Species–area relationships in Mediterranean-climate plant communities

Aim To determine the best‐fit model of species–area relationships for Mediterranean‐type plant communities and evaluate how community structure affects these species–area models. Location Data were collected from California shrublands and woodlands and compared with literature reports for other Mediterranean‐climate regions. Methods The number of species was recorded from 1, 100 and 1000 m² nested plots. Best fit to the power model or exponential model was determined by comparing adjusted r² values from the least squares regression, pattern of residuals, homoscedasticity across scales, and semi‐log slopes at 1–100 m² and 100–1000 m². Dominance–diversity curves were tested for fit to the lognormal model, MacArthur's broken stick model, and the geometric and harmonic series. Results Early successional Western Australia and California shrublands represented the extremes and provide an interesting contrast as the exponential model was the best fit for the former, and the power model for the latter, despite similar total species richness. We hypothesize that structural differences in these communities account for the different species–area curves and are tied to patterns of dominance, equitability and life form distribution. Dominance–diversity relationships for Western Australian heathlands exhibited a close fit to MacArthur's broken stick model, indicating more equitable distribution of species. In contrast, Californian shrublands, both postfire and mature stands, were best fit by the geometric model indicating strong dominance and many minor subordinate species. These regions differ in life form distribution, with annuals being a major component of diversity in early successional Californian shrublands although they are largely lacking in mature stands. Both young and old Australian heathlands are dominated by perennials, and annuals are largely absent. Inherent in all of these ecosystems is cyclical disequilibrium caused by periodic fires. The potential for community reassembly is greater in Californian shrublands where only a quarter of the flora resprout, whereas three quarters resprout in Australian heathlands. Other Californian vegetation types sampled include coniferous forests, oak savannas and desert scrub, and demonstrate that different community structures may lead to a similar species–area relationship. Dominance–diversity relationships for coniferous forests closely follow a geometric model whereas associated oak savannas show a close fit to the lognormal model. However, for both communities, species–area curves fit a power model. The primary driver appears to be the presence of annuals. Desert scrub communities illustrate dramatic changes in both species diversity and dominance–diversity relationships in high and low rainfall years, because of the disappearance of annuals in drought years. Main conclusions Species–area curves for immature shrublands in California and the majority of Mediterranean plant communities fit a power function model. Exceptions that fit the exponential model are not because of sampling error or scaling effects, rather structural differences in these communities provide plausible explanations. The exponential species–area model may arise in more than one way. In the highly diverse Australian heathlands it results from a rapid increase in species richness at small scales. In mature California shrublands it results from very depauperate richness at the community scale. In both instances the exponential model is tied to a preponderance of perennials and paucity of annuals. For communities fit by a power model, coefficients z and log c exhibit a number of significant correlations with other diversity parameters, suggesting that they have some predictive value in ecological communities.     

Scrub communities along a climatic gradient in the southern Balkans: maquis,pseudomaquis and shibljak

The research deals with three scrub communities/formations in the southern Balkans, maquis, pseudomaquis and shibljak, and their changes along the main (macro)ecological gradient. We performed DCA analysis and, since the first axis correlates with climatic data, the projection of relevés on this axis was accepted as proxy for the (macro)ecological gradient, and the species turnover along this gradient was elaborated. Linear regression was used to test the turnover of life forms, chorotypes and the life strategy of species (syntaxonomical affiliation). It was found that life forms, except hemicryptophytes and nanophanerophytes, do not change significantly along the gradient change; most chorotypes change significantly; among species, life strategies (syntaxonomical affiliation), the proportion of species of deciduous and evergreen forests and scrub, as well as species of dry grassland, change significantly. The classification established three clusters, representing three formations and we tested differences among them. It was established that stenomediterranean and Eurasian species and species of perennial grasslands differentiate all three clusters and some groups only two of them. We found that discontinuous formations can be established along a continuous gradient. Communities were also classified within the present synsystematic system.     

Climate and land use change impacts on Mediterranean high-mountain vegetation in the Apennines since the 1950s

Background: High-mountain ecosystems are centres of plant diversity that are particularly sensitive to land-use and climate change.

