Are islands more susceptible to plant invasion than continents? A test using Oxalis pes‐caprae L. in the western Mediterranean

Aim We tested the relative vulnerability of islands to Oxalis pes‐caprae L. invasion compared to mainland regions. Oxalis pes‐caprae is a South African annual geophyte that reproduces via bulbils, and has spread in many Mediterranean and temperate regions of the world where introduced. Our study is one of the first detailed regional analyses of the occurrence and local abundance of a non‐native plant. Methods We conducted an extensive survey (2000 sampling points) to examine local and coarse‐scale patterns in both the occurrence and abundance of O. pes‐caprae on islands and in neighbouring mainland regions of Spain. Location We analysed occurrence (number of samples where present) and abundance (percentage cover) on two Balearic Islands (Menorca and Mallorca) and in two mainland administrative provinces of Spain (Murcia and València). Results Oxalis pes‐caprae occurrence was consistently higher on islands. Occurrence varied among habitats, being the highest in tree groves and the lowest in forests and shrublands. It was never found in these two habitats on the mainland. Mean O. pes‐caprae abundance was greatest in tree groves on the mainland, and in field margins and old fields on the islands. However, in general there were not significant differences in local abundances between island and mainland locations. Main conclusions These findings suggest that local processes (such as the biotic resistance of plant communities) are less important than coarse‐scale phenomena (such as environmental driving forces) in explaining differences in the invasion patterns observed between islands and adjacent mainland regions. We suggest that O. pes‐caprae has occupied a larger proportion of available habitats on islands due to: (1) its strong dependence on domestic animal and human‐mediated dispersal which are probably greater on the islands than in mainland areas, and (2) the smaller area encompassed by islands that, over a comparable period of time, enables a greater proportion of available habitats to be colonized (and hence higher occurrence) than equivalent larger mainland areas.     

Does invasion by an alien plant species affect the soil seed bank?

Questions: How does invasion affect old‐field seed bank species richness, composition and density? How consistent are these effects across sites? Does the soil seed bank match vegetation structure in old‐fields? Location: Menorca, Balearic Islands, Spain, western Mediterranean basin. Methods: We monitored seed germination in soils from old‐fields that were both uninvaded and invaded (legacy effect) by the annual geophyte Oxalis pes‐caprae. We also added O. pes‐caprae bulbs to uninvaded soils to test O. pes‐caprae interference with seedling emergence (competitive effect). We compared species composition in the seed bank with that of the vegetation. Results: Species richness in the seed bank and in the vegetation was not significantly different between invaded and uninvaded areas. Uninvaded areas did not have larger seed banks than invaded areas. More seedlings, especially of geophytes, emerged when O. pes‐caprae bulbs were added to the soil. Species similarity between invaded and uninvaded areas was higher in the seed bank (74%) than in the vegetation (49%). Differences in species composition were as important as differences among sites. The degree of species similarity between the seed bank and the vegetation was very low (17%). Conclusions: Despite invasion by O. pes‐caprae not affecting species richness, the variation in the seed bank species composition in invaded and uninvaded areas, and the differences between the seed bank and the mature vegetation, highlights that even if the invader could be eradicated the vegetation could not be restored back to the exact composition as found in uninvaded areas.     

Sexual reproduction of the pentaploid,short‐styled Oxalis pes‐caprae allows the production of viable offspring

Reproduction is a key factor for the successful establishment and spread of introduced species. Oxalis pes‐caprae is a tristylous species with a self‐ and morph‐incompatibility sexual system that, in the invaded range of the western Mediterranean Basin, has been found to reproduce asexually because only the pentaploid, short‐styled morph (5x S‐morph) was introduced. The objective of this study was to test the ability of the 5x S‐morph of O. pes‐caprae to produce viable offspring in the absence of compatible mates, exploring the hypothesis that new morphs could have emerged by sexual reproduction events of the initially introduced morph. Pollen germination, pollen tube development, fruit and seed production, seed germination and offspring ploidy levels were analysed after controlled hand‐pollinations to assess self‐ and morph‐incompatibility and production of viable gametes by the 5x S‐morph. The self‐incompatibility system is still operating, but a partial breakdown in the morph‐incompatibility system combined with the production of viable gametes was observed, allowing sexual reproduction of the 5x S‐morph in the invaded range. The ability of the 5x S‐morph to reproduce sexually may have major consequences for the dynamics of invasive populations of O. pes‐caprae and could be one of the factors involved in the occurrence of new floral morphs in this invaded range.     

