Perennial nitrophilous vegetation of the northern Iberian Peninsula

Abstract. The syntaxonomy of the perennial nitrophilous vegetation assigned to the Artemisietea vulgaris of the northern part of the Iberian Peninsula (Basque Country and surrounding areas) was revised. The study area is of biogeographic importance due to its transitional character - here, the ruderal vegetation of the Mediterranean and that of temperate Europe meet. Numerical ordination of the communities was performed in order to reveal systematic relations between the syntaxa. Two subclasses, Artemisienea vulgaris and Onopordenea, encompassing five orders, Convolvuletalia, Galio-Alliarietalia, Artemisietalia, Onopordetalia, Carthametalia lanati, and seven alliances with 12 associations and two rankless communities were distinguished.     

Morphological and molecular data support recognition of a new rupicolous species of Pinguicula (Lentibulariaceae) from the Iberian Peninsula

A new rupicolous species, Pinguicula saetabensis, belonging to P. sect. Pinguicula is described from calcareous cliffs of central-southern Valencia province, in the eastern Iberian Peninsula. It has previously been confused with P. mundi and P. vallisneriifolia, two close allies endemic to southern Spain which share some morphological traits and a similar habitat. However, some peculiarities allow recognition of those Valencian plants at the specific rank. Data on morphology, ecology, biogeography and conservation are reported for the new species, and its taxonomic affinities are discussed on the basis of phylogenetic analyses of the internal transcribed spacer region of nuclear ribosomal DNA. Furthermore, the presence of one population of P. vallisneriifolia is confirmed in that province, also based on morphological and molecular data.     

Spatial variation in a grassland on soil rich in heavy metals

Abstract. The vegetation in a grassland area in the prefecture of Kilkis (N. Greece), known for its surface Cu-mineralization, was studied. 43 quadrats were established along a transect through an area where the vegetation formed patches of different size. Cover-abundance and frequency estimates for all species were made. Normal Association Analysis revealed five quadrat groups characterized by Trifolium scabrum, Linaria pelisseriana, Anthoxanthum ovatum, Gypsophila muralis and Minuartia hirsuta ssp. falcata. 43 soil samples were analysed for pH, organic C, CaCO₃, total content of Fe, Zn, Pb, Cu, Mn, Ca, Mg, K, Na and soil texture. From a discriminant analysis performed on soil data five soil groups resulted, which are highly related to the five vegetation groups. Thymus sibthorpii, Minuartia hirsuta ssp. falcata and Rumex acetosella are the most important taxa with respect to physiognomy and patchi-ness of the vegetation. The number of species in each group of quadrats is affected by the relative favourableness or severity of the soil conditions. Metal contents, mainly Zn, Cu, Mg and Na, and soil texture are considered to be among the main factors controlling the structure and physiognomy of the vegetation.     

Influence of management on the species composition of epiphytic cryptogams in Danish Fagus forests

Abstract. The purpose of this study is to evaluate and to quantify the influence of management on the epiphytic species composition in Danish Fagus sylvatica forests. Trunks were chosen at random within selected blocks in both managed and unmanaged Fagus stands.117 trunks in 43 sample plots in three forests were investigated. Cover was estimated for 72 lichens, 29 bryophytes and two fungi on vertical trunks 0.3 ‐2 m above ground. 18 explanatory variables were recorded and arranged in three groups: eight management‐related variables, six micro‐climatic‐ and soil variables, and four geographical‐ and geological variables. Canonical Correspondence Analysis was used to test and to quantify the explaining power of the three groups of variables applying the variation partitioning approach. Detrended Correspondence Analysis was used to evaluate whether important gradients were ignored. It was concluded that the traditional ‘shelterwood uniform system’ used in Fagus forests has a marked influence on the epiphytic species composition. Thus, management‐related variables explained more than one third of the total variation explained, which equals 10.9% of the total variation. Microclimatic and soil variables explained 9.0%, and geographical and geological variables explained 6.0%. The following three recommendations are suggested to reduce the significant influence of management on the epiphytic species composition. First, it should be attempted to ensure a continuous occurrence of some big trees per ha to be left for natural death and decomposition. Second, regular thinning is necessary, especially in the first part of rotation. Third, a multi‐layered forest, where wind and desiccation effects are minimized, is recommended.     