Aims: We investigated the ecological trends associated with land use and climate change since the 1950s in different vegetation types in high-mountain habitats in the central Apennines.

Methods: We analysed temporal changes in: Pinus mugo scrub, calcareous subalpine grasslands and alpine scree vegetation, comparing historical and recent vegetation records from vegetation plots from two periods (1955–1980 and 1990–2014) for their ecological indicator values (Landolt temperature and nutrient indicators) and structural traits (growth forms) over time using generalised linear models (GLMs).

Results: We observed significant temporal differences in the ecology and structure of the analysed habitats. In the Pinus mugo scrub we detected a reduction of subalpine and herbaceous species and in calcareous alpine screes we observed an increment of the lower montane, montane and subalpine species and of dwarf shrubs. Conversely, subalpine grasslands were stable over time.

Conclusions: Ecological changes that have occurred in the Central Apennines, following changes in type and intensity of land use and recent warming are consistent with those observed in other European mountains, for which climate and land-use changes are claimed as the main driving forces.     

Shrubland formations and associations in mediterranean-desert transitional zones of northwestern Baja California

The area between Ensenada and EI Rosario (Baja California, Mexico) has long been considered as a transitional zone in which two great ecoclimatic regions (Mediterranean and Tropical-Desert) overlap. The floristic and biotypical diversity of this area was evaluated by analyzing its shrubland formations from a phytosociological point of view. This phytosociological study, carried out according to the Braun-Blanquet method and supported by cluster analysis, describes sixteen shrubland associations from Northwestern Baja California.Floristic diversity of the transitional zone was evaluated using two indices, endemic value (EV) and endemic community value (ECV), which are related to the degree of endemism in the flora and plant associations. The phytosociological analysis showed that the high number of shrubland associations found in this area reflected its transitional character. The closer the associations are to the transitional zone, the higher their biotypical and floristic diversity.Abbreviations ADE Adenostoma fasciculatum - AES Fraxinus trifoliata-Aesculus parryi - BER Bergerocactus emoryi-Agave shawii - DES Echinocereus engelmannii-Agave deserti - ENC Viguiera deltoidea-Encelia asperifolia - EUR Eurotia lanata-Yucca schidigera - FOU Agave cerulata-Fouquieria columnaris - FRA Atriplex julacea-Frankenia palmeri - HYM Baccharis glutinosa-Hymenoclea monogyra - KEC Clematis lasiantha-Keckiella antirrhinoides - LAR Ambrosia chenopodifolia-Larrea tridentata - LYC Ephedra californica-Lycium brevipes - MAH Malosma laurina-Heteromeles arbutifolia - MUN Salvia munzii-Artemisia californica - ROS Rosa minutifolia-Aesculus parryi - SAL Salvia apiana-Viguiera laciniata     

Effects of elevated atmospheric CO2 on fine root length and distribution in an oak-palmetto scrub ecosystem in central Florida

Atmospheric CO₂ concentration is rising and it has been suggested that a portion of the additional carbon is being sequestered in terrestrial vegetation and much of that in below-ground structures. The objective of the present study was to quantify the effects of elevated atmospheric CO₂ on fine root length and distribution with depth with minirhizotrons in an open-top chamber experiment in an oak-palmetto scrub ecosystem at Kennedy Space Centre, Florida, USA. Observations were made five times over a period of one and a half years in three ambient chambers (350 p.p.m. CO₂), three CO₂ enriched chambers (700 p.p.m. CO₂), and three unchambered plots. Greater root length densities were produced in the elevated CO₂ chambers (14.2 mm cm^?2) compared to the ambient chambers (8.7 mm cm^?2). More roots may presumably lead to more efficient acquisition of resources. Fine root abundance varied significantly with soil depth, and there appeared to be enhanced proliferation of fine roots near the surface (0–12 cm) and at greater depth (49–61 cm) in the elevated CO₂ chambers. The vertical root distribution pattern may be a response to availability of nutrients and water. More studies are needed to determine if increased root length under CO₂ enriched conditions actually results in greater sequestering of carbon below ground.     

