The phytogeography of European and Mediterranean heath species (Ericoideae, Ericaceae): a quantitative analysis

The geographic ranges of heath species in Europe and the Mediterranean and their relationships with environmental (climatic and ecogeographic) variables and biological features are analysed by means of multivariate methods. In particular, twinspan classifications into floristic elements and floristic regions, DCA floristic ordinations, CCA environment-constrained ordinations and CCA biology-constrained ordinations are carried out. Results of the analyses show a correspondence with conventional regionalization analyses based on broader criteria, and less correspondence with numerical analyses of other taxonomic groups at a similar scale. This lack of fit depends on the particular history and ecology of the taxonomic groups under study. A number of climatic (temperature and water stress) and geographic (coast length) variables are associated with different types of heaths according to their geographical ranges (continental, Mediterranean, Atlantic). Biological features of heaths account for a small part of the variation in range, but the association of temperate heaths with a preference for acid soils, of Mediterranean heaths with pubescence, and of Atlantic heaths with plant height is of interest. Heath species richness throughout Europe and the Mediterranean is analysed by multiple regression analyses and, apart from a strong influence of area size, a significant effect of water conditions, temperature and proximity to sea is detected. The area with the highest heath species richness is Western Mediterranean. Additional classification, ordination and multiple regression analyses of heaths in the Iberian Peninsula (Spain and Portugal, the areas with the highest heath diversity) revealed similar patterns to those found in Europe and the Mediterranean. The effect of heterogeneity of the studied units at this latter scale is removed in the Iberian analysis because of the relative homogeneity of the units considered at this scale.     

The vegetation of seacliffs and headlands in New South Wales,Australia

A phytosociological survey of seacliff and headland vegetation on the central and south coast of New South Wales was carried out. Analysis of the quadrat data led to the recognition of 15 communities; the occurrence of a number of other vegetation types is briefly discussed. The vegetation types fall into three major groupings: grasslands, scrub and heathland, although the coincidence between structural and floristic boundaries is poor. Ordination of the communities indicates that the major factors varying between communities are exposure and soil fertility, the grasslands and scrubs occurring on more fertile soils than the heathlands. Amongst the woody communities it is suggested that the two major groups recognized fall into different phytosociological alliances, although lack of studies of Australian vegetation limits the applicability of a conventional hierarchical classification. The true heathlands correspond closely with the alliance Leptospermion, previously recognized in Victoria. The scrub communities are assigned to a new alliance, the Westringio-Banksion integrifoliae.     

Analyse pollinique des dépôts lacustresdu Pléistocène inférieur de Banyoles (Bañolas,site de la Bòbila Ordis - Espagne): un élément nouveau dans la reconstitution de l'histoire paléoclimatique des régions méditerranéennes d'Europe occidentale

Clays from Bòbila Ordis near Banyoles have yielded a rich pollen flora. Three pollen zones are recognizable: the lower and the middle parts are dominated by components of a deciduous forest (with numerous exotic taxa); in the upper part, Pinus pollen and NAP (especially Compositae) are in a majority with an increase of mediterranean taxa towards the top. This floristic evolution may be connected with a climatic change characterized by a decrease in humidity which is itself linked to a world cooling. We suggest a possible relation with the pollen flora published by H. Elhai. The pollen diagram of Bòbila Ordis is the second so far established from Lower Pleistocene deposits in the Mediterranean countries of Western Europe; chronologically it follows the Bernasso (Languedoc) pollen diagram and confirms the latter results.     