Assessing the risk of freshwater fish introductions into the Iberian Peninsula

1. Preventing the introduction of species likely to become invaders is the best management option to deal with biological invasions. A data set consisting of native, introduced and species not currently present in Iberian Peninsula (n = 167 species) was used to identify freshwater fish species that are likely to be introduced and become successful invaders in the near future. 2. Principal component analysis (PCA) of species traits was used to determine species likely to be introduced, assuming that the traits of species introduced in the future will resemble those of previously introduced species. The likelihood of introduction was calculated as the proportion of neighbour species (in the space defined by the PCA) that have been introduced to the Iberian Peninsula and, together with metrics related to different stages of invasion, was used to construct a region‐specific risk index (Iberian risk index). 3. Introduced species had higher index values compared with native species or species currently absent from the region. The Iberian risk index was positively related to the results of an independent risk analysis for freshwater fish as well as to the geographical spread of species previously introduced to the Iberian Peninsula. 4. Iberian risk index values were used to establish a cut‐off value for estimating the probability of a successful invasion. This threshold value was used to construct a list of 20 species to be included in a ‘watch list’ to prevent freshwater fish invasions in the Iberian Peninsula.     

Establishment of Fagus sylvatica and Fraxinus excelsior in an old-growth beech forest

Abstract. Distribution of tree seedlings, forest architecture, light conditions, ground vegetation and humus conditions were studied in a 45 m × 100 m area including multiple gaps in an old-growth beech forest. Gaps were created after some beech trees had been felled in severe storms in February 1990. A group of adult ash trees is found near the study site. The data were analyzed by Correspondence Analysis. Young seedlings (< 4 yr), of both Fraxinus (a sun species) and Fagus (a shade species), were most abundant under the crown of beech trees in semi-shade conditions, and where beech litter did not accumulate. Differences in the dissemination of Fraxinus and Fagus explained differences in the establishment of the two species. In contrast, older seedlings of beech established before the storms were more numerous in the gaps, suggesting a change in the ecological requirements of beech seedlings in the course of time.     

Genetic diversity analysis and conservation implications for the Iberian threatened populations of the irano-turanian relict Krascheninnikovia ceratoides (Chenopodiaceae)

Krascheninnikovia ceratoides (L.) Gueldenst. (Chenopodiaceae) is an irano‐turanian steppe plant with a striking geographical vicariance between the western Mediterranean and the central Asian regions. Its westernmost populations are located in the north‐east part of the Iberian Peninsula, in the narrow middle Ebro and Alfambra valleys, where they are threatened and have been catalogued as ‘vulnerable’. Genetic diversity and structure was studied in 150 individuals from five populations of these two valleys using inter‐simple sequence repeats (ISSR). A total of 121 informative bands allowed the identification of 150 genotypes. Nei's genetic diversity found in all natural populations (h = 0.448) was higher than that of other geographical restricted species. Analysis of molecular variance and spatial correlations analyses showed a strong genetic differentiation among populations (35.88%) and among valleys (22.33%). The unweighted pair‐group method of arithmetical averages and Neighbour‐joining clusterings further demonstrated a substructure of two population cores within the Ebro valley. Our genetic data suggest that the high levels of genetic diversity and the strong genetic structure found among the northern Iberian populations and ranges of K. ceratoides might result from the summed effects of its tetraploidy, its outbreeding nature, and its success in colonizing newly altered areas. However, the ISSR data might also reflect the existence of an ancient wider distribution range of the plant in the Iberian Peninsula coupled with a more recent history of habitat fragmentation. Different management conservation guidelines are recommended for this plant after our genetic analysis: a microreserve of approximately 2 km² is proposed for the highly threatened population of Osera (Ebro valley), whereas seed collection strategies and occasional population reinforcements are suggested for the remaining populations. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 92 , 419–429.     