Spatial heterogeneity in the determinants of woody plant invasion of lowland heath

Questions: 1. What is the scale and extent of spatial variability in factors affecting Betula invasion of heaths? 2. How much effect does each factor have on within‐patch patterns of invasion? 3. How can this understanding aid in managing Betula invasions? Location: Lowland heath of southern England. Methods: Determinants of Betula (both B. pubescens and B. pendula) invasion: biomass density, necromass density, mean vegetation height, P‐availability, soil water content and total Betula seed bank density, were measured at two sites on a 5‐ha sampling grid. Spatial pattern was assessed using geostatistics. Contributions of each determinant to within‐site heterogeneity in predicted Betula seedling densities were estimated by varying variables over their full and interquartile ranges in a statistical model derived from experimental data. Results: Salient spatial trends were revealed: strong autocorrelation over distances of < 50 m for soil factors and more extensive autocorrelation (0 to > 150 m) in vegetation variables and Betula seed bank densities. The latter resulted in single across‐site gradients, the former small, distinct patches. All patterns were overlain with variance that was present at distances of < 17.6 m. Variables displaying spatial pattern also accounted for within‐site heterogeneity in predicted Betula seedling densities but their relative contribution to this varied between sites. Conclusions: Identifiable spatial autocorrelation in factors controlling patch‐scale invasion patterns allows managers to target invasion prone patches, potentially reducing management intensities. Furthermore, management effort may be optimised by spatially de‐coupling Betula seed from safe‐sites. This plan may adaptable to the management of other weeds and open‐land ecosystems.     

Population dynamics of the West Indian sweetpotato weevil Euscepes postfasciatus (Fairmaire): a simulation analysis

The West Indian sweetpotato weevil Euscepes postfasciatus (Fairmaire) is a major pest of the sweet potato Ipomoea batatas (L.) Lam. and this weevil is a target of an eradication program using the Sterile Insect Technique in Okinawa Prefecture, Japan. Understanding the population ecology is essential in the planning of an eradication program; hence, a host‐plant infestation survey and light trap survey have been conducted to monitor the population dynamics of the weevil on Kume Island (Okinawa Prefecture), which is the target area of the trial weevil eradication project. Seasonal tendencies of weevil density were found in these field surveys, but the tendency found in the host‐plant infestation survey was not seen every year, and the effectiveness of the light trap is somewhat suspect. To confirm the reliability of the tendency observed in these field surveys, the present study attempted to explain the tendency by a seasonal temperature change using a temperature‐based model of weevil population dynamics. The seasonal changes of weevil density differed according to host plants and host‐plant fields. The seasonal changes of weevil density inside the host plant Ipomoea indica and outside the host plants in I. indica fields were consistent with those predicted by the model. However, those inside the host plant Ipomoea pes‐caprae in the host‐plant infestation survey were contrary to the predicted ones, and those observed outside host plants in I. pes‐caprae fields by the light trap survey were not in good agreement with the predicted ones. It was concluded that the seasonal change of the weevil density observed in I. indica and I. indica fields can be explained by a seasonal temperature change, but factors other than seasonal temperature change are needed to explain those in I. pes‐caprae and I. pes‐caprae fields.     

Phenyl Cinnamate Derivatives from Oxalis pes‐caprae

During the screening of Mediterranean invasive plants, Oxalis pes‐caprae was identified as promising species. The fresh leaves and twigs of the plant were crumbled and extracted with AcOEt. The solution was concentrated, and separated into acidic and neutral fractions. The crude neutral residue was fractionated by chromatographic procedures, followed by structure elucidation on the basis of ¹H‐ and ¹³C‐NMR, and MS data analysis, and six new phenyl cinnamate derivatives were identified. The phytotoxic effects of the isolated compounds on the germination and growth of the dicotyledon Lactuca sativa L. (lettuce) were studied in the concentration range from 10^?4 to 10^?7 M .     