Disentangling environmental correlates of vascular plant biodiversity in a Mediterranean hotspot

We determined the environmental correlates of vascular plant biodiversity in the Baetic‐Rifan region, a plant biodiversity hotspot in the western Mediterranean. A catalog of the whole flora of Andalusia and northern Morocco, the region that includes most of the Baetic‐Rifan complex, was compiled using recent comprehensive floristic catalogs. Hierarchical cluster analysis (HCA) and detrended correspondence analysis (DCA) of the different ecoregions of Andalusia and northern Morocco were conducted to determine their floristic affinities. Diversity patterns were studied further by focusing on regional endemic taxa. Endemic and nonendemic alpha diversities were regressed to several environmental variables. Finally, semi‐partial regressions on distance matrices were conducted to extract the respective contributions of climatic, altitudinal, lithological, and geographical distance matrices to beta diversity in endemic and nonendemic taxa. We found that West Rifan plant assemblages had more similarities with Andalusian ecoregions than with other nearby northern Morocco ecoregions. The endemic alpha diversity was explained relatively well by the environmental variables related to summer drought and extreme temperature values. Of all the variables, geographical distance contributed by far the most to spatial turnover in species diversity in the Baetic‐Rifan hotspot. In the Baetic range, climate was the most significant driver of nonendemic species beta diversity, while lithology and climate were the main drivers of endemic beta diversity. Despite the fact that Andalusia and northern Morocco are presently separated by the Atlantic Ocean and the Mediterranean Sea, the Baetic and Rifan mountain ranges have many floristic similarities – especially in their western ranges – due to past migration of species across the Strait of Gibraltar. Climatic variables could be shaping the spatial distribution of endemic species richness throughout the Baetic‐Rifan hotspot. Determinants of spatial turnover in biodiversity in the Baetic‐Rifan hotspot vary in importance between endemic and nonendemic species.     

Steppes, savannahs, forests and phytodiversity reservoirs during the Pleistocene in the Iberian Peninsula

A palaeobotanical analysis of the Pleistocene floras and vegetation in the Iberian Peninsula shows the existence of patched landscapes with Pinus woodlands, deciduous and mixed forests, parklands (savannah-like), shrublands, steppes and grasslands. Extinctions of Arctotertiary woody taxa are recorded during the Early and Middle Pleistocene, but glacial refugia facilitated the survival of a number of temperate, Mediterranean and Ibero-North African woody angiosperms. The responses of Iberian vegetation to climatic changes during the Pleistocene have been spatially and temporarily complex, including rapid changes of vegetation in parallel to orbital and suborbital variability, and situations of multi-centennial resilience or accommodation to climatic changes. Regional characteristics emerged as soon as for the Middle Pleistocene, if not earlier: Ericaceae in the Atlantic coast indicating wetter climate, thermo-mediterranean elements in the south as currently, and broad-leaf trees in the northeastern. Overall, steppe landscapes and open Pinus woodlands prevailed over many continental regions during the cold spells of the Late Pleistocene. The maintenance of a high phytodiversity during the glacials was linked to several refuge zones in the coastal shelves of the Mediterranean and intramountainous valleys. Northern Iberia, especially on coastal areas, was also patched with populations of tree species, and this is not only documented by palaeobotanical data (pollen and charcoal) but also postulated by phylogeographical models.     

Ecological study of pine forest clearings along the French Atlantic sand dunes: Perspectives of restoration

The 18th century afforestation campaign carried out in western France to fix sand dunes has left only a small surface of non-forested dunes. As dune plantations have only a small conservation value, it would be of great interest to restore grey dunes. But, for the moment, there is no proof that pine felling would lead to grey dunes. So, a study has been carried out on pine forest clearings along the French Atlantic shoreline. Floral and ecological data were carried out and analysed with two statistical tools, CA (canonical analysis) and CCA (canonical correspondence analysis). The clearings appear very different from one bank of the Gironde river to the other. South of the river, the clearing vegetation develops on acid soils and possess some thermophilous species. North of the river, the clearings are composed mostly of species growing best on calcareous soils. Apart from geographical variations, clearings showed two patterns of response: development of closed thickets and tall heathlands; or an open vegetation similar to growing heathlands and grasslands. The cover, the height, the floral richness per strata as well as pedological characteristics are the parameters which explain the best floristic composition of pine clearings. The vigour of pine forests and the degree of exposure to coastal influences determine whether close or open vegetation develops. Where there is an open vegetation, restoration of grey dunes may be possible. In the other case, alternative solutions, such as the restoration of dune woodlands or dune heath, may provide best conservation values.     

