall was 52% (low) of the long-term average. Dominant alien plants included the annual grasses Bromus rubens, Bromus trinii, and Schismus spp., and the forb Erodium cicutarium. Species richness and dominance of alien annual plants were slightly higher where disturbance was high, and much higher where soil nutrients were high. B. rubens and B. trinii were most dominant in washlets and in the Larrea-north microhabitats during both years. These two species evolved in mesic ecosystems, and appeared to be particularly limited by soil nutrients at this site. Schismus spp. and E. cicutarium were also most dominant in washlets, but their dominance varied between interspaces in 1994 and the Larrea-south microhabitat in 1995. Monitoring to detect the invasion of new annual plants should focus on regions of high rainfall and nitrogen deposition and on washes and beneath-canopy microhabitats. The ecological range of each alien species should be evaluated separately, because their evolutionary origins may greatly affect their patterns of invasion and dominance in the Mojave Desert. This revised version was published online in July 2006 with corrections to the Cover Date.     

Allometric growth, disturbance regime, and dilemmas of controlling invasive plants: a model analysis

Disturbed communities are observed to be more susceptible to invasion by exotic species, suggesting that some attributes of the invaders may interact with disturbance regime to facilitate invasion success. Alternanthera philoxeroides, endemic to South America, is an amphibious clonal weed invading worldwide. It tends to colonize disturbed habitats such as riparian zones, floodplain wetlands and agricultural areas. We developed an analytical model to explore the interactive effects of two types of physical disturbances, shoot mowing and root fragmentation, on biomass production dynamics of A. philoxeroides. The model is based on two major biological assumptions: (1) allometric growth of root (belowground) vs. shoot (aboveground) biomass and (2) exponential regrowth of shoot biomass after mowing. The model analysis revealed that the interaction among allometric growth pattern, shoot mowing frequency and root fragmentation intensity might lead to diverse plant ‘fates’. For A. philoxeroides whose root allocation decreases with growing plant size, control by shoot mowing was faced with two dilemmas. (1) Shoot regrowth can be effectively suppressed by frequent mowing. However, frequent shoot mowing led to higher biomass allocation to thick storage roots, which enhanced the potential for faster future plant growth. (2) In the context of periodic shoot mowing, individual shoot biomass converged to a stable equilibrium value which was independent of the root fragmentation intensity. However, root fragmentation resulted in higher equilibrium population shoot biomass and higher frequency of shoot mowing required for effective control. In conclusion, the interaction between allometric growth and physical disturbances may partially account for the successful invasion of A. philoxeroides; improper mechanical control practices could function as disturbances and result in exacerbated invasion.