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.     

Effects of exotic and native tree leaf litter on soil properties of two contrasting sites in the Iberian Peninsula

Aims

We assessed the effects of native and exotic tree leaf litter on soil properties in two contrasting scenarios. The native Quercus robur and Pinus pinaster tree species coexist with the aliens Eucalyptus globulus and Acacia dealbata in acid soils of NW Spain. The native trees Fraxinus angustifolia and Ulmus minor coexist with the aliens Ailanthus altissima, Robinia pseudoacacia and Ulmus pumila in eutrophic basic riparian soils in Central Spain.

Methods

Four plastic trays per species were filled with homogenized top-soil of the site and covered with leaf litter. Before and after 9?months of incubation, litter mass, soil pH, organic matter, mineral and total N were measured. Available mineral N (NO ₃ ^? -N and NH ₄ ⁺ -N) was assessed every 2?months.

Results

Soil biological activity was higher in the basic than in the acid soil. Litter of the exotic trees tended to decompose less than litter of native species, probably due to the presence of secondary metabolites in the former. Soil pH, mineral and total N responded differently to different litter types, irrespective of their exotic or native origin (acid soil), or was similar across litter treatments (basic riparian soil). The similar response of the basic soil to the addition of different litter types may be due to the low contrast of litter quality between the species. E. globulus litter inhibitied soil microbial activity much more than the rest of the studied litter types, leading to a drastic impoverishment of N in soils.

Conclusion

Litter of exotic N-fixing trees (A. dealbata and R. pseudoacacia) did not increase soil N pools because of the inhibition of microbial activity by secondary compounds. Therefore, secondary metabolites of the litter played a major role explaining exotic litter impact on soil properties.     

Ground-truthing the impact of invasive species: spatio-temporal overlap between native least chub and introduced western mosquitofish

Western mosquitofish (Gambusia affinis) are one of the most destructive introduced species in the American West. The negative impact of introduced species on native taxa depends on their spatio-temporal overlap, which will determine the availability of refugia for native species. Experiments on the mechanisms underlying the interactions between introduced and native species rarely address habitat use, overlap, and refugia because individuals are confined to enclosures. In a previous study we used cages, microcosms, and aquaria to show that mosquitofish could prey on and out-compete native least chub (Iotichthys phlegethontis). In this study, we examined the spatio-temporal overlap between mosquitofish and least chub under natural conditions. We found periods of overlap and partitioning in the seasonal and diel habitat use of these species. Both species used shallow habitats during the day and night throughout the spring when least chub were spawning. Predation by adult mosquitofish on young least chub during the spring likely explains the reduction in least chub recruitment in the presence of mosquitofish. During the summer least chub avoided mosquitofish by exclusively using cooler habitats, or by occupying deeper, cooler habitats during the day when mosquitofish were active, and shallower, warmer habitats at night when mosquitofish were inactive. A shift to cooler habitats in the presence of mosquitofish may result in decreased growth and fecundity of least chub. However, we suggest that a reduction of warm shallow habitat may have a disproportionately greater negative impact on mosquitofish than species native to temperate regions, such as least chub. Habitat manipulations that reduce or eliminate warm habitats may ameliorate the harmful effects of mosquitofish and promote the long-term persistence of native taxa.     

Effects of herbivory and competition on an introduced plant in decline

Anthemiscotula was introduced to Denmark 500 years ago, and its distribution is presently limited and in decline. A manipulative field experiment was performed to investigate the effects of native plant competitors and native invertebrate herbivores on its performance. Generally, both herbivory and competition treatments had great impact, and when both factors were operating, the effects were additive for all variables except plant height. Although A. cotula showed plasticity in growth, resource allocation and flowering timing, it was unable to adjust to competition and compensate for losses due to herbivory sufficiently to ensure and restore its achene production. This vulnerability, combined with improved cereal cleaning techniques and thus fewer reintroductions of A. cotula seeds, may be the cause of its current decline. A. cotula responded to herbivory by prolonging its flowering period, a “bet-hedging” strategy. In Denmark this strategy is unreliable since risks of sub-optimal conditions are much greater in August–October. Received: 12 December 1997 / Accepted: 14 October 1998     