Xerie Mediterranean-type shrubland associations of Alta and Baja California and the community/continuum debate

A survey of the xeric shrublands of Pacific coastal North America from San Francisco to El Rosario (Mexico), including the inner Channel Islands, was conducted using 99 sample sites of 0.063 ha size. TWINSPAN classification and DECORANA ordination confirmed the existence of two plant formations, distinguishable physiognomically: coastal sage scrub and coastal succulent scrub. Within coastal sage scrub, four floristic associations were recognized: Diablan, Venturan, Riversidian and Diegan. Within coastal succulent scrub, two floristic associations were defined: Martirian and Vizcainan. These associations occur in distinct geographical regions following the coastline, with the Riversidian association occurring in the basin inland from Venturan and Diegan regions. Their locations are strongly correlated with differences in evapotranspirative stress regimes. The Channel Island sites show affinities to several of the mainland associations. The Venturan association can be further subdivided floristically into two subassociations, dominated by Salvia mellifera and S. leucophylla respectively. These subassociations which are coextensive geographically at a regional scale, typically do not intermingle at a local scale but often meet along sharp boundaries in the landscape. The dominant species segregate by moisture preference, S. mellifera preferring coarser-texture soils and more southerly aspects than S. leucophylla. Richness and equitability of these sites are depressed relative to other xeric shrubland sites, reflecting the fact that the two subassociations partition the Venturan flora into substantially non-overlapping subsets of species. This segregation of associates between the two Salvia dominance types strongly suggests biotic influence of the dominants on subordinate species, perhaps mediated by allelopathy. This biotic interaction, leading to relatively strong floristic subassociations segregating independently in the landscape, would provide an example of the holistic community structure referred to by Clements and his followers, embedded within a larger pattern of continuity in species distributions.Nomenclature follows Munz & Keck (1959), Munz (1974) and Wiggins (1980).I am grateful to the following for research assistance: S. Coon, E. Hobbs, S. Lavinger, J. F. O'Leary, K. R. Preston, B. Rich and A. Troeger. I also thank the numerous public and private land owners who permitted access to the study sites. This research is based upon work supported by the National Science Foundation under grant DEB 76-81712.     

Relating species abundance distributions to species-area curves in two Mediterranean-type shrublands

Abstract. Based on both theoretical and empirical studies there is evidence that different species abundance distributions underlie different species‐area relationships. Here I show that Australian and Californian shrubland communities (at the scale from 1 to 1000 m²) exhibit different species‐area relationships and different species abundance patterns. The species‐area relationship in Australian heathlands best fits an exponential model and species abundance (based on both density and cover) follows a narrow log normal distribution. In contrast, the species‐area relationship in Californian shrublands is best fit with the power model and, although species abundance appears to fit a log normal distribution, the distribution is much broader than in Australian heathlands. I hypothesize that the primary driver of these differences is the abundance of small‐stature annual species in California and the lack of annuals in Australian heathlands. Species‐area is best fit by an exponential model in Australian heathlands because the bulk of the species are common and thus the species‐area curves initially rise rapidly between 1 and 100 m². Annuals in Californian shrublands generate very broad species abundance distributions with many uncommon or rare species. The power function is a better model in these communities because richness increases slowly from 1 to 100 m² but more rapidly between 100 and 1000 m² due to the abundance of rare or uncommon species that are more likely to be encountered at coarser spatial scales. The implications of this study are that both the exponential and power function models are legitimate representations of species‐area relationships in different plant communities. Also, structural differences in community organization, arising from different species abundance distributions, may lead to different species‐area curves, and this may be tied to patterns of life form distribution.     

