Wet dune slacks occur as small, naturally fragmented systems in the dune landscape, isolated from other slacks. We studied the effects of slack isolation and area on the rate and direction of primary succession in a chronosequence of dune slacks. The results indicate that important changes occur in community characteristics over a period of 50 yr. Total cover and number of species increase as a result of the endogenous succession process, during which organic matter and nutrients accumulate. Consequently, competitive interactions shift from competition for nutrients to competition for light. Local factors thus determine, at least partially, the community composition in the slack. However, differences in community composition with increasing age are smaller when the slack is more isolated or smaller, suggesting a slower succession rate and biomass accumulation. Together with a lower contribution of slow dispersing species in more isolated slacks, this indicates that species accumulation is dispersal limited and thus influenced by regional factors. The stochastic variation resulting from this dispersal limited species accumulation causes a divergent successional pathway.

Nomenclature: Lambinon et al. (1998).

Abbreviation: MIV = Mean indicator value.

The phenology of spring leaf replacement was studied in a population of 46 evergreen scrub oaks (Quercus cornelius-mulleri) at the edge of the Mojave Desert in each of five years over the period 1990–2001. The oaks occupied a site that spanned rocky slopes to sandy bajadas. The site receives variable annual rainfall (estimated 12-yr average 195 mm; range in study years 67–706 mm). The spatial coordinates of all individuals were recorded, and in April, when leaf replacement was underway, individual replacement phenologies were assessed. Shrub sizes were recorded in three separate years, and in 2001 water potentials were measured.

Individuals vary greatly in their timing of leaf replacement within years, and also between years. Many individuals with an early phenology one year are significantly later in the following year, and vice versa. While we detected weak influences on leaf replacement phenology due to shrub size, position within the site, and a genetic component, stronger influences were attributable to the phenology of the shrub in prior years, and to the phenology of neighbours within years. Neighbouring individuals that are close and/or large are significantly disparate in phenology, with one early and the other late. A potential mechanism of local resource depletion associated with costs to an early phenology is discussed.

Abbreviation: LRP = Leaf replacement phenology; PSA = Photosynthetic area; RPH = Ranked leaf replacement phenologies.

Nomenclature: Nixon & Steele (1981).

Response of a species to an environmental variable may be modeled and predicted using a wide spectrum of different functions. Contrary to other functions (Gaussian, polynomial etc), all parameters of the β-function are interpretable in ecological terms. However, computational difficulties in the determination of the β-function parameters initiated controversial debates on the applicability and usefulness of this function in vegetation modelling and gradient analysis. We propose a simple algorithm for fitting the β-function to observed data. Analytic properties of the algorithm (its ability to recover the known species responses along gradients) are tested using a series of simulated data. In most cases the algorithm correctly estimated parameters of the simulated responses.

Woody legumes growing in dry climates can increase soil nutrient content and facilitate plant growth in their understorey. We investigated differences in soil fertility and herbaceous community in relation to the presence and absence of the legume Retama sphaerocarpa in a shrubland in a mediterranean type climate. The results showed a higher content of limiting nutrients for plant growth, such as phosphorus and nitrogen, under the R. sphaerocarpa canopy. Herbaceous biomass, cover and nitrogen content increased below the canopy. However, species richness and diversity were diminished in the presence of a dense canopy of R. sphaerocarpa. Nitrogen isotopic analysis did not indicate a clear relationship between symbiotic fixation in R. sphaerocarpa and nitrogen content of soil and plants under its canopy. Nevertheless, herbs growing in the understorey showed a linear correlation between foliar N content and ¹⁵N values. The existence of a dense shrub canopy induced a smaller monthly variation in herb richness, diversity and biomass, suggesting that it provides a stable microhabitat facilitating herbaceous vegetation establishment and growth. The ‘shelter’ effect was more important when environmental conditions became stressful for herbaceous plants.

Nomenclature: Tutin et al. (1968–1980); Devesa (1995).

Plant cover was visually estimated by five observers, independent of each other, in a species-rich grassland in the Bílé Karpaty Mts., southeastern Czech Republic, in seven plots ranging from 0.001 to 4 m². Variation of total plant cover among the observers was high at small scales: 0.001 - 0.016 m²; coefficient of variation, CV = 35 to 45%, but much lower at larger scales: 0.06 - 4 m²; CV = 7 to 15%. Differences between visual estimates of plant cover of individual species made by different observers were affected by plot size, total cover and morphology of particular plants. CV of the cover of individual species ranged from 0 to 225% and decreased with increasing plot size. For abundant plants the CV attained ca. 50%, independent of plot size. In spite of a very high number of sterile plants with similar leaf morphology and colour, the observed variation in cover estimates in the studied grassland was comparable with results reported from other vegetation types. Differences between estimates by individual observers were often larger than usual year to year changes in undisturbed grasslands. Therefore, I suggest that to avoid difficulties in the interpretation of results based on plant cover data obtained from visual estimates, several observers should always work together, adjusting their extreme estimates.

