Survival and long-term maintenance of tertiary trees in the Iberian Peninsula during the Pleistocene: first record of Aesculus L. (Hippocastanaceae) in Spain

The Italian and Balkan peninsulas have been places traditionally highlighted as Pleistocene glacial refuges. The Iberian Peninsula, however, has been a focus of controversy between geobotanists and palaeobotanists as a result of its exclusion from this category on different occasions. In the current paper, we synthesise geological, molecular, palaeobotanical and geobotanical data that show the importance of the Iberian Peninsula in the Western Mediterranean as a refugium area. The presence of Aesculus aff. hippocastanum L. at the Iberian site at Cal Guardiola (Tarrasa, Barcelona, NE Spain) in the Lower–Middle Pleistocene transition helps to consolidate the remarkable role of the Iberian Peninsula in the survival of tertiary species during the Pleistocene. The palaeodistribution of the genus in Europe highlights a model of area abandonment for a widely-distributed species in the Miocene and Pliocene, leading to a diminished and fragmentary presence in the Pleistocene and Holocene on the southern Mediterranean peninsulas. Aesculus fossils are not uncommon within the series of Tertiary taxa. Many appear in the Pliocene and suffer a radical impoverishment in the Lower–Middle Pleistocene transition. Nonetheless some of these tertiary taxa persisted throughout the Pleistocene and Holocene up to the present in the Iberian Peninsula. Locating these refuge areas on the Peninsula is not an easy task, although areas characterised by a sustained level of humidity must have played an predominant role.     

Late Holocene ecological history of Pinus pinaster forests in the Sierra de Gredos of central Spain

This article describes the patterns and processes of vegetation change and fire history in the Late Holocene (c. 2400 calendar year BP) palaeoecological sequence of Lanzahíta, Sierra de Gredos in central Spain, and provides the first Iberian pollen sequence undertaken within a monospecific Pinus pinaster woodland. These new data reassess not only the autochthonous nature of this pine species in the region and the Iberian Peninsula, but also the naturalness of well-developed cluster pine forests. Conflicts of palaeoecological evidence with phytosociological models of vegetation dynamics in the study region, and the relationships of P. pinaster with fire occurrence in Mediterranean-type ecosystems, are discussed.     

Effects of <Emphasis Type="Italic">Pinus pinea</Emphasis> litter on seed germination and seedling performance of three Mediterranean shrub species

Pinus pinea L. is a coniferous tree inhabiting the Mediterranean Basin, which appears widely distributed in the Iberian Peninsula, frequently associated to Mediterranean shrub species, such as Cistus salvifolius L., C. libanotis L. and Halimium halimifolium (L.) Willk. (Cistaceae). This study investigated the effects (chemical and physical) of P. pinea needles on the seed germination and early growth of these three shrub species. Laboratory experiments were carried out in order to assess an expected negative effect by using different treatments to seeds (physical layer and two aqueous extracts concentrations). Seed germination of all species was inhibited when treated with aqueous extracts and the degree of inhibition was higher when using the extract with the highest concentration, while no effect was observed with the physical layer treatment. Seedling performance was also sensitive to the different treatments, showing the most stressed values with the high concentration extract. The results from this study highlight the importance of allelopathy of P. pinea needles as a mechanism for controlling seed germination and seedling performance of some Mediterranean shrub species.     

Climatic niche and neutral genetic diversity of the six Iberian pine species: a retrospective and prospective view

Quaternary climatic fluctuations have left contrasting historical footprints on the neutral genetic diversity patterns of existing populations of different tree species. We should expect the demography, and consequently the neutral genetic structure, of taxa less tolerant to particular climatic extremes to be more sensitive to long‐term climate fluctuations. We explore this hypothesis here by sampling all six pine species found in the Iberian Peninsula (2464 individuals, 105 populations), using a common set of chloroplast microsatellite markers, and by looking at the association between neutral genetic diversity and species‐specific climatic requirements. We found large variation in neutral genetic diversity and structure among Iberian pines, with cold‐enduring mountain species (Pinus uncinata, P. sylvestris and P. nigra) showing substantially greater diversity than thermophilous taxa (P. pinea and P. halepensis). Within species, we observed a significant positive correlation between population genetic diversity and summer precipitation for some of the mountain pines. The observed pattern is consistent with the hypotheses that: (i) more thermophilous species have been subjected to stronger demographic fluctuations in the past, as a consequence of their maladaptation to recurrent glacial cold stages; and (ii) altitudinal migrations have allowed the maintenance of large effective population sizes and genetic variation in cold‐tolerant species, especially in more humid regions. In the light of these results and hypotheses, we discuss some potential genetic consequences of impending climate change.     