Comparative phylogeography of endemic cyprinids in the south-west Iberian Peninsula: evidence for a new ichthyogeographic area

The comparative phylogeography and evolutionary history of three native cyprinid genera, Squalius (subfamily Leuciscinae), Chondrostoma (subfamily Leuciscinae) and Barbus (subgenus Luciobarbus ; subfamily Cyprininae), were examined focusing mainly in the South-Western region of the Iberian Peninsula, where recently described endemic species are present with considerably restricted distribution areas. In order to accomplish that the variation at the complete cytochrome b gene (1140 bp) was analysed for specimens from the South-Western region, and also for representatives of the three genera from all over the Iberian Peninsula. Data indicate different evolutionary histories, with distinct time periods of colonization between the two cyprinid subfamilies in the Iberian Peninsula. Four new Iberian ichthyogeographic areas are accordingly proposed based on congruent phylogeographic and geological evidences: the South-Western, the South-Eastern, the Atlantic and the Mediterranean. Evidence was provided for the older isolation of the South-Western area in the Miocene during the Endorheic Drainages phase, designating a clearly defined and distinct ichthyogeographic area. A new molecular clock calibration is proposed for the subgenus Luciobarbus .     

Genotyping an ALAD polymorphism with real-time PCR in two populations from the Iberian Peninsula

Lead-induced toxicity varies among individuals partly because of genetic differences in their susceptibility to the metal’s effects. One genetic polymorphism associated with lead toxicity is a G-to-C transversion at position 177 in the coding region of delta-aminolevulinic acid dehydratase (ALAD), originating from two codominant alleles (ALAD1 and ALAD2). We examined the distribution of this single nucleotide polymorphism in two populations from the Iberian Peninsula. Genomic DNA was extracted from whole blood, and a real-time PCR assay was designed to determine ALAD polymorphic distribution. The distribution of MspI polymorphism was similar in the two populations studied, and allelic frequencies were comparable to those obtained in other studies of Caucasians. Further studies are needed to assess fully the functional significance of this polymorphism and its influence on the toxicokinetics of lead.     

Genetic evidence for multiple events of hybridization between wolves and domestic dogs in the Iberian Peninsula

Hybridization between wild species and their domestic counterparts may represent a major threat to natural populations. However, high genetic similarity between the hybridizing taxa makes the detection of hybrids a difficult task and may hinder attempts to assess the impact of hybridization in conservation biology. In this work, we used a combination of 42 autosomal microsatellites together with Y-chromosome microsatellite-defined haplotypes and mtDNA sequences to investigate the occurrence and dynamics of wolf-dog hybridization in the Iberian Peninsula. To do this, we applied a variety of Bayesian analyses and a parallel set of simulation studies to evaluate (i) the differences between Iberian wolves and dogs, (ii) the frequency and geographical distribution of hybridization and (iii) the directionality of hybridization. First, we show that Iberian wolves and dogs form two well-differentiated genetic entities, suggesting that introgressive hybridization is not a widespread phenomenon shaping both gene pools. Second, we found evidence for the existence of hybridization that is apparently restricted to more peripheral and recently expanded wolf populations. Third, we describe compelling evidence suggesting that the dynamics of hybridization in wolf populations is mediated by crosses between male dogs and female wolves. More importantly, the observation of a population showing the occurrence of a continuum of hybrid classes forming mixed packs may indicate that we have underestimated hybridization. If future studies confirm this pattern, then an intriguing avenue of research is to investigate how introgression from free-ranging domestic dogs is enabling wolf populations to adapt to the highly humanized habitats of southern Europe while still maintaining their genetic differentiation.     