Plot shape effects on plant species diversity measurements

Abstract. Question: Do rectangular sample plots record more plant species than square plots as suggested by both empirical and theoretical studies? Location: Grasslands, shrublands and forests in the Mediterranean‐climate region of California, USA. Methods: We compared three 0.1‐ha sampling designs that differed in the shape and dispersion of 1‐m² and 100‐m² nested subplots. We duplicated an earlier study that compared the Whittaker sample design, which had square clustered subplots, with the modified Whittaker design, which had dispersed rectangular subplots. To sort out effects of dispersion from shape we used a third design that overlaid square subplots on the modified Whittaker design. Also, using data from published studies we extracted species richness values for 400‐m² subplots that were either square or 1:4 rectangles partially overlaid on each other from desert scrub in high and low rainfall years, chaparral, sage scrub, oak savanna and coniferous forests with and without fire. Results: We found that earlier empirical reports of more than 30% greater richness with rectangles were due to the confusion of shape effects with spatial effects, coupled with the use of cumulative number of species as the metric for comparison. Average species richness was not significantly different between square and 1:4 rectangular sample plots at either 1‐ or 100‐m². Pairwise comparisons showed no significant difference between square and rectangular samples in all but one vegetation type, and that one exhibited significantly greater richness with squares. Our three intensive study sites appear to exhibit some level of self‐similarity at the scale of 400 m², but, contrary to theoretical expectations, we could not detect plot shape effects on species richness at this scale. Conclusions: At the 0.1‐ha scale or lower there is no evidence that plot shape has predictable effects on number of species recorded from sample plots. We hypothesize that for the mediterranean‐climate vegetation types studied here, the primary reason that 1:4 rectangles do not sample greater species richness than squares is because species turnover varies along complex environmental gradients that are both parallel and perpendicular to the long axis of rectangular plots. Reports in the literature of much greater species richness recorded for highly elongated rectangular strips than for squares of the same area are not likely to be fair comparisons because of the dramatically different periphery/area ratio, which includes a much greater proportion of species that are using both above and below‐ground niche space outside the sample area.     

Predicting the distribution of shrub species in southern California from climate and terrain-derived variables

Abstract. Generalized additive, generalized linear, and classification tree models were developed to predict the distribution of 20 species of chaparral and coastal sage shrubs within the southwest ecoregion of California. Mapped explanatory variables included bioclimatic attributes related to primary environmental regimes: averages of annual precipitation, minimum temperature of the coldest month, maximum temperature of the warmest month, and topographically-distributed potential solar insolation of the wettest quarter (winter) and of the growing season (spring). Also tested for significance were slope angle (related to soil depth) and the geographic coordinates of each observation. Models were parameterized and evaluated based on species presence/absence data from 906 plots surveyed on National Forest lands. Although all variables were significant in at least one of the species’ models, those models based only on the bioclimatic variables predicted species presence with 3–26% error. While error would undoubtedly be greater if the models were evaluated using independent data, results indicate that these models are useful for predictive mapping – for interpolating species distribution data within the ecoregion. All three methods produced models with similar accuracy for a given species; GAMs were useful for exploring the shape of the response functions, GLMs allowed those response functions to be parameterized and their significance tested, and classification trees, while some-times difficult to interpret, yielded the lowest prediction errors (lower by 3–5%).     

Observations on North-West European limestone grassland communities: Phytosociological and ecological notes on chalk grasslands of Southern England

Summary In 1970 chalk grasslands were studied in southern England, according to the principles of the French-Swiss School. The communities described belong to the associationCirsio-Brometum Shimwell 1971, emend., allianceMesobromion erecti (Br. Bl. & Moor 1938) Oberd. 1957, orderBrometalia W. Koch (1926 n.n.) Br.-Bl. 1936, of the classFestuco-Brometea Br.-Bl. 1936, emend. Tüx. 1961. Within theCirsio-Brometum three new sub-associations are distinguished: 1) subass. nov.anthoxanthetosum odorati; 2) subass. nov.Cornetosum sanguineae; and 3) subass. nov.campanuletosum rotundifoliae.Special attention is paid to the relationship that exists between the management of these communities and the seral stages that lead to the development of serub following the decreasing influence of grazing. Ulex europaeus scrub occurring on calcareous soil was also studied. The cryptogams of the chalk grasslands, mainly Musci, were investigated in detail.Plant nomenclature follows Heukels & van Ooststroom (1970) or for taxa not mentioned by these, Clapham et al. (1968) for phanerograms and Margadant (1959) and van der Wijk et al. (1969) for bryophytes.Contribution to the Symposium on Plant species and Plant communities, held at Nijmegen, 11 12 November 1976, on the occasion of the 60th birthday of Professor Victor Westhoff.