Large‐scale assessment of regeneration and diversity in Mediterranean planted pine forests along ecological gradients

Aim There is increasing concern regarding sustainable management and restoration of planted forests, particularly in the Mediterranean Basin where pine species have been widely used. The aim of this study was to analyse the environmental and structural characteristics of Mediterranean planted pine forests in relation to natural pine forests. Specifically, we assessed recruitment and woody species richness along climatic, structural and perturbation gradients to aid in developing restoration guidelines. Location Continental Spain. Methods We conducted a multivariate comparison of ecological characteristics in planted and natural stands of main Iberian native pine species (Pinus halepensis, Pinus pinea, Pinus pinaster, Pinus nigra and Pinus sylvestris). We fitted species‐specific statistical models of recruitment and woody species richness and analysed the response of natural and planted stands along ecological gradients. Results Planted pine forests occurred on average on poorer soils and experienced higher anthropic disturbance rates (fire frequency and anthropic mortality) than natural pine forests. Planted pine forests had lower regeneration and diversity levels than natural pine forests, and these differences were more pronounced in mountain pine stands. The largest differences in recruitment – chiefly oak seedling abundance – and species richness between planted and natural stands occurred at low‐medium values of annual precipitation, stand tree density, distance to Quercus forests and fire frequency, whereas differences usually disappeared in the upper part of the gradients. Main conclusions Structural characteristics and patterns of recruitment and species richness differ in pine planted forests compared to natural pine ecosystems in the Mediterranean, especially for mountain pines. However, management options exist that would reduce differences between these forest types, where restoration towards more natural conditions is feasible. To increase recruitment and diversity, vertical and horizontal heterogeneity could be promoted by thinning in high‐density and homogeneous stands, while enrichment planting would be desirable in mesic and medium‐density planted forests.     

Growth patterns in relation to drought-induced mortality at two Scots pine (Pinus sylvestris L.) sites in NE Iberian Peninsula

Drought-related tree mortality has become a widespread phenomenon. Scots pine (Pinus sylvestris L.) is a boreal species with high ecological amplitude that reaches its southwestern limit in the Iberian Peninsula. Thus, Iberian Scots pine populations are particularly good models to study the effects of the increase in aridity predicted by climate change models. A total of 78 living and 39 dead Scots pines trees were sampled at two sites located in the NE of the Iberian Peninsula, where recent mortality events have been recorded. Annual tree rings were used to (1) date dead trees; (2) investigate if there was an association between the occurrence of tree death and severe drought periods characterized by exceptionally low ratios of summer precipitation to potential evapotranspiration (P/PET); and (3) to compare the growth patterns of trees that died with those of surviving ones. Mixed models were used to describe the relationships between tree growth (in terms of basal area increment, BAI, and the percentage of latewood, LW%) and climate variables. Our results showed a direct association between Scots pine mortality and severe drought periods characterized by low summer water availability. At the two sites, the growth patterns of dead trees were clearly distinguishable from those of the trees that survived. In particular, the BAI of dead trees was more sensitive to climate dryness (low P/PET_summer, high temperatures) and started to decline below the values of surviving neighbors 15–40 years before the time of death, implying a slow process of growth decline preceding mortality.     

Okadaic acid and trifluoperazine enhance <Emphasis Type="Italic">Agrobacterium</Emphasis>-mediated transformation in eastern white pine

Mature zygotic embryos of recalcitrant Christmas tree species eastern white pine (Pinus strobus L.) were used as explants for Agrobacterium tumefaciens strain GV3101-mediated transformation using the uidA (β-Glucuronidase) gene as a reporter. Influence of the time of sonication and the concentrations of protein phosphatase inhibitor (okadaic acid) and kinase inhibitor (trifluoperazine) on Agrobacterium-mediated transformation have been evaluated. A high transformation frequency was obtained after embryos were sonicated for 45–50 s, or treated with 1.5–2.0 μM okadaic acid or treated with 100–200 μM trifluoperazine, respectively. Protein phosphatase and kinase inhibitors enhance Agrobacterium-mediated transformation in eastern white pine. A 2–3.5-fold higher rate of hygromycin-resistant callus was obtained with an addition of 2 μM okadaic acid or 150 μM trifluoperazine or sonicated embryos for 45 s. Stable integration of the uidA gene in the plant genome of eastern white pine was confirmed by polymerase chain reaction (PCR), Southern and northern blot analyses. These results demonstrated that a stable and enhanced transformation system has been established in eastern white pine and this system would provide an opportunity to transfer economically important genes into this Christmas tree species. Communicated by W. H. Wu     