108 years of change in spatial pattern following selective harvest of a Pinus ponderosa stand in northern Arizona,USA

Questions: How did an initial tree harvest in 1894 influence the spatial and temporal patterns of Pinus ponderosa recruitment? How do these patterns compare to our understanding of P. ponderosa stand dynamics prior to Euro‐American settlement? How might spatial pattern information, particularly with respect to patch characteristics, inform current restoration and management practices? Location: A 2.59‐ha permanent sample plot in the Fort Valley Experimental Forest, Flagstaff, Arizona. The plot was selectively harvested in 1894 and measured in 1909 and 2002. Methods: We used historical stem‐map and ledger data, contemporary data, and dendrochronological techniques to reconstruct stand structure (tree size, age, location) in three scenarios: (1) unharvested (1909), (2) harvested (1909), and (3) contemporary (2002). We used Clark and Evans' R, Ripley's K(t) univariate analysis, and correlogram analysis to assess the spatial pattern in each scenario. We also used Ripley's K₁₂(t) bivariate analysis and tree age data to examine spatial and temporal recruitment patterns as observed in the contemporary scenario. Results and Conclusions: The unharvested stand was aggregated at scales up to 28 m. The selective harvest accentuated the spatial patchiness of the stand in 1909 and changed spatial patterns by homogenizing tree size within patches. By 2002, the stand was a single patch dominated by small trees. Post‐harvest recruitment patterns were not spatially random; Pinus seedlings initially established in natural grass openings and then proceeded to fill‐in stump patches created by harvesting. Knowledge of spatial pattern should be explicitly incorporated into restoration activities in these forests.     

Spatial and Temporal Dynamics of a Restored Population of Oryza rufipogon in Huli Marsh, South China

Studying the process of population restoration is helpful for managing and preserving endangered species. A population of Oryza rufipogon (wild rice), an endangered species, was reintroduced in 1993 into Huli Marsh. We conducted a detailed survey over a 5‐year period (1997–2001) to evaluate the present status of the population and to further our understanding of its habitat requirements and the population model. The population was surveyed using 2 × 2–m quadrats in mid‐September of each year. In total, 2,683 quadrats were surveyed covering the whole O. rufipogon reserve during each survey. The population's spatial distribution was mapped, and the maps were used to examine the relationship between patch replacement and water depth. The individual number of O. rufipogon increased steadily from 1993 to 2001. The patch number, patch area, mean patch size, and largest patch size increased over this time period, and Korcak patchiness exponents decreased. On average, 83% of the patches persisted from one year to the next. There was a significant positive correlation between the initial patch size and the size the following year. The probability of patch disappearance decreased as patch size increased. Fifty‐eight percent of the patches were located at water depths between 20 and 30 cm. Water depth had no significant effect on the patch transition from O. rufipogon to other species. The loss and gain of O. rufipogon patches were statistically correlated with the patch areas in different water depths. Our results show that the population of O. rufipogon can successfully be reintroduced to the original habitat after appropriate environmental conditions have been restored. We recommend the following transplantation practices: transplant many smaller patches rather than a few larger patches, use transplant patch sizes of at least 20 m², and transplant into sites vegetated with species with different regeneration niches from the transplanted species.     

Structural segregation and scales of spatial dependency in Abies amabilis forests

Question: Is a mosaic structure apparent in the spatial distribution of trees in old‐growth Abies amabilis forests? Location: Montane forests of the western Cascade Range, Washington, USA. Methods: Maps of tree locations were created for study areas located in two, 300‐year old stands and a single 600‐year old stand. Stand structure parameters were calculated using several subsample quadrats sizes (56.25 ‐ 306.25 m²), which were drawn randomly with replacement at a density of 250 quadrats per ha from the stem maps in the computing environment. Spatial cross‐covariance functions between different canopy strata were estimated using the spline cross‐correlogram. Results: Negative spatial correlation (segregation) between subcanopy tree density and areas of high overstorey occupancy was detected. Understorey and midstorey tree densities were positively spatially correlated. These general trends were apparent across the range of observational scales investigated. Significant spatial correlation between canopy strata was observed at spatial scales of 12 ‐ 44 m and extended to the largest scales in the 600‐year old stand. Conclusion: The observed spatial segregation between canopy strata supports the hypothesis that old A. amabilis forests form fine‐scale structural mosaics. Structural segregation at small scales may be due to competitive interactions as well as exogenous forcing of tree locations (e.g. by mortality due to pathogens or disturbance), however segregation at large scales in the 600‐year old stand is likely due to exogenous factors alone. This study reinforces the idea that horizontal heterogeneity is an emergent property of old‐growth forests.     