Aquatic vegetation and hydrology of a braided river floodplain

The response of aquatic vegetation to the complex hydrology of a river floodplain was investigated on a stretch of former braided channels of the Rhône River near Lyon, France, which is fed by three different water inputs: underground water, surface flow from a small tributary and river overflow during spates. Water temperature, pH, conductivity, alkalinity, sulfate, nitrate, and phosphate content were measured monthly at seven sampling stations. Principal Component Analysis (PCA) of these data revealed two gradients: a gradient of spatial heterogeneity and a gradient of temporal fluctuations. A PCA of samples based on substrate grain size, flow velocity, channel width and depth, height and shade of bank vegetation suggested the division of the braided system into four zones. Along these former channels, the vegetation varies from communities with Potamogeton coloratus of nutrient-poor water, to meso-eutraphent communities with Callitriche platycarpa, Elodea canadensis, and Ranunculus trichophyllus. Correspondence Analysis (CA) of the floristic data revealed four zones. Regarding the habitat parameters investigated, one zone is assumed to be linked to the instability and the nutrient inputs of a former channel (running water from the tributary and flood disturbances), while another is linked to the stability and nutrient-poor conditions of the other former channel (supplied by underground water). The other two zones are considered as ecotones, each of them occurring in a former channel, upstream of the confluences. The location as well as the floristic and functional differences between these two ecotones are discussed.     

Shrub vegetation on tropical granitic inselbergs in French Guiana

Abstract. In French Guiana, inselbergs in the form of granite outcrops rise abruptly from the surrounding rain forest. They constitute isolated islands of a special type of vegetation restricted to this peculiar substrate. Shrub granitic vegetation, organized in thickets on open exposed rocks of inselbergs, are described using the Braun‐Blanquet method combined with Correspondence Analysis. This phytosociological study revealed only one particular shrub community on each inselberg, including predominantly evergreen and sclerophyllous shrubs, especially microphanerophytes, belonging to the Clusiaceae, Myrtaceae and Bombacaceae. These outcrop communities exhibit species endemic to the Guianas region and also species rare in French Guiana. Affinities with flora of other inselbergs and vegetation types in South America are examined and discussed. Reasons for observed floristic and structural changes in each community are also discussed.     

Effects of disturbance caused by traditional Spanish rural land use on the regeneration of Cytisus multiflorus

Abstract. Cytisus multiflorus is a leguminous matorral shrub native to the NW Iberian Peninsula, where it is one of the most important species in the extension of matorral at the expense of set-aside agricultural land. Dehesas have traditionally been used for extensive livestock raising and matorral was periodically burnt, cut or pulled out. The two latter practices are now out of use. However, burning is more frequent than in the past. The effects of fire, cutting, and pulling out of C. multiflorus on its regeneration was studied in order to ascertain whether the presently increasing distribution of the species relates to fire-stimulated regeneration or to the reduction of other traditional practices. Three years after treatment two sets of parameters were determined: 1. Plant origin: seedlings and different ramet types (ramets=resprout clumps), density, weight, and biomass as well as the percentage of resprouting. 2. Seed persistence at various soil depths. The possible mechanisms of breaking dormancy and plant emergence in different years after fire were studied in other experiments. The results suggest that the regeneration mechanism in C. multiflorus is stimulated by fire, but it is not an exclusive relationship. Stimulation occurs also after other disturbances leading to the total elimination of aerial biomass. The present expansion of the species does not appear to result from the abandonment of some traditional practices, such as cutting or pulling out, but from frequent fires (resulting in aerial-biomass elimination).     