Deterministic Plio-Pleistocene extinctions in the European cool-temperate tree flora

Extinctions have been important in the shaping of modern phytogeographic patterns. A classic example is the heavy Plio‐Pleistocene losses that have caused Europe to have a depauperate temperate tree flora compared to eastern North America and eastern Asia. To investigate the mechanisms involved in this extinction event, I test the hypothesis that the present European status (extinct, relictual, or widespread) of cool‐temperate tree genera found in Pliocene Europe is predictable from their modern climatic requirements. As a prerequisite for this analysis, I test for genus‐level conservatism in climatic requirements by comparing congeneric values across Europe, eastern Asia and North America, and find strong evidence hereof. I find a high degree of ecological determinism in the fate of European Pliocene tree genera, still widespread taxa being more tolerant of cold growing season and winter temperatures than extinct and relictual taxa, and relictual taxa being more drought tolerant than extinct taxa.     

Phenological and biogeographical aspects of coastal dune plant communities in southern Brazil

The coastline of southern Brazil is characterized by vast sand dunes. From March 1982 to April 1984 the dune vegetation was sampled along a transect between the Atlantic shore and an inland freshwater marsh, and phenological cycles of plant species were observed at monthly intervals.The occurrence of 23 annual species in the local dunes and peak germination, growth and flowering of 21 perennial species during spring, summer and fall, suggest that unfavourable temperatures and inundation of slacks during winter are seasonally limiting factors for this flora.Differences in the floristic composition between this region and its southern and northern extremes in the southwestern Atlantic together with a marked seasonality of the total species number and of germination and flowering periods, suggest that this area corresponds to a warm temperate biogeographic transition zone between northern tropical and southern cold temperate regions.     

Biogeographic analysis of Jurassic-Early Cretaceous wood assemblages from Gondwana

The terrestrial biogeography of Gondwana during Jurassic-Early Cretaceous times is poorly resolved, and the flora is usually considered to have been rather uniform. This is surprising given the size of Gondwana, which extended from the equator to the South Pole. Documenting Gondwanan terrestrial floristic provincialism in the Jurassic-Early Cretaceous times is important because it provides a historical biogeographic context in which to understand the tremendous evolutionary radiations that occurred during the mid-Cretaceous. In this paper, the distribution of Jurassic-Early Cretaceous fossil wood is analysed at generic level across the entire supercontinent. Specifically, wood assemblages are analyzed in terms of five climatic zones (summer wet, desert, winter wet, warm temperate, cool temperate) established on the basis of independent data. Results demonstrate that araucarian-like conifer wood was a dominant, cosmopolitan element, whereas other taxa showed a greater degree of provincialism. Indeed, several narrowly endemic morphogenera are recognizable from the data. Finally, comparisons with Laurasian wood assemblages indicate strong parallelism between the vegetation of both hemispheres.     

Floristic changes in the Iberian Peninsula and Balearic Islands (south-west Europe) during the Cenozoic