Aggression and competition for shelter between a native and an introduced crayfish in Europe

1. The introduced North American crayfish, Pacifastacus leniusculus Dana, is expanding its range in Europe and locally often replaces two native crayfish species, Astacus astacus L. and Austropotamobius pallipes Lereboullet. Pacifastacus leniusculus is also expected to invade the habitat of a third native crayfish, the endangered Austropotamobius torrentium Schrank. Interspecific aggressive interactions and competition for shelter were experimentally studied in the laboratory to assess the potential impact of P. leniusculus on A. torrentium .
2. Neither species was inherently dominant in aggressive interactions, but dominance was strongly size-dependent, favouring the larger and faster growing species, P. leniusculus .
3. Access to limited shelter was generally determined by aggressive dominance, although species-specific preference also influenced the outcome of competition for shelter. Austropotamobius torrentium had a higher preference for experimental shelters and often defended these even against larger P. leniusculus .
4. In accordance with theoretical models of animal conflicts, agonistic interactions between equally sized contestants were more severe than between animals of different size.
5. The P. leniusculus used in the present experiment were infected with the crayfish plague, Aphanomyces astaci , to which the animals are resistant. The crayfish transmitted the disease to non-resistant A. torrentium which died ≈ 2 weeks after contact with P. leniusculus .     

Morphological dispersion of introduced Hawaiian finches: evidence for competition and a Narcissus effect

Summary Morphological analyses were conducted on finch species introduced to Oahu, Hawaii. As many as 25 species of finches representing four families (Emberizidae, Fringillidae, Ploecidae, Estrildidae) have been introduced to Oahu. Of these, 15 species currently have established wild populations. When compared with pools of 25 and 23 species introduced to Oahu, the 15 surviving species are morphologically overdispersed. This result is consistent with the hypothesis that competition has influenced the outcome of these introductions. We also tested the hypothesis that surviving introduced finches in one habitat (sugarcane fields) were morphologically overdispersed. When the seven surviving finch species found in sugarcane were compared with the pools of 25 and 23 species, they too were morphologically overdispersed. However, when the seven sugarcane species were compared with a species pool consisting of only the surviving 15 species found across Oahu, they were not morphologically overdispersed. This result suggests that morphological analyses of community structure based only on comparisons of assemblages of surviving species may be biased by a Narcissus effect.     

Alu insertion polymorphisms in NW Africa and the Iberian Peninsula: evidence for a strong genetic boundary through the Gibraltar Straits

An analysis of 11 I Alu insertion polymorphisms (ACE, TPA25, PV92, APO, FXIIIB, D1, A25, B65, HS2.43, HS3.23, and HS4.65) has been performed in several NW African (Northern, Western, and Southeastern Moroccans, Saharawi; Algerians; Tunisians) and Iberian (Basques, Catalans, and Andalusians) populations. Genetic distances and principal component analyses show a clear differentiation of NW African and Iberian groups of samples, suggesting a strong genetic barrier matching the geographical Mediterranean Sea barrier. The restriction to gene flow may be attributed to the navigational hazards across the Straits, but cultural factors must also have played a role. Some degree of gene flow from sub-Saharan Africa can be detected in the southern part of North Africa and in Saharawi and Southeastern Moroccans, as a result of a continuous gene flow across the Sahara desert that has created a south-north cline of sub-Saharan Africa influence in North Africa. Iberian samples show a substantial degree of homogeneity and fall within the cluster of European-based genetic diversity.     