Widespread resistance of Mediterranean island ecosystems to the establishment of three alien species

Although some invasive plants are cosmopolitan, not all ecosystems are invaded to the same degree. Yet there is little experimental work on how ecosystem resistance to invasion at the establishment phase differs among ecosystems. We conducted two field sowing experiments in two consecutive years to examine establishment of the deciduous tree Ailanthus altissima, the succulent subshrub Carpobrotus spp. and the annual geophyte Oxalis pes‐caprae in coastal dunes, shrublands and oldfields in more than 200 sites across six Mediterranean Basin islands differing in climatic conditions and local species richness. Establishment success (i.e. percentage of plots with at least one seedling) and rates (i.e. seedling to sown seed ratio) were low, especially for Ailanthus even when accounting for differences in seed viability. Oxalis was capable of producing a new cohort of seedlings the year following planting. By contrast, all Ailanthus seedlings and half the Carpobrotus seedlings died following the first summer. Differences in establishment success and rates among ecosystems were species‐, island‐ and year‐dependent. Differences in precipitation and mean temperature were associated with differences in establishment rates across sites. Establishment rates tended to be positively correlated with cumulative precipitation and negatively with mean T^a. Unexpectedly, native species richness was not a good predictor of seedling establishment, except for higher Oxalis establishment success in species rich habitats. By conducting field sowing tests at multiple sites across a region we found that except for Oxalis, Mediterranean island ecosystems are quite resistant to invader establishment. These results suggest that differences in the degree of invasion between ecosystems and islands might be more dependent upon the influence of invasion event factors (e.g. propagule pressure) or factors acting at a later life‐history stages rather than differences in the resistance imposed by ecosystems to invader recruitment. Moreover, our results support the notion that in Mediterranean ecosystems invasions are highly idiosyncratic events and strongly dependent on water availability conditions.     

The spatio‐temporal forest patch dynamics inferred from the fine‐scale synchronicity in growth chronology

Question: Abrupt increments in tree radial growth chronology are associated with gap formations derived from disturbances. If a forest has been primarily controlled by fine‐scale disturbances such as single tree‐fall, do these release events spatio‐temporally synchronize at a fine scale such as 10 m and 5 years? Is it possible to quantify spatio‐temporal patterns of synchronicity from tree rings and long‐term inventories, and associate them with spatial forest patch dynamics? How and to what extent can we reconstruct the fine‐scale synchronized growth and spatio‐temporal forest patch dynamics from currently available information? Location: Cores were taken from Abies sachalinensis trees in a coniferous/deciduous mixed forest in the Shiretoko Peninsula, Hokkaido, northern Japan. Methods: We first eliminated short‐term fluctuations and highlighted growth trends over the mid‐term using a time‐series smoothing technique. This helped identify release events, we then conducted fine‐scale spatial analyses on released A. sachalinensis primarily with cluster analysis. Results: We specified the unit scale of synchronicity at 10 m, and classified released A. sachalinensis trees into spatially separated regions. Only once during the recent 50 years was extensive synchronicity over 40 m found. Most of the released A. sachalinensis were isolated, with non‐released A. sachalinensis present in nearby, implying imperfect synchronization. The ambiguous 20–30 m A. sachalinensis patches present in the current forest were the result of connected and overlapping patches smaller than 10 m associated with different disturbances and different responses of understorey trees. Conclusion: Tree‐ring series, long‐term census and fine‐scale spatio‐temporal analyses revealed that this forest community has been controlled by two types of disturbance: frequent small disturbances such as single tree‐fall and less frequent multiple tree‐falls.     