Aim  The aim of this work was to identify the main changes in the flora and vegetation of the Iberian Peninsula over the Cenozoic Era, to record the disappearance of taxa associated with these changes and to determine the influence of climate and human activity on these events.
Location  The Iberian Peninsula and Balearic Islands.
Methods  A critical review was made of the palaeobotanical literature with the aims of detecting patterns of floristic change and extracting information on the disappearance of different taxa over the Cenozoic. These data are viewed alongside the most recent data for the climate of this period. A critical analysis is made of the role of Palaeotropical and Arctotertiary taxa in the forest communities of the Iberian Peninsula throughout the Cenozoic.
Results  Although the Eocene–Oligocene transition was a time when many taxa disappeared, the most outstanding events occurred between the end of the Oligocene and throughout the Miocene. Substantial floristic changes took place over this period, including the disappearance of 177 Palaeotropical taxa. This was probably related to acute cooling and aridification; no evidence exists that the Messinian Salinity Crisis had any important effect in the Iberian Peninsula. The last great disappearance of Palaeotropical taxa (36 in total) ended in the Middle–Late Piacenzian; Arctotertiary taxa were most affected during the Pleistocene. The Lower–Middle Pleistocene transition, best represented by marine isotopic stages (MIS) 36–34 and 20–18 and characterized by a change in glacial cyclicity, was the time of the last notable disappearance of taxa.
Main conclusions  This work provides the first chronogram of extinctions for the Iberian flora, and records the disappearance of 277 taxa during the Cenozoic. A clear relationship was detected between the main climatic events and the latest appearances of the different taxa.     

European post-glacial forests: compositional changes in response to climatic change

Abstract. Multivariate analysis of an extensive palyno-logical database for Europe has enabled reconstruction of broad-scale vegetation history. Whereas many major features of present vegetation patterns were established early in the Holocene, floristic composition of the forests has changed continuously up to the present day. For example, although ‘mixed deciduous forests’ had reached approximately their present extent in northwest Europe by 8000 B.P., Tilia peaked in abundance in these forests during the middle post glacial, whereas Pinus was most abundant in these forests during the early post-glacial and Fagus increased in abundance only in recent millennia. Pollen-climate response surfaces for major pollen taxa show how their distribution and abundance patterns relate to contemporary climate. Past forest-compositional changes were responses to climatic changes, the nature of which can be inferred from pollen-climate response surfaces. Post-glacial climate changes have been different in magnitude and direction in different regions of Europe. For example, in recent millennia the vegetation changes indicate decreasing summer temperatures in northern Europe but increasing summer temperatures in the Mediterranean region. The way in which vegetation responded to past climatic changes gives insight into the likely response of vegetation to future climate changes induced by the ‘greenhouse effect’.     

Quaternary refugia are associated with higher speciation rates in mammalian faunas of the Western Palaearctic

Quaternary climate changes have contributed to shape the biogeographic distribution of extant species. The combination of climatic niche conservatism and glacial‐interglacial cycles forced many species to retract their range limits for surviving under the advance of Pleistocene ice‐sheets. Refugia offered geographical opportunities for species to retreat, persist and, later on, begin recolonization processes under favourable environmental conditions. Here we explore the hypothesis that refugia have been not only crucial for the survival of multiple species but also acted as speciation centres for Western Palaearctic mammals. We define ‘recurrent massive refugia’ as those geographical regions that have historically accumulated the highest levels of co‐occurring species for several Quaternary cycles. Our assemblage‐level analyses identify the existence of refugia within the Iberian Peninsula and the Atlantic French margin that were recurrently selected by most mammals. The topographic heterogeneity, climatic stability and microhabitat availability of these refugial areas may have offered suitable habitat conditions for multiple species during different climatic events over time. Using a Bayesian analysis of macroevolutionary mixtures we detected that the higher level of divergence and accumulative evolutionary changes in mammals of the Western Palaearctic are found in refugia. The continuous retractions and expansions of species’ ranges during the Pleistocene promoted temporal changes in the composition and richness of communities in this biogeographic region. The reorganization of ecological composition driven by cyclical climatic events may have favoured the emergence of biotic interactions and ecological responses conducive to novel selective pressures. Our findings suggest, first, that multiple climatic changes in the form of glacial‐interglacial transitions during the Quaternary have left a detectable imprint on the observed geographical patterns of species richness in mammalian faunas of the Western Palaearctic, and second, highlight the importance of refugia for the preservation of species (‘museums’) and as centers of speciation and endemism (‘cradles’) as well.     