The Carboniferous floras of the Iberian Peninsula: A synthesis with geological connotations

A general review is presented of the Carboniferous floral records in the Iberian Peninsula in the context of the geological history and distribution of the different basins. Mississippian floras are found in Sierra Morena, where major strike-slip faults brought in terranes of diverse provenance. Lower Pennsylvanian floras are represented in the Peñarroya-Belmez-Espiel (Córdoba) and Villanueva del Río y Minas (Sevilla) coalfields of SW Spain (also strike-slip controlled), at La Camocha, near Gijón (Asturias), and in other parts of northern Spain. Middle Pennsylvanian is represented near Oporto, but more completely in the Central Asturian Coalfield, as well as other, more limited localities in NW Spain and the Pyrenees. Upper Pennsylvanian (Stephanian Stage-Cantabrian to Stephanian B substages) floras are splendidly represented in NW Spain. Uppermost Pennsylvanian (Stephanian C-Autunian) floras are present in the strike-slip controlled Douro and Buçaco basins of North Portugal, the Pyrenees, Central Spain (Ciudad Real, Guadalajara, Zaragoza), and SW Spain (Guadalcanal and Valdeviar in Sevilla province).A complete succession of megafloral zones is presented. This includes a new Annularia spicata Zone at the top of the Pennsylvanian (equivalent to middle to upper Autunian which has been often attributed to the Lower Permian). The information is summarised in charts compiled from a selection of the most significant species. A number of floral elements are illustrated including the zonal indices. Brief taxonomic comments are provided in the Appendix.     

The Cenozoic vegetation of the Iberian Peninsula: A synthesis

The aim of this work is to provide a first approach to the evolution of Iberia's vegetation during the Cenozoic (with the exclusion of the Quaternary). The Palaeogene was floristically defined by Palaeotropical elements forming tropical/subtropical rainforests, mangrove swamps, edaphically-mediated laurophyllous forests and leguminous-sclerophyllous communities. During the Miocene, Iberian landscapes were drastically modified due to geographic and climatic changes (mainly cooling and aridification) changes. Open, steppe-like environments developed towards the interior of the peninsula and Arctotertiary elements invaded mountainous and riparian ecosystems, coexisting with or becoming part of evergreen, broadleaved forests of Palaeotropical species. From the Late Miocene onwards these forests suffered changes due to the extinction of taxa, the impact of environmental change on the survivors, and the perturbations caused by the arrival of further Arctotertiary elements. However, several Palaeotropical taxa overcame the environmental and climatic changes of the Miocene and Pliocene to form a part of the modern flora of the Iberian Peninsula.     

Comparison of competitive ability between native and introduced salmonids: evidence from pairwise contests

Brown trout, Salmo trutta, and rainbow trout, Oncorhynchus mykiss, have been introduced to freshwaters in Hokkaido, Japan. Today, it is recognized that these introduced salmonids have negative impacts on native salmonids such as white-spotted charr, Salvelinus leucomaenis, and masu salmon, O. masou. In particular, interspecific competition may be an important mechanism that could contribute to the exclusion for native salmonids. In this study, experimental pairwise contests were conducted to compare interference competitive ability between native and introduced salmonids. We demonstrated that brown trout were competitively superior to white-spotted charr and masu salmon whereas rainbow trout were superior to white-spotted charr. We suggest that introduced brown trout negatively impact both white-spotted charr and masu salmon, and introduced rainbow trout negatively impact white-spotted charr.     

The non-omphalinoid species of Arrhenia in the Iberian Peninsula

A taxonomic study of the species with nutant, pleurotoid and cyphelloid basidiomata of the genus Arrhenia in the Iberian Peninsula is presented. This study is based on the examination of recent specimens collected in the field and from dried herbarium collections. Five species and one variety of this genus are recognized on the basis of morphological and ecological features: Arrhenia acerosa var. acerosa, A. acerosa var. tenella, A. auriscalpium, A. lobata, A. retiruga and A. spathulata. Arrhenia auriscalpium is new to Spain, where it is not restricted to alpine zones of the Eurosiberian region and is more common in the Mediterranean region than previously reported. In contrast, A. acerosa var. tenella currently is known from the Eurosiberian subalpine belt and represents the first report to the Iberian Peninsula, and A. retiruga is known only from the mesomediterranean to supramediterranean belts of the Mediterranean region. Arrhenia acerosa var. acerosa, A. lobata and A. spathulata are widespread in both the Eurosiberian and the Mediterranean regions. Non-Iberian materials of A. acerosa var. tenella and A. spathulata also were studied for comparison. Cyphella cochlearis var. subsessilis, Dictyolus lagunae, Leptoglossum muscigenum var. azonum and Pleurotellus acerosus var. tenellus are lectotypified. Cyphella cochlearis var. subssesilis, Dictyolus lagunae and Leptoglossum muscigenum var. azonum are synonymized with Arrhenia spathulata, and Cyphella cochlearis var. auriformis is placed in synonymy with A. auriscalpium. These taxa are illustrated, described and discussed, based on Iberian material with emphasis on features of the basidioma, pigment of the pileipellis, presence or absence of clamps and shape and size of the basidiospores. A diagnostic key also is given.     