Niche characteristics of Danish woody species as derived from coenoclines

Abstract. A main floristic coenocline is extracted from a Danish forest vegetation data set with Detrended Correspondence Analysis. Huisman‐Olff‐Fresco (HOF) modelling of 262 species occurrences relative to site scores along the coenocline is compared with Gaussian and Generalized Linear Modelling. HOF models allow for responses of variously skewed forms, which may result from interspecific interactions. Most species have symmetric (60%) or skewed (29%), and only few species have flat or monotone response curves. This provides evidence for the generality of unimodal plant species response curves, provided the gradient in question comprises a sufficient amount of compositional turnover. The Gaussian niche width is correlated with density of optima and probability of occurrence of the species. Detailed analysis of HOF estimates for 28 woody species indicates that early succession species such as Pinus sylvestris and Populus tremula have wider niches than middle to late successional canopy species such as Alnus glutinosa, Carpinus betulus, Fagus sylvatica, Fraxinus excelsior, Quercus robur and Tilia cordata. A marked overlap in niche range along the coenocline is interpreted in terms of species competition. Rubus idaeus intrudes into the niches of all other shrub species. Several canopy tree species, such as Fagus sylvatica, Fraxinus excelsior and Quercus robur overlap more than 60% of the ranges of all other species, while their internal overlaps are fairly equal, presenting evidence for their likely co‐existence given more natural forest conditions. The combined analysis of coenoclines with HOF modelling of species behaviour is the most robust among the compared methods for estimating model parameters and niche characteristics of plant species. It is recommended for future ecological studies.     

Changes in landscape structure in the northwestern Alps over the last 7000 years: lessons from soil charcoal

Abstract. Current land‐use a bandonment and the current rise in temperature in the Alps both suggest that tree limits may change. When it is assumed that the climate of the early mid‐Holocene between 8000 and 5000 yr before present is analogous to that of the predicted climate of the late 21st century, palaeo‐ecological studies of the early Holocene may provide data for the prediction of the vegetation pattern in a century from now. It appears that mid‐Holocene charcoal assemblages can be used to reconstruct the spatial patterns of the vegetation before, or during, the practice of slash‐and‐burn. Correspondence analysis (CA) of charcoal assemblages shows that an important ecological gradient is determined by elevation. However CA also shows that charcoal assemblages in profiles between 1700 and 2100 m a.s.l. are roughly stratified: the more recent assemblages from the topmost centimetres of soil are intermediate between the lowermost assemblages and assemblages from higher elevations. This suggests that the woody communities at the highest elevation were located at lower elevations at a later date. The taxonomic diversity of the soil charcoal assemblages has been compared to that of present‐day phytosociological relevés after transformation to charcoal‐equivalent data. This comparison revealed that the vegetation pattern along the altitudinal gradient in the mid‐Holocene was different from that at present. The assemblages indicate that some communities disappeared, that Picea is a late‐Holocene invading species, and that there is no strict modern analogue for the vegetation structure prior to that of 3000 yr ago. The past structure of the woody vegetation was also different from that of today. Although past vegetation is not a good analogue for predicting future vegetation patterns, it still has potential as an indicator for the potential presence of tree species where there is none today. If we assume a temperature rise, and take into account current trends of landscape use abandonment, then we can expect strong vegetation dynamics at the upper tree line in the future: Abies alba may expand to occupy elevations of ca. 1800–2000 m in mixed communities with Picea abies, Pinus sylvestris and hardwood species, and Pinus cembra may expand up to 2500–2700 m a.s.l.     

Phylogeny of western Mediterranean Leptodirini, with an emphasis on genital characters (Coleoptera: Leiodidae: Cholevinae)