Diminishing Spatial Heterogeneity in Soil Organic Matter across a Prairie Restoration Chronosequence

Habitat restoration resulting in changes in plant community composition or species dominance can affect the spatial pattern and variability of soil nutrients. Questions about how these changes in soil spatial heterogeneity develop over time at restoration sites, however, remain unaddressed. In this study, a geostatistical approach was used to quantify changes over time in the spatial heterogeneity of soil organic matter (SOM) across a 26‐year chronosequence of tallgrass prairie restoration sites at FermiLab, outside of Chicago, Illinois. We used total soil N and C as an index of the quantity of SOM. We also examined changes in C:N ratio, which can influence the turnover of SOM. Specifically, the spatial structure of total N, total C, and C:N ratio in the top 10 cm of soil was quantified at a macroscale (minimum spacing of 1.5 m) and a microscale (minimum spacing of 0.2 m). The magnitude of spatial heterogeneity (MSH) was characterized as the proportion of total sample variation explained by spatially structured variation. At the macroscale, the MSH for total N decreased with time since restoration (r²= 0.99, p < 0.001). The decrease in spatial heterogeneity over time corresponded with a significant increase in the dominance of the C₄ grasses. At the microscale, there was significant spatial structure for total N at the 4‐year‐old, 16‐year‐old, and 26‐year‐old sites, and significant spatial structure for total C at the 16‐year‐old and 26‐year‐old sites. These results suggest that an increase in dominance of C₄ grasses across the chronosequence is homogenizing organic matter variability at the field scale while creating fine‐scale patterns associated with the spacing of vegetation. Areas of higher soil moisture were associated with higher soil N and C at the two oldest restoration sites and at the native prairie site, potentially suggesting patches of increased belowground productivity in areas of higher soil moisture. This study is one of the first to report significant changes over time in the spatial structure of organic matter in response to successional changes initiated by restoration.     

Exploring spatiotemporal patterns in early stages of primary succession on former lignite mining sites

Questions: 1. Does random colonization predominate in early stages of primary succession? 2. Do pioneer species facilitate the establishment of later arriving species? 3. Does an initially random distribution change to an aggregated pattern with ongoing succession? Location: Lignite mining region of Lower Lusatia, eastern Germany. Methods: Individual plants were mapped along a 2 m × 28 m transect during three successive years and classified into two groups (1) the pioneer Corynephorus canescens and (2)‘all other species’. Using the pair‐correlation function, univariate point pattern analysis was carried out by applying a heterogeneous Poisson process as null model. Bivariate analysis and a toroidal shift null model were applied to test for independence between the spatial patterns of the two groups separately for each year, as well by exploring spatiotemporal patterns from different years. Results: In the first year Corynephorus and ‘all other species’ showed an aggregated pattern on a spatial scale > 40 cm and in the second and third years a significant attraction for distances between 4 and 12 cm, with an increasing radius in the third year. The analyses of interspecific spatiotemporal dynamics revealed a change from independence to attraction between distances of 4 cm and 16 cm when using Corynephorus as focal species. However, applying ‘all other species’ as focal points results in a significant attraction at distances up to 60 cm in the first year and a diminishing attraction in the second and third years with distances ≤ 6 cm. Conclusions: Facilitative species‐species interactions are present in early stages of primary succession, resulting mainly from pioneer species acting as physical barriers and their ability to capture diaspores being drifted by secondary dispersal along the substrate surface. However, due to gradual establishment of perennial species and their ability of lateral extension by vegetative dispersal, facilitation may influence spatial pattern formation predominantly on short temporal and fine spatial scales.     

Within‐stand spatial structure and relation of boreal canopy and understorey vegetation

Question: How do spatial patterns and associations of canopy and understorey vegetation vary with spatial scale along a gradient of canopy composition in boreal mixed‐wood forests, from younger Aspen stands dominated by Populus tremuloides and P. balsamifera to older Mixed and Conifer stands dominated by Picea glauca? Do canopy evergreen conifers and broad‐leaved deciduous trees differ in their spatial relationships with understorey vegetation? Location: EMEND experimental site, Alberta, Canada. Methods: Canopy and understorey vegetation were sampled in 28 transects of 100 contiguous 0.5 m × 0.5 m quadrats in three forest stand types. Vegetation spatial patterns and relationships were analysed using wavelets. Results: Boreal mixed‐wood canopy and understorey vegetation are patchily distributed at a range of small spatial scales. The scale of canopy and understorey spatial patterns generally increased with increasing conifer presence in the canopy. Associations between canopy and understorey were highly variable among stand types, transects and spatial scales. Understorey vascular plant cover was generally positively associated with canopy deciduous tree cover and negatively associated with canopy conifer tree cover at spatial scales from 5–15 m. Understorey non‐vascular plant cover and community composition were more variable in their relationships with canopy cover, showing both positive and negative associations at a range of spatial scales. Conclusions: The spatial structure and relation of boreal mixed‐wood canopy and understorey vegetation varied with spatial scale. Differences in understorey spatial structure among stand types were consistent with a nucleation model of patch dynamics during succession in boreal mixed‐wood forests.     