Nomenclature: Marhold & Hindák (1998).

A spatially explicit model was developed to study the relationships between the dynamics and spatial structure of forest stands. The objective was to test whether tree spatial structure can be used as an indicator of stand dynamics. The model simulates the growth, mortality and recruitment of trees in a multi-specific and uneven-aged stand. It includes deterministic and stochastic processes so that repeated simulations do not lead to the same stand but provide several possible results for a given dynamic (defined by a set of parameters). Second-order neighbourhood analyses were used to characterize the resulting spatial structures. They showed a high variability for a given set of parameters. Only the main trends in the spatial structure can be interpreted. Sensitivity analyses, concerning the influence of competition on spatial structure, showed that in heterogeneous stands confounding effects can hinder the interpretation of the spatial structure if all the trees are considered. The spatial structure of the canopy trees alone proved easier to interpret as it is directly linked to post recruitment competition. Inference on the dominant modality of competition (one-sided or two-sided) based on the spatial structure proved difficult.

This study examined the influence of the fungal endophyte Neotyphodium lolii on the competitive interactions between its perennial, cool season host, Lolium perenne (perennial ryegrass), and a warm season, annual grass, Digitaria sanguinalis (large crabgrass), in densely planted stands (>1000 plant.m⁻²) in the glasshouse. Endophyte infection had little or no effect on L. perenne tiller production, above- or below-ground biomass or root:shoot ratio in monoculture. However, endophyte infection significantly reduced L. perenne tiller production and above-ground biomass in mixtures with D. sanguinalis. Conversely, D. sanguinalis had significantly higher above-ground biomass and yielded more seed (g) when competing with endophyte infected L. perenne. An apparent trade-off between allocation of resources to reproductive vs root tissues was observed in D. sanguinalis – root:shoot ratio was significantly lower when competing with endophyte infected L. perenne. Results indicate negative ecological consequences for harbouring the fungal endophyte when competing with the fast growing annual grass in newly established stands. These findings underscore the existence of a physiological cost of harbouring the fungal endophyte which is often overlooked.

Woody plants are increasing in many grassland and savanna ecosystems around the world. As a case in point, the Edwards Plateau of Texas, USA, is a vast region (93 000 km²) in which rapid woody encroachment appears to be occurring. The native vegetation (prior to the Anglo-European settlement 150 – 200 yr ago) and the biogeochemical consequences of woody encroachment in this region, however, are poorly understood. To assess these matters we measured plant and soil δ¹³C, soil organic C and soil N content from grasslands and two important woody patch types (mature Quercus virginiana clusters and Juniperus ashei woodlands) in this region. Soil δ¹³C values showed that relative productivity of C₃ species has increased in grassland and both woody habitats in recent times. δ¹³C of SOC in grasslands and Q. virginiana clusters increased with depth from the litter layer to 30 cm (grasslands = − 21 to − 13‰; Q. virginiana clusters = − 27 to − 17‰) and were significantly different between habitats at all depths, indicating that Q. virginiana has been a long-term component of the landscape. In J. ashei woodlands, soil δ¹³C values (at 20 – 30 cm depth) near the woodland edge (−13‰) converged with those of an adjacent grassland (−13‰) while those from the woodland interior (−15‰) remained distinct, indicating that the woodland has been present for many years but has recently expanded. Concentrations and densities of SOC and total N were generally greater in woody patches than in grasslands. However, differences in the amount of SOC and N stored beneath the two woody patch types indicates that C and N sequestration potentials are species dependent.

Nomenclature: Jones et al. (1997).

Abbreviations: SOC = soil organic carbon; SOM = soil organic matter.