Abstract The tribe Leptodirini (Leiodidae: Cholevinae) is one of the largest radiations of Coleoptera in the subterranean environment. Although subjected to systematic and evolutionary studies, the phylogeny remains poorly understood. We assessed the phylogeny of the western Mediterranean lineages (Iberian Peninsula, Pyrenees and Sardinia) based on a cladistic analysis of fourteen characters of external morphology and twenty characters of the male and female genitalia, studied in 182 species belonging to thirty‐nine genera. We tested the monophyly of the traditional two main divisions of the group (infraflagellates and supraflagellates), as well as that of some ‘phyletic series’. The final matrix contained fifty‐eight terminal taxa, twenty‐four of which had different character state combinations. The strict consensus of the sixty most parsimonious trees recovered a monophyletic Leptodirini, but not their separation into infraflagellates and supraflagellates. The supraflagellates formed a paraphyletic group with respect to the infraflagellates (corresponding to our sampled ‘Speonomus’ series), with Notidocharis sister to all other included Leptodirini, and Speonomidius sister to Leptodirini excluding Notidocharis. The series ‘Spelaeochlamys’, including the Sardinian genera but excluding Pseudochlamys, was recovered as monophyletic with weak support. The ‘Quaestus’ series formed a polytomy with Pseudochlamys plus the ‘Speonomus’ series (including Bathysciola), which was recovered as monophyletic with strong support. Speonomus, Bathysciola, Quaestus and Troglophyes were para‐ or polyphyletic. Our results suggested the respective monophyletic origin of the Leptodirini from the Pyrenees (Pseudochlamys plus the ‘Speonomus’ series) and the Mediterranean coast plus Sardinia (series ‘Spelaeochlamys’). On the contrary, the Leptodirini of the Atlantic north coast of the Iberian Peninsula (series ‘Quaestus’ and ‘Speonomidius’) were not monophyletic.     

Environmental control on the biogeographical distribution of Desmanella (Soricomorpha,Mammalia) in the Miocene of the Iberian Peninsula

Abstract: This paper reports the first record of Desmanella (Mammalia, Soricomorpha) from the Granada Basin in southern Iberian Peninsula, which represents its south‐westernmost occurrence in all Eurasia. It is a controversial taxon whose systematic assignment has been discussed for a long time. This genus belongs to the family Talpidae, a group of insectivores that include extant moles, shrew moles and desmans. Desmanella was very abundant in the late Miocene and early Pliocene of Eurasia, including several basins of northern Iberian Peninsula, but it has not been found until now in southern Iberia. Previous studies have shown that the faunal record and the climatic conditions during the Neogene in the Granada Basin were different from other areas of the Iberian Peninsula. The particular distribution of Desmanella supports the hypothesis that specific climatic features (wetter than neighbouring areas) prevailed in the Granada Basin during the late Turolian (late Miocene).     

The plant communities of theAsplenietea trichomanis in the SW Iberian Peninsula

The rupicolous vegetation of theAsplenietea trichomanis in the south-western Iberian Peninsula is analysed using the methods of the Zürich-Montpellier School. Phytosociological tables, and biogeographical, ecological and floristic data are given for a number of syntaxa. A phytosociological synopsis of the studied vegetation was made for the entire region. Four new syntaxa are described: theCoincyo logirostri-Dianthetum lusitani, Jasiono marianae-Dianthetum lusitani jasionetosum tomentosae, Digitali thapsi-Dianthetum lusitani conopodietosum ramosi andSedo hirsuti-Polypodietum cambrici polypodietosum interjecti.     

Tropical forest structure and diversity in relation to altitude and disturbance in a Biosphere Reserve in central India

Questions: Is species diversity affected in protected areas where human activities are permitted or tolerated? On plots of a fixed size, does stem density alone predict number of species? Are differences in density related to disturbance and altitude? Location: Achanakmar‐Amarkantak Biosphere Reserve, central India. Methods: 42 sites, each with three replicate 10‐m radius plots, were examined. All trees (≥ 30 cm GBH) in each plot were measured for girth at breast height. α‐diversity, species richness and evenness were calculated for each site. The sites were ordinated by Nonmetric Multidimensional Scaling (NMS) using relative importance values of component species. Correspondence Analysis was used to broadly delineate communities. Anthropogenic disturbances were recorded in terms of percentage of trees lopped, scale of lopping, number of domestic livestock dung piles and foot trails (both livestock and people) for each plot. Results: The NMS analysis exhibited a near linear arrangement of sites with no evidence of discrete vegetation zones. NMS axes were significantly related to altitude and disturbance scores. With increasing elevation, basal area increased but number of species, α‐diversity and its components declined monotonically. The number of species and indices of species diversity were positively associated with tree lopping and also with total disturbance. Number of species was controlled by stem density only in plots not dominated by Shorea robusta. Conclusions: Recent levels of human disturbance are associated with higher species diversity in this biosphere reserve. There is some evidence that stands at all altitudes follow the same successional pattern to dominance by Shorea, a successional pattern that also results in decreased diversity without disturbance.     