Spatial correlations of Diceroprocta apache and its host plants: evidence for a negative impact from Tamarix invasion

Abstract 1. The hypothesis that the habitat‐scale spatial distribution of the Apache cicada Diceroprocta apache Davis is unaffected by the presence of the invasive exotic saltcedar Tamarix ramosissima was tested using data from 205 1‐m² quadrats placed within the flood‐plain of the Bill Williams River, Arizona, U.S.A. Spatial dependencies within and between cicada density and habitat variables were estimated using Moran's I and its bivariate analogue to discern patterns and associations at spatial scales from 1 to 30 m. 2. Apache cicadas were spatially aggregated in high‐density clusters averaging 3 m in diameter. A positive association between cicada density, estimated by exuvial density, and the per cent canopy cover of a native tree, Goodding's willow Salix gooddingii, was detected in a non‐spatial correlation analysis. No non‐spatial association between cicada density and saltcedar canopy cover was detected. 3. Tests for spatial cross‐correlation using the bivariate I_YZ indicated the presence of a broad‐scale negative association between cicada density and saltcedar canopy cover. This result suggests that large continuous stands of saltcedar are associated with reduced cicada density. In contrast, positive associations detected at spatial scales larger than individual quadrats suggested a spill‐over of high cicada density from areas featuring Goodding's willow canopy into surrounding saltcedar monoculture. 4. Taken together and considered in light of the Apache cicada's polyphagous habits, the observed spatial patterns suggest that broad‐scale factors such as canopy heterogeneity affect cicada habitat use more than host plant selection. This has implications for management of lower Colorado River riparian woodlands to promote cicada presence and density through maintenance or creation of stands of native trees as well as manipulation of the characteristically dense and homogeneous saltcedar canopies.     

Weed species differ in their ability to emerge in no‐till systems that include cover crops

No‐till cropping systems that include cover crops could lead to important changes in weed communities by decreasing some annual weed populations. In this study, we predicted that seed burial depth and the presence of cover crop would affect the emergence and initial growth success of annual weed species. We tested two factors on 14 weed species in a greenhouse: the seed burial depth of weeds (buried versus soil surface) and the presence/absence of a cover crop (ryegrass). We counted the emerged seedlings and measured the height of weeds and cover crops (Hweed, Hcover), the dry matter content of weeds and cover crops (DMCweed, DMCcover) and the number of leaves of weeds (NLweed) on 1433 weed and 390 ryegrass individuals. Emergence of five weed species (AMBEL, ANGAR, BROST, CENCY and EPHHE) was affected by the seed location (?10.3% on average for unburied seeds), five other weed species (ALOMY, CAPBP, SONAS, VERPE and VLPMY) were affected by cover (on average ?9.5% for seeds emerged in the presence of cover crop), and four weed species (GERDI, LAMPU, POAAN and VIOAR) were not affected by either. Weed growth of all weed species also decreased with the presence of a cover crop (on average Hweed: ?49.9%, DMCweed: ?87.2% and NLweed: ?55.4%) and for unburied seeds (on average Hweed: ?33.7%, DMCweed: ?70.6% and NLweed: ?43.3%), with various responses according to species. This study indicates that annual weeds could be disadvantaged by no‐till systems using cover crops.     

A grid‐based method for sampling and analysing spatially ambiguous plants

Abstract. Spatial data can provide much information about the interrelations of plants and the relationship between individuals and the environment. Spatially ambiguous plants, i.e. plants without readily identifiable loci, and plants that are profusely abundant, present non‐trivial impediments to the collection and analysis of vegetation data derived from standard spatial sampling techniques. Sampling with grids of presence/absence quadrats can ameliorate much of this difficulty. Our analysis of 10 fully‐mapped grassland plots demonstrates the applicability of the grid‐based approach which revealed spatial dependence at a much lower sampling effort than mapping each plant. Ripley's K‐function, a test commonly used for point patterns, was effective for pattern analysis on the grids and the gridded quadrat technique was an effective tool for quantifying spatial patterns. The addition of spatial pattern measures should allow for better comparisons of vegetation structure between sites, instead of sole reliance on species composition data.     