Distribution and population genetic structure of the Mediterranean pine shoot beetle Tomicus destruens in the Iberian Peninsula and Southern France

1 The Mediterranean pine shoot beetle Tomicus destruens has long been indistinguishable from its congeneric Tomicus piniperda. Both species attack pines, and can be found in sympatry. The geographical distribution of T. destruens is still unclear in most of the Mediterranean Basin. 2 We aimed to describe the geographical distribution and zones of sympatry of both species in the Iberian Peninsula and France, and to study the molecular phylogeographical pattern of T. destruens. 3 Tomicus spp. adults were sampled in Portugal, Spain and France, and a portion of the mitochondrial genes COI and COII was sequenced for 84 individuals. Sequences were aligned to a data set previously obtained from French localities. 4 Tomicus destruens was found in all populations, except for one locality in Portugal and in the Landes (France). It was in sympatry with T. piniperda in two locations on Pinus pinaster and one location on Pinus radiata. 5 Within‐population genetic diversity was high, but we found a significant pattern of spatial distribution of genetic variation, as well as a significant effect of the host tree. 6 The data suggest the existence of two glacial refugia, from which T. destruens recolonized its current range. One refugium was located in Portugal where the beetle probably evolved on P. pinaster. The corresponding haplotypes show a West–East frequency gradient. The other refugium was probably in the eastern range, where the beetles evolved on Pinus halepensis and P. pinea. The corresponding haplotypes show an East–West frequency gradient.     

The American brine shrimp as an exotic invasive species in the western Mediterranean

The hypersaline environments and salterns present in the western Mediterranean region (including Italy, southern France, the Iberian Peninsula and Morocco) contain autochthonous forms of the brine shrimp Artemia, with parthenogenetic diploid and tetraploid strains coexisting with the bisexual species A. salina. Introduced populations of the American brine shrimp A. franciscana have also been recorded in these Mediterranean environments since the 1980s. Based on brine shrimp cyst samples collected in these countries from 1980 until 2002, we were able to establish the present distribution of autochthonous brine shrimps and of A. franciscana, which is shown to be an expanding invasive species. The results obtained show that A. franciscana is now the dominant Artemia species in Portuguese salterns, along the French Mediterranean coast and in Cadiz bay (Spain). Co-occurrence of autochthonous (parthenogenetic) and American brine shrimp populations was observed in Morocco (Mar Chica) and France (Aigues Mortes), whereas A. franciscana was not found in Italian cyst samples. The results suggest these exotic A. franciscana populations originate as intentional or non-intentional inoculations through aquacultural (hatchery effluents) or pet market activities, and suggest that the native species can be rapidly replaced by the exotic species. This revised version was published online in July 2006 with corrections to the Cover Date.     

Long-term changes in tree-ring–climate relationships at Mt. Patscherkofel (Tyrol,Austria) since the mid-1980s

Although growth limitation of trees at Alpine and high-latitude timberlines by prevailing summer temperature is well established, the loss of thermal response of radial tree growth during last decades has repeatedly been addressed. We examined long-term variability of climate–growth relationships in ring width chronologies of Stone pine (Pinus cembra L.) by means of moving response functions (MRF). The study area is situated in the timberline ecotone (ca. 2,000–2,200 m a.s.l.) on Mt. Patscherkofel (Tyrol, Austria). Five site chronologies were developed within the ecotone with constant sample depth (≥19 trees) throughout most of the time period analysed. MRF calculated for the period 1866–1999 and 1901–1999 for ca. 200- and ca. 100-year-old stands, respectively, revealed that mean July temperature is the major and long-term stable driving force of Pinus cembra radial growth within the timberline ecotone. However, since the mid-1980s, radial growth in timberline and tree line chronologies strikingly diverges from the July temperature trend. This is probably a result of extreme climate events (e.g. low winter precipitation, late frost) and/or increasing drought stress on cambial activity. The latter assumption is supported by a <10% increase in annual increments of ca. 50-year-old trees at the timberline and at the tree line in 2003 compared with 2002, when extraordinary hot and dry conditions prevailed during summer. Furthermore, especially during the second half of the twentieth century, influence of climate variables on radial growth show abrupt fluctuations, which might also be a consequence of climate warming on tree physiology.     