Long‐term dynamics of the hemiparasite Rhinanthus angustifolius and its relationship with vegetation structure

Questions: 1. How are the long‐term dynamics of the root hemiparasite Rhinanthus angustifolius related to vegetation structure, grassland management and climate? 2. Does R. angustifolius have a long‐term impact on standing crop and community composition? Location : A formerly fertilized grassland, part of a larger brook valley system in the nature reserve ‘Drentsche Aa’, near Groningen, The Netherlands. Methods : Vegetation development has been monitored since 1973 in 54 permanent plots in nine management regimes without fertilizer application. Results : 1. The hemiparasite established when standing crop was less than 600 g.m^?2 and performed best under annual haymaking using machinery. Since its appearance, the population fluctuated stochastically, with two peaks. Coinciding collapses in six adjacent grasslands and comparison with an integrated climatic index suggest that the population collapses are induced by spring drought. 2. We did not find a relationship between total standing crop and R. angustifolius cover. Cover of grasses was negatively related to the abundance of the hemiparasite in the same and the previous year. Forb cover tended to increase with the parasite. Conclusions : R. angustifolius shows stochastic population fluctuations, mainly determined by spring drought, to which this species is probably highly vulnerable because of its parasitic and annual life style without a persistent seed bank. The hemiparasite also shows long‐term relationships with grass cover (negative) and forb cover (positive), but it seems to have no lasting impact on standing crop.     

Relationships between vegetation zonation and altitude in a salt-marsh system in northwest Spain

Abstract. Vegetation zonation in salt marshes has traditionally been attributed largely to altitudinal differences, since altitude determines the temporal pattern of tidal flooding and is thus closely related to proximate determinants of the distribution of species and plant communities. We investigated the distribution of vascular plants and plant communities along a series of altitudinal transects in two salt marshes in the northwestern Iberian Peninsula. Our results indicate that altitudinal range varies significantly both among species and among communities, and confirm that salt-marsh vegetation characteristics (species cover and composition) can be predicted on the basis of altitude, particularly at the lower levels of the profile.     

Above‐ground biomass distribution among species during early old‐field succession

Abstract. The interdependence of species richness and plant biomass has widely been accepted as a general biodiversity rule. However, there is no information about how relationships are established during colonization and how total biomass is distributed among plants. The main objective of this study was to determine the role of several factors which we have hypothetized as affecting biomass distribution among species in an early old‐field community. To test this hypothesis, we conducted an experiment in a deeply ploughed Agropyro‐Rumicion crispi community in the Basque Country (Spain). Light, water and soil nitrogen content were factorially manipulated resulting in eight experimental treatments. We also examined several site features, which could potentially influence final biomass distribution: seed bank composition, soil physico‐chemical heterogeneity and cover and density of the dominant plant species in the community. A partition hypothesis testing approach using Redundancy Analysis was conducted to determine the fraction of biomass distribution variability assigned to each treatment and site feature. The most important species, in terms of biomass, were Agropyron repens, Sinapis arvensis, Arrhenatherum bulbosum and Picris echioides. As a general conclusion, biomass distribution among species during early secondary succession primarily depends on light availability and nitrogen supply. Several soil variables, such as magnesium, calcium and clay contents, also explain a relevant fraction of the biomass distribution among plant species. On the contrary, we found no effect of seed bank composition on biomass distribution. Finally, the total species number and cover of dominant species such as Sinapis arvensis, may determine final biomass distribution.