We examined the changes in prevalence of nine plant traits – including the presence of woody stem tissue, leaf longevity, nitrogen fixation, seed longevity, dispersal vector, pollination vector, and clonal growth form – across a riparian/forest-understory ecotone. This ecotone, found along headwater streams in boreal mixed-wood forests, supports four distinct vegetation zones: streambank, riparian, transition, and upland forest understory. The objective of this study was to identify specific trait patterns that may indicate functional responses to the changes in environmental factors such as nutrient availability and wind exposure that occur across the ecotone. The suites of plant species traits found in each zone were distinct, with a strong change in the prevalence of several traits. Wind and insect pollination, wind and vertebrate diaspore dispersal, and deciduous and evergreen leaves showed the greatest change in prevalence between the vegetation types. Some traits, including insect pollination and vertebrate diaspore dispersal, were strongly correlated within species. The consistent co-occurrence of pairs of traits in the same species suggests common responses by very different traits to the same environmental factor. This study demonstrates that an ecotone can be characterized not only as a discontinuity in species distributions or environmental factors, but also as a discontinuity in the trait spectrum. Examining ecotones from a trait perspective has strong potential for identifying the environmental factors and associated species functional responses that encourage the development of distinct vegetation boundaries.

Nomenclature: Newmaster et al. (1998).

We analysed the floristic turnover, or β-diversity patterns, of woody legumes in Baja California, the 1300 km long peninsula in NW Mexico. From the specimen labels of nine herbaria, we digitized 4205 records from 78 species and projected them onto a grid of 205 cartographic cells (20′ longitude × 15′ latitude). We found that the E–W turnover from the Pacific coast to the Sea of Cortés was high (ca. 60% change from one cell to the next) and that it was significantly associated with topographic variation across the peninsula. An analysis of between-site similarities showed that floristic regions in the peninsula have well defined boundaries, and that floristic similarities tend to extend for long distances along the coastal corridor of the Sea of Cortés. Finally, a numerical classification of Baja California based on the floristic attributes of the cells separated the peninsula into five well defined regions that coincide very closely with regionalizations, previously proposed by various authors using descriptive methods, and with the areas of endemism described for the peninsula in a previous paper. We conclude that the ecological regions of woody legumes in the peninsula clearly reflect the two main environmental forces that act on it, namely, the 1300 km long temperate–tropical climatic gradient and the dramatic E–W gradient running from the Pacific Ocean ca. 100 km across the mountain range into the Sea of Cortés.

Nomenclature: Wiggins (1980), Hickman (1993), Skinner and Pavlik (1994), and International Legume Database & Information Service,  www.ildis.org/legumes.html.

Vegetation science has relied on untested paradigms relating to the shape of species response curves along environmental gradients. To advance in this field, we used the HOF approach to model response curves for 112 plant species along six environmental gradients and three ecoclines (as represented by DCA ordination axes) in SE Norwegian swamp forests. Response curve properties were summarized in three binary response variables: (1) model unimodal or monotonous (determinate) vs. indeterminate; (2) for determinate models, unimodal vs. monotonous and (3) for unimodal models, skewed vs. symmetric. We used logistic regression to test the influence, singly and jointly, of seven predictor variables on each of three response variables. Predictor variables included gradient type (environmental or ecocline) and length (compositional turnover); species category (vascular plant, moss, Sphagnum or hepatic), species frequency and richness, tolerance (the fraction of the gradient along which the species occurs) and position of species along each gradient. The probability for fitting a determinate model increased as the main occurrence of species approached gradient extremes and with increasing species tolerance and frequency and gradient length. Appearance of unimodal models was favoured by low species tolerance and disfavoured by closeness of species to gradient extremes. Appearance of skewed models was weakly related to predictors but was slightly favoured by species optima near gradient extremes. Contrary to the results of previous studies, species category, gradient type and variation in species richness along gradients did not contribute independently to model prediction. The overall best predictors of response curve shape were position along the gradient (relative to extremes) and tolerance; the latter also expressing gradient length in units of compositional turnover. This helps predicting species responses to gradients from gradient specific species properties. The low proportion of skewed response curves and the large variation of species response curves along all gradients indicate that skewed response curves is a smaller problem for the performance of ordination methods than often claimed. We find no evidence that DCA ordination increases the unimodality, or symmetry, of species response curves more than expected from the higher compositional turnover along ordination axes. Thus ordination axes may be appropriate proxies for ecoclines, applicable for use in species response modelling.

Abbreviations: GAM = Generalized Additive Model; GLM = Generalized linear model; HOF = Huisman-Olff-Fresco; LNMDS = Local Non-metric Multidimensional Scaling.

Nomenclature: Lid & Lid (1994); Frisvoll et al. (1995); Krog et al. (1994), except for Polytrichastrum G.L.Sm., which is not recognized as distinct from Polytrichum Hedw. Several groups of related taxa were treated collectively.