A floristic analysis of the lowland dipterocarp forests of Borneo

Aim To (1) identify floristic regions in the lowland (below 500 m a.s.l.) tropical dipterocarp rain forest of Borneo based on tree genera, (2) determine the characteristic taxa of these regions, (3) study tree diversity patterns within Borneo, and (4) relate the floristic and diversity patterns to abiotic factors such as mean annual rainfall and geographical distance between plots. Location Lowland tropical dipterocarp rain forest of Borneo. Methods We used tree (diameter at breast height ≥ 9.8 cm) inventory data from 28 lowland dipterocarp rain forest locations throughout Borneo. From each location six samples of 640 individuals were drawn randomly. With these data we calculated a Sørensen and Steinhaus similarity matrix for the locations. These matrices were then used in an UPGMA clustering algorithm to determine the floristic relations between the locations (dendrogram). Principal coordinate analysis was used to ordinate the locations. Characteristic taxa for the identified floristic clusters were determined with the use of the INDVAL method of Dufrene & Legendre (1997) . Finally, Mantel analysis was applied to determine the influence of mean annual rainfall and geographical distance between plots on floristic composition. Results A total of 77 families and 363 genera were included in the analysis. On average a random sample of 640 trees from a lowland dipterocarp forest in Borneo contains 41.6 ± 3.8 families and 103.0 ± 12.7 genera. Diversity varied strongly on local scales. On a regional scale, diversity was found to be highest in south‐east Borneo and central Sarawak. The most common families were Dipterocarpaceae (21.9% of trees) and Euphorbiaceae (12.2% of trees). The most common genera were Shorea (12.3% of trees) and Syzygium (5.0% of trees). The 28 locations were clustered in geographically distinct floristic regions. This was related to the fact that floristic similarity depended strongly on the geographical distance between plots and similarity in mean annual rainfall. Conclusions We identified five main floristic regions within the lowland dipterocarp rain forests of Borneo, each of which had its own set of characteristic genera. Mean annual rainfall is an important factor in explaining differences in floristic composition between locations. The influence of geographical distance on floristic similarity between locations is probably related to the fact that abiotic factors change with distance between plots. Borneo's central mountain range generally forms an effective dispersal barrier for the lowland tree flora. Diversity patterns in Borneo are influenced by the mid‐domain effect, habitat size and the influence of past climatic changes (ice ages during the Pleistocene).     

Zonobiomes, zonoecotones and azonal vegetation along the Pacific coast of North America

In this study of the Pacific coast of North America, from Baja California to Alaska, we evaluated the hypothesis that the floristic composition of azonal vegetation determines areas and distribution limits similar to those of the corresponding zonobiomes (ZB), and does so in response to the same macroclimatic changes occurring on the continental scale. To this end, 686 vascular plants of the different habitats found in 279 sites along this coastal strip were recorded. Using an objective classification system (Average Linkage Clustering) and factorial analysis, floristic data acquired in fieldwork were classified into groups, which were in turn related to regional macroclimates. Our main finding was that the azonal coastal vegetation follows a distribution model that is closely linked to the corresponding macroclimate. The four ZB of the northern Pacific coast show a flora and azonal vegetation characteristic to each zonobiome; the latitudinal limits of the azonal vegetation practically coinciding with those already established for the zonal vegetation. The Boreal and Temperate ZB show high percentages of broadly distributed elements. The floristically richest zonobiome in terms of endemic taxa is the Mediterranean zonobiome, whereas the flora of Baja California is characterized by a high number of taxa related to Neotropical flora, especially to those showing links with South America. Data on the geographical distribution and habitats of the 247 most significant coastal species are also provided.     