Patterns of nuclear and mitochondrial DNA variation in Iberian populations of <Emphasis Type="Italic">Emys orbicularis</Emphasis> (Emydidae): conservation implications

The European pond turtle (Emys orbicularis) is threatened and in decline in several regions of its natural range, due to habitat loss combined with population fragmentation. In this work, we have focused our efforts on studying the genetic diversity and structure of Iberian populations with a fine-scale sampling (254 turtles in 10 populations) and a representation from North Africa and Balearic island populations. Using both nuclear and mitochondrial markers (seven microsatellites, ∼1048 bp nDNA and ∼1500 bp mtDNA) we have carried out phylogenetic and demographic analyses. Our results show low values of genetic diversity at the mitochondrial level although our microsatellite dataset revealed relatively high levels of genetic variability with a latitudinal genetic trend decreasing from southern to northern populations. A moderate degree of genetic differentiation was estimated for Iberian populations (genetic distances, F _ST values and clusters in the Bayesian analysis). The results in this study combining mtDNA and nDNA, provide the most comprehensive population genetic data for E. orbicularis in the Iberian Peninsula. Our results suggest that Iberian populations within the Iberian–Moroccan lineage should be considered as a single subspecies with five management units, and emphasize the importance of habitat management rather than population reinforcement (i.e. captive breeding and reintroduction) in this long-lived species.     

The value of leaf cuticle characteristics in the identification and classification of Iberian Mediterranean members of the genus Pinus

This study reports the value of leaf cuticle characteristics in the identification and classification of Iberian Mediterranean species of the genus Pinus (P. nigra subsp. salzmannii, P. pinaster, P. pinea and P. halepensis), with the aim of using these characters to identify isolated cuticles and stomata in palynology slides. Preparations were made of the cuticles of pine needles belonging to one natural Iberian population of each of the above species. A number of epidermal morphological characteristics were then recorded with the aim of distinguishing these species from one another. The structure of the stomatal complex (the shape and arrangement of the subsidiary cells) was different in each species. The aperture of the epistomatal chamber was significantly smaller in P. pinea than in the other species examined, and the variables recorded for the thickening of the guard cells provided relationships that clearly distinguished all four taxa. The width and length of the stomata and the upper woody lamellae, the central distance between the external limits of the medial lamellae borders and the length of the stem were the most useful variables in this respect. The present results contribute to the ongoing discussion regarding the taxonomic classification of the members of Pinus, and provide valuable clues for the identification of Iberian Mediterranean pine species from small pine needle fragments or isolated stomata. After validation of the present results for multiple populations, these results could also be used to help identify fossil leaf macroremains and the scattered/isolated stomata commonly observed in palaeopalynological samples. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 161 , 436–448.     

Breeding performance and longevity of Tomicus destruens on Mediterranean and continental pine species

The pine shoot beetle Tomicus destruens (Wollaston) (Coleoptera: Curculionidae: Scolytinae) is one of the main pests of Mediterranean forests, where it is oligophagous on Mediterranean pines. However, possible global warming may make the insect move to higher latitudes and altitudes, allowing it to attack new pine species. In this respect, the aim of the present article was to assess both the acceptance and performance of T. destruens offered host and non‐host pine species. A no‐choice breeding experiment was set up under laboratory conditions, using logs of three Mediterranean (Pinus pinea L., Pinus pinaster Miller, and Pinus halepensis Aiton) and two continental (Pinus nigra Arnold and Pinus sylvestris L.) pine species. Log debarking at the end of adult emergence assessed parent fecundity, egg, and larval mortality. The quality of callow adults emerging from each tested pine was evaluated on the basis of their longevity on a semiartificial diet. Tomicus destruens colonised all tested pine species, but did not reproduce in Scots pine, taking about 79 days to complete development with no differences among pines tested. The best breeding performance, evaluated as female fecundity and adult production, was observed in P. halepensis, and the lowest in P. pinaster. On average, adults emerging from P. pinea survived longer (83 days) than from other pines, and adult longevity was the lowest in males emerging from P. nigra. Austrian pine, which under natural conditions is usually not a host of T. destruens, allowed insect development and adult production similar to P. pinea and P. pinaster.     