The montane rain forest in Chiapas, Mexico has been subject to continuous human disturbance, mainly by wood extraction, cattle grazing and fire. To address the effect of this kind of disturbance regime on regeneration, survival and growth of naturally established tree seedlings were evaluated during a 3-yr period in three forest stands which varied in frequency and intensity of human disturbance. Frequency of cattle browsing on transplanted seedlings of Cornus disciflora, Liquidambar styraciflua and Oreopanax xalapensis planted on different facing slopes was evaluated over an 18-mo period, their survival and growth was also determined. The density of seedlings, number of species and survival of naturally recruited tree seedlings decreased with increased intensity of disturbance. Only seedlings of Pinus and Quercus species were recorded in heavily disturbed conditions. Seedlings grew more vigorously under moderately disturbed than in undisturbed or heavily disturbed conditions. More extreme microclimatic variation (air temperature, relative humidity and solar radiation) was recorded in heavily vs moderately disturbed forest. Survival and growth of transplanted tree seedlings differed with species. Grazing damage varied among seedlings located on different facing slopes. Plants located on NE facing slopes survived better than those on other slope positions. The mean proportion of surviving individuals of L. styraciflua (80%) and C. disciflora (78%) was higher than those for O. xalapensis (60%). A higher number of C. disciflora and O. xalapensis seedlings were grazed than L. styraciflua in any slope orientation treatment. The results suggest that human disturbance negatively affects natural recruitment, reduces richness of tree seedlings and favours a small number of tree species.

Abbreviations: PF = Pinus forest; PPFD = Photosynthetic photon flux density; PQLF = Pinus-Quercus-Liquidambar forest; QPF = Quercus-Podocarpus forest; RDGR = Relative basal diameter growth rate; RHGR = Relative height growth rate; TMRF = Tropical montane rain forest.

Nomenclature: Breedlove (1981).

This study deals with the habitat restriction of Mammillaria pectinifera, a threatened cactus species, confined to a few low density localities of the Tehuacán valley in tropical Mexico. We analysed the patterns of presence/absence of M. pectinifera in relation to the presence/absence of 48 other plant species, and the variation of environmental factors in 120 sampling plots. A Principal Components Analysis revealed a clear segregation between plots with and without individuals of M. pectinifera. A classification analysis resulted in four groups: two with low prevalence and two with high prevalence of M. pectinifera. Paired comparisons between plots with and without M. pectinifera allowed the characterization of its patterns of occurrence related to the variation of environmental factors. M. pectinifera was found on deep alkaline soils with relatively high surface stoniness and high water retention capacity, showing low species richness compared with plots where it was absent. The limited distribution of M. pectinifera in the Tehuacán Valley seems to be related to particular requirements of this species, being restricted to certain suitable habitat patches. Nevertheless, it is likely that other aspects, such as poor dispersal and establishment abilities, or biotic interactions could be associated with the observed patterns.

Nomenclature: Dávila-Aranda et al. (1993).

We evaluated variability in cover estimation data obtained by (1) two sampling teams who double sampled plots and (2) one team that used two methods (line intercepts and visual estimation of cover classes) to characterize vegetation of herbaceous wetlands. Species richness and cover estimates were similar among teams and among methods, but one sampling team scored cover higher than the other. The line intercept technique yielded higher cover estimates but lower species richness estimates than the cover class method. Cluster analyses of plots revealed that 36% and 11% of plots sampled consecutively by two teams or using two methods, respectively, were similar enough in species composition and abundance to be paired together in the resulting clustering tree. Simplifying cover estimate data to presence/absence increased the similarity among both teams and methods at the plot scale. Teams were very similar in their overall characterization of sites when cover estimation data were used, as assessed by cluster analysis, but methods agreed best on their overall characterization of sites when only presence/absence data were considered. Differences in abundance estimates as well as pseudoturnover contribute to variability. For double sampled plots, pseudoturnover was 19.1%, but 57.7% of pseudo-turnover cases involved taxa with ≦ 0.5% cover while only 3.4% involved taxa with > 8% cover. We suggest that vegetation scientists incorporate quality control, calibrate observers and publish their results.

Branch direction in Carnegiea gigantea (saguaro) cacti affects PAR interception and CO₂ uptake, and may influence reception of photomorphogenic cues. Branch direction data were collected in 29 saguaro populations over Arizona to determine if the southerly bias observed in previous studies at two locales can be generalized to the American range of the saguaro, and to determine the effect of shading by nurse plants on branch direction. Locally observed S bias from previous studies can be extended and generalized to include the American saguaro range. Branches were statistically uniform across environmental variability regarding temperature and precipitation. The S bias was not found in saguaros shaded by large nurse plants, supporting the assertion that branch direction occurs, in part, to maximize receipt of insolation.