Clustered animal burrows yield higher spatial heterogeneity

An understanding of the relationships between spatial heterogeneity and disturbance regime is important for establishing the mechanisms necessary to maintain biodiversity. Our objective was to examine how the configuration of disturbance by burrowing rodents (Siberian marmot) affected the spatial heterogeneity of vegetation and soil nutrient properties. We established three 2500-m² (50 m × 50 m) isolated-burrows plots and three 2500-m² clustered-burrows plots in a Mongolian grassland. Each plot was subdivided into 4-m² quadrats, and the plant species richness, percent coverage, and soil nutrient properties in the quadrats were surveyed. Spatial heterogeneity was calculated for vegetation using the mean dissimilarity of species composition among sample quadrats, and geostatistical analysis was used to calculate soil properties. Heterogeneous patches of plants such as Achnatherum splendens and higher nutrient concentrations were found only near the clustered burrows. As a result, spatial heterogeneities of vegetation and soil nutrient properties were higher in the clustered colony than those in the isolated colony. The configuration of disturbance patches affected the spatial heterogeneity at the landscape level through the spatial pattern of disturbance frequency.     

A test for spatial relationships between neighbouring plants in plots of heterogeneous plant density

Abstract. Maps of plant individuals in (x, y) coordinates (i.e. point patterns) are currently analysed through statistical methods assuming a homogeneous distribution of points, and thus a constant density within the study area. Such an assumption is seldom met at the scale of a field plot whilst delineating less heterogeneous subplots is not always easy or pertinent. In this paper we advocate local tests carried out in quadrats partitioning the plot and having a size objectively determined via a trade‐off between squared bias and variance. In each quadrat, the observed pattern of points is tested against complete spatial randomness (CSR) through a classical Monte‐Carlo approach and one of the usual statistics. Local tests yield maps of p‐values that are amenable to diversified subsequent analyses, such as computation of a variogram or comparison with co‐variates. Another possibility uses the frequency distribution of p‐values to test the whole point pattern against the null hypothesis of an inhomogeneous Poisson process. The method was demonstrated by considering computer‐generated inhomoge‐neous point patterns as well as maps of woody individuals in banded vegetation (tiger bush) in semi‐arid West Africa. Local tests proved able to properly depict spatial relationships between neighbours in spite of heterogeneity/clustering at larger scales. The method is also relevant to investigate interaction between density and spatial pattern in the presence of resource gradients.     

Patterns of resource tracking by avian frugivores at multiple spatial scales: two case studies on discordance among scales

Scaling is relevant for the analysis of plant‐frugivore interaction, since the ecological and evolutionary outcomes of seed dispersal depend on the spatial and temporal scale at which frugivory patterns emerge. We analyse the relationship between fruit abundance and frugivore activity at local and landscape spatial scales in two different systems composed, respectively, by the bird‐dispersed woody plants Juniperus communis and Bursera fagaroides, and their frugivore assemblages. We use a hierarchical approach of nested patchiness of fruit‐resource, where patches are defined by individual plants within site, at the local scale, and by sites within region, at the landscape scale. The structure of patches is also described in terms of contrast (differences in fruit availability among patches) and aggregation (spatial distribution of patches). For J. communis, frugivore activity was positively related to fruit availability at the landscape scale, this pattern seldom emerging at the local scale; conversely, B. fagaroides showed a general trend of positive local pattern that disappeared at the landscape scale. These particular trends might be partially explained by differences in contrast and aggregation. The strong contrast among plants within site together with a high aggregation among sites would promote the B. fagaroides pattern to be only local, whereas in J. communis, low aggregation among sites within region would favour a sharp landscape‐scale pattern. Both systems showed discordant patterns of fruit‐resource tracking among consecutive spatial scales, but the sense of discordance differed among systems. These results, and the available multi‐scale frugivory data, suggest that discordance among successive scales allows to link directly frugivory patterns to resource‐tracking mechanisms acting at particular scales, resulting, thus, more informative than concordance observational data, in which landscape patterns might result from accumulated effect of local mechanisms. In this context, we propose new methodological approaches for a better understanding of the hierarchical behavioural mechanisms underpinning the multi‐scale resource tracking by frugivores.