Numerical syntaxonomy of the Asplenietalia petrarchae in the Iberian Peninsula

Abstract. A numerical syntaxonomic study of rupicolous vegetation of the Asplenietalia petrarchae in the Iberian Peninsula is described. The study is based on 37 local tables (OSUs) of different rupicolous communities from limestone cliffs found in the eastern half of the Iberian Peninsula. Three classification approaches were followed: (1) ‘hard partition’ techniques, (2) ‘fuzzy partition’ and (3) ordination partition. In order to find a direct solution, we tried an evaluation system based on the inertia associated with a CCA, Canonical Correspondence Analysis, and its relationships with the inertia of a similar CA, Correspondence Analysis. The most effective solutions were found through spatial partitions, and the best partition was obtained with CA considering five segments per axis. Among the fuzzy partitions the best solution was obtained with two clusters and a fuzziness value of 1.20. The best hierarchical partition was Complete Linkage with Chord Distance as a resemblance function. With the GRID procedure applied to the CA diagram, three different groups were distinguished. The first group is related to the Saxifragion boisseriano-reuterani, including five OSUs from Baetic ranges; the second group is very close to the Cosentinio-Lafuenteion rotundifoliae and includes 11 OSUs from thermo-mediterranean areas; the third group, including 21 OSUs basically related to meso-mediterranean areas, is strongly related to the Teucrion buxifolii.     

Fire and bones: Bronze Age III in the North-Eastern Iberian Peninsula

Between the third millennium BP and the fifth century AD, there are numerous necropolises that house urns in the Iberian Peninsula. One example is the "Pi de la Lliura" (Vidreres, Girona). However, there is a dearth of research concerning these structures. The "Pi de la Lliura" housed 43 structures, which contained a total of 47 vessels with human remains. Of these 43 structures, 22 were totally or partially excavated at the laboratory. The evidence from the fragments indicates cremation at a temperature of 650-700 degrees C. Part of the cremated corpse was then deposited in an urn. One of the most unique characteristics of the necropolis is the high frequency of individuals younger than 20 years old. The mortality of sub-adults is high in any prehistoric necropolis, but it is even higher in a cremation necropolis. "Pi de la Lliura" is a very small necropolis, where corpses were treated similarly over a short period.     

Explanation for the decline of bacteria introduced into lake water

The sizes of the populations of individual bacterial species diminished following their addition to water from lakes with different trophic levels at temperatures of 5, 10, 15, and 30°C. Some species persisted after their initial reduction in cell numbers, but others were undetectable after 3 to 15 days. The decline of these introduced bacteria was not a result of their inoculation at higher densities than are found in nature. The death of most of the test species was not the result of starvation, abiotic factors, bdellovibrios, or bacteriophages. Despite the presence of lytic bacteria, the lake water did not have lytic activity against the test species. Protozoan predation was a significant factor in the fall in bacterial population sizes because protozoa increased in numbers as the bacterial density fell, the suppression of protozoa led to the elimination or delay of the decline of the bacteria, and the addition of protozoa to lake water in which indigenous protozoa were suppressed produced the same pattern of bacterial elimination as in untreated lake water.