Abbreviations: OPCNM = Organ Pipe Cactus National Monument; SNP = Saguaro National Park.

Harsh conditions in arid and semi-arid environments make seedling establishment rare. Plant recruitment in arid environments often occurs only in years with above average rainfall or in safe sites under the canopy of nurse plants that provide shelter from high temperatures and low moisture. Associations of establishing seedlings with adult plants are referred to as nurse-protégé interactions and are thought to be commensalisms in which seedlings benefit from the micro-environment created by adult plants with no effect for the latter. This phenomenon is thought to be more frequent in harsh than in mild environments and appears to occur frequently in deserts and arid and semi-arid biomes. Here, we investigate whether nurse-protégé interactions are more common in arid environments by searching the published literature from the previous 92 years using the terms nurse plants, protégé plants, facilitation, nucleation and facultative mutualism. We then quantitatively compared these reports from arid zones to other environments. A total of 296 papers were found which referred to nurse-protégé interactions. More than half (158) focused on arid and semi-arid zones. This information was also used to explore hypotheses of potential causative forces that might have selected for such interactions in the arid zones such as seed trapping, nutrient, moisture, protection from browsing or trampling and support availability. Because of the large number of different nurse species (147, from 98 genera and 40 families) and protégé species (429, from 273 genera and 84 families), described across a diversity of environments, we suggest that there may be more than one causative factor selecting for nurse-protégé interactions in arid and semi-arid environments.

Nomenclature: See references in ^{(App. 1)}.

In order to investigate how scale (grain size) affects the relationships between species richness and environmental drivers (such as stress and disturbance), we collected 12 nested quadrats (from 0.25 m² to 1023 m²) from seven remnant oak savannas located in the floodplain of the Chippewa River in western Wisconsin, USA. Large and small-scale richness were not significantly correlated, suggesting that small-scale richness is not strongly controlled by sampling effects of the local species pool. Linear and curvilinear regressions between species richness and disturbance, canopy cover, biomass, and soil organic matter were dependent on sampling scale (grain size). Disturbance by fire was strongly related to richness at small scales, while tree canopy cover was strongly related to richness at larger scales. While there was some evidence suggesting the transition from disturbance to canopy effects occurs between 10 and 100 m², the transition was not particularly abrupt. The results cast doubt on the general importance of local species pools in affecting small-scale richness as well as our ability to make generalizations that do not explicity include scale.

Nomenclature: Gleason & Cronquist (1991).

Persistence by longevity has been rarely considered as an alternative to regeneration by seeding for plants showing multiple demographic strategies. We propose a conceptual model of multiple demographic strategies for long-lived plants in stable habitats, shifting from regeneration by seeding to persistence by longevity and/or vegetative reproduction, along gradients of abiotic stress or interspecific competition. Regeneration by seeding would be promoted under low abiotic stress or under low competition, whereas persistence by longevity and/or vegetative reproduction would predominate at high levels of abiotic stress or competition. We test this model with two threatened species of the Mediterranean region, the shrub Juniperus communis, a widely distributed species which maintains relict populations in the Mediterranean mountains thanks to great adult longevity and Pinguicula vallisneriifolia, a palaeo-endemic herb relying on a perennial habit and vegetative reproduction under drought imposed stress or high competition at late successional phases. As a main consequence, multiple demographic strategies enhance a plant's ability to exploit environmental heterogeneity at different spatial (patches, localities, regions within the species' distribution area) and temporal (individual life span, glacial-interglacial cycles) scales. The potential of multiple demographic dynamics based on persistence and regeneration must be considered as a major ecological trait determining the long-term viability of peripheral populations of relict species as well as the inertia against extinction of many threatened endemisms, thereby contributing to the maintenance of the high plant diversity characterizing the Mediterranean region.

VEGAN adds vegetation analysis functions to the general-purpose statistical program R. Both R and VEGAN can be downloaded for free. VEGAN implements several ordination methods, including Canonical Correspondence Analysis and Non-metric Multidimensional Scaling, vector fitting of environmental variables, randomization tests, and various other analyses of vegetation data. It can be used for large data. Graphical output can be customized using the R language's extensive graphics capabilities. VEGAN is appropriate for routine and research use, if you are willing to learn some R.

Abbreviation: MDS = Multidimensional Scaling.