Influence of bombardment with BA-coated microprojectiles on shoot organogenesis fromPhlox paniculata L. andPinus pinea L. tissues

Summary Leaves ofPhlox paniculata L. were bombarded with BA-coated gold microprojectiles with the Biolistic PDS/1000-He gene gun. Increasing the concentration of BA from 0.01 to 0.1 mg/μl increased the mean number of adventitious shoots from 7 to 16 shoots per leaf. Bombardment of cotyledons ofPinus pinea L. (Italian stone pine) with BA-coated gold particles significantly reduced the frequency of shoot organogenesis by about 40%. However, induced shoot regenerants from bombarded explants of bothP. paniculata andP. pinea were longer (>1 cm) than those derived from control explants after 3 and 5 wk following bombardment, respectively.     

Pine volatiles mediate host selection for oviposition by Thaumetopoea pityocampa (Lep., Notodontidae)

The oviposition behaviour and host selection by females of the pine processionary moth, Thaumetopoea pityocampa, was studied under two conditions: in a field insectary and in a pine stand. An oviposition choice test was conducted in an insectary cage, using artificial Christmas trees (ACTs) baited with extracts from four pine species: Pinus pinea, Pinus pinaster, Pinus halepensis and Pinus brutia, plus a control. Females oviposited significantly more egg batches on the ACT baited with a P. brutia extract, while no oviposition occurred on the control ACT. In a large P. pinea stand, two groups of randomly selected trees were marked and baited respectively with an extract of P. brutia, which was the preferred species under insectary conditions and with a solvent, to act as control. Results showed that oviposition in the field followed an aggregated pattern, fitting a negative binomial distribution and that trees baited with P. brutia extracts, received a significantly larger number of egg‐batches than control trees. In parallel, the volatiles emitted by all pine extracts tested where analysed by gas chromatography and mass spectrometry (GC‐MS), coupled to a time‐of‐flight (TOF) mass analyser after solid phase microextraction (SPME), resulting in the identification of 26 compounds. Comparative chromatograms showed qualitative differences among the pine species used, some compounds being present in one of them only. Findings demonstrate for the first time that: (i) T. pityocampa females discriminate among bouquets extracted from different host pine species and exhibit oviposition preferences; and (ii) olfactory cues play an important role in mediating the selection process.     

Prehistoric Pinus woodland dynamics in an upland landscape in northern Scotland: the roles of climate change and human impact

Pollen, microscopic charcoal, palaeohydrological and dendrochronological analyses are applied to a radiocarbon and tephrochronologically dated mid Holocene (ca. 8500–3000 cal b.p.) peat sequence with abundant fossil Pinus (pine) wood. The Pinus populations on peat fluctuated considerably over the period in question. Colonisation by Pinus from ca. 7900–7600 cal b.p. appears to have had no specific environmental trigger; it was probably determined by the rate of migration from particular populations. The second phase, at ca. 5000–4400 cal b.p., was facilitated by anthropogenic interference that reduced competition from other trees. The pollen record shows two Pinus declines. The first at ca. 6200–5500 cal b.p. was caused by a series of rapid and frequent climatic shifts. The second, the so-called pine decline, was very gradual (ca. 4200–3300 cal b.p.) at Loch Farlary and may not have been related to climate change as is often supposed. Low intensity but sustained grazing pressures were more important. Throughout the mid Holocene, the frequency and intensity of burning in these open Pinus–Calluna woods were probably highly sensitive to hydrological (climatic) change. Axe marks on several trees are related to the mid to late Bronze Age, i.e., long after the trees had died.     