The age of Calluna stands moderates post‐fire regeneration rate and trends in northern Calluna heathlands

Questions: Does stand age influence the direction and rate of post‐fire successional dynamics in coastal Calluna heaths and can old degraded heath vegetation be restored through reintroduction of fire? Location: Coastal heaths in the Tarva archipelago, central Norway. Methods: We investigated revegetation dynamics after experimental fires set in young (8 years since last fire) and old (>50 years since last fire) grazed heath stands. A repeated measures design was used, with floristic data recorded in permanent plots in the post‐fire successions (n=12) over a 7‐year period. The data were analysed using multivariate ordination techniques (PCA, RDA and PRC) and mixed effects models. Results: The age of Calluna stands strongly influenced post‐fire succession, different trends due to age explained 10.4% of variation in floristic data. Young heath showed faster succession towards pre‐fire community composition than old heath, and this could partially be explained by succession‐related factors: young heath had lower cover of mosses and lichens in the pre‐burned vegetation, and lower cover of litter early in succession. Young heath had a less pronounced overall community response to fire than old heath. Vegetative regeneration of C. vulgaris was absent in both old and young heath, but Calluna still re‐established as the dominant species within 5–7 years in both young and old stands. Regeneration dynamics were also affected by habitat conditions, different trends due to habitat explained 6% of variation. Conclusions: Our study demonstrates that old stands do develop characteristic heathland vegetation and structure after fire, and while potential invasives into the system such as trees and rhizomatous species are present, they do not impair Calluna regeneration or vegetation development towards the target heathland community composition and structure. Further, as our young stands are only in their second fire rotation after restoration, we suggest that characteristic dynamics of managed heathlands can re‐establish relatively rapidly, even in severely degenerated sites (>50 years since last fire). Site‐specific factors also need to be considered. We conclude that there is restoration potential in old heaths, despite slow dynamics in the first rotation.     

Distribution and diversity of Arctic-Alpine species in the Balkans

The distributions of 77 Arctic-Alpine species in the Balkans are mapped and the centers of their richness and diversity presented. Within the Dinaric Alps these are Mts Vranica, Durmitor, and Prokletije; in the Scardo-Pindhic mountains, ?arplanina–Rudoka–Korab form a continuous chain; in the Rhodope-Rila mountain system there are Mts Vito?a, Rila, and Pirin; while in the Balkan mountain system there are the West and Central part of Stara planina. A comparison of floristic richness and distribution of Arctic-Alpine flora in relation to altitude, geographical location, and geological substrate is made. Correlations between floristic richness and geographical distance of the Balkan mountains from the two main centers of Arctic-Alpine flora in Central Europe are also provided.     

Phytogeographical analysis of seed plant genera in China

BACKGROUND AND AIMS: A central goal of biogeography and ecology is to uncover and understand distributional patterns of organisms. China has long been a focus of attention because of its rich biota, especially with respect to plants. Using 290 floras from across China, this paper quantitatively characterizes the composition of floristic elements at multiple scales (i.e. national, provincial and local), and explores the extent to which climatic and geographical factors associated with each flora can jointly and independently explain the variation in floristic elements in local floras. METHODS: A study was made of 261 local floras, 28 province-level floras and one national-level flora across China. Genera of seed plants in each flora were assigned to 14 floristic elements according to their worldwide geographical distributions. The composition of floristic elements was related to climatic and geographical factors. KEY RESULTS AND CONCLUSIONS: Variations in percentages of cosmopolitan, tropical and temperate genera among local floras tend to be greater at higher latitudes than at lower latitudes. Latitude is strongly correlated with the proportions of 13 of the 14 floristic elements. Correlations of the proportions of floristic elements with longitude are much weaker than those with latitude. Climate represented by the first principal component of a principal component analysis was strongly correlated with the proportions of floristic elements in local floras (|r| = 0.75 +/- 0.18). Geographical coordinates independently explained about four times as much variation in floristic elements as did climate. Further research is necessary to examine the roles of water-energy dynamics, geology, soils, biotic interactions, and historical factors such as land connections between continents in the past and at present in creating observed floristic patterns.