Morphometric and chemical fruit variability of selected stone pine trees (Pinus pinea L.) grown in non-native environments

Few studies have been conducted on Stone pine (Pinus pinea) morphology, cone to kernel yield (KY) and composition. To study the species variability, a morphometric and chemical characterization of pine nuts was performed across the species distribution in Chile. Seventy-six highly productive trees were selected in three macrozones. Ten cones per tree were harvested, and cone weight, in-shell pine nut number cone^?1 and size and weight of in-shell pine nuts and kernels were measured; KY was determined. Chemical and nutritional characterization of pine nuts was performed in 38 trees. Differences among macrozones were tested with mixed linear models. Partial correlation was used to test for correlation between cone and pine nut morphometric data, chemical features, and climate. Canonical correlation coefficients between groups were tested. Selected trees differed in cone and in-shell pine nut size, kernel morphometry, and KY, with lowest trait values being generally found in the coast. KY was higher than in Europe. Chemical composition was similar to values reported in producer countries, homogeneous among macrozones, with the exception of oleic acid and potassium. Detected differences would respond to the environment and to the interaction genotype?×?environment given the restricted origins of the genetic material, confirming the species adaptability.     

Reconstructing the history of an invasion: the toxic phytoplankton species <Emphasis Type="Italic">Gymnodinium catenatum</Emphasis> in the Northeast Atlantic

The phytoplankton species Gymnodinium catenatum is responsible for major worldwide losses in aquaculture due to shellfish toxicity. On the West coast of the Iberian Peninsula, toxic blooms have been reported since the mid-1970s. While the recent geographical spread of this species into Australasia has been attributed to human-mediated introduction, its origin in the Northeast Atlantic is still under debate. Gymnodinium catenatum forms a highly resistant resting stage (cyst) that can be preserved in coastal sediments, building-up an historical record of the species. Similar cyst types (termed microreticulate) are produced by other non-toxic Gymnodinium species that often co-occur with G. catenatum. We analysed the cyst record of microreticulate species in dated sediment cores from the West Iberian shelf covering the past ca. 150 years. Three distinct morphotypes were identified on the basis of cyst diameter and paracingulum reticulation. These were attributed to G. catenatum (35.6–53.3 μm), G. nolleri (23.1–36.4 μm), and G. microreticulatum (20.5–34.3 μm). Our results indicate that G. catenatum is new to the NE Atlantic, where it appeared by 1,889 ± 10, expanding northwards along the West Iberian coast. The earliest record is from the southernmost sample, while in the central Portuguese shelf the species appears in sediments dated to 1,933 ± 3, and in the North, off Oporto, in 1,951 ± 4. On the basis of the cyst record and toxic bloom reports, we reconstruct the invasive pathway of G. catenatum in the NE Atlantic. Although human-mediated introduction cannot be discarded, the available evidence points towards natural range expansion, possibly from NW Africa.     

Invasion biology of Australian ectomycorrhizal fungi introduced with eucalypt plantations into the Iberian Peninsula

In the last two centuries, several species of Australian eucalypts (e.g. Eucalyptus camaldulensis and E.␣globulus) were introduced into the Iberian Peninsula for the production of paper pulp. The effects of the introduction of exotic root-symbitotic fungi together with the eucalypts have received little attention. During the past years, we have investigated the biology of ectomycorrhizal fungi in eucalypt plantations in the Iberian Peninsula. In the plantations studied, we found fruit bodies of several Australian ectomycorrhizal fungi and identified their ectomycorrhizas with DNA molecular markers. The most frequent species were Hydnangium carneum, Hymenogaster albus, Hysterangium inflatum, Labyrinthomyces donkii, Laccaria fraterna, Pisolithus albus, P. microcarpus, Rhulandiella berolinensis, Setchelliogaster rheophyllus, and Tricholoma eucalypticum. These fungi were likely brought from Australia together with the eucalypts, and they seem to have facilitated the establishment of eucalypt plantations and their naturalization. The dispersion of Australian fungal propagules may be facilitating the spread of eucalypts along watercourses in semiarid regions increasing the water lost. Because ectomycorrhizal fungi are obligate symbionts, their capacity to persist after eradication of eucalypt stands, and/or to extend beyond forest plantations, would rely on the possibility to find compatible native host trees, and to outcompete the native ectomycorrhizal fungi. Here we illustrate the case of the Australasian species Laccaria fraterna, which fruits in Mediterranean shrublands of ectomycorrhizal species of Cistus (rockroses). We need to know which other Australasian fungi extend to the native ecosystems, if we are to predict environmental␣risks associated with the introduction of Australasian ectomycorrhizal fungi into the Iberian Peninsula. This revised version was published online in July 2006 with corrections to the Cover Date.