Issues in Bioinvasion Science The editors wish to acknowledge the following referees: Benigno Elvira, Universidad Complutense de Madrid, Spain Jose´ Esteban Dura´n, Instituto Nacional de Investigacio´n y Tecnolog´ıa Agraria y Alimentaria, Spain Piero Genovesi, National Wildlife Institute, Italy Fe´lix Llamas Garc´ıa, Universidad de Leo´n, Spain Marco Masseti, Universita` degli Studi di Firenze, Italy Nicola´s Pe´rez Hidalgo, Universidad de Leo´n, Spain Mar´ıa Antonia Ribera Siguan, Universidad de Barcelona, Spain Daniel Simberloff, University of Tennessee, USA Daniel Sol, McGill University, Canada Issues in Bioinvasion Science EEI 2003: a Contribution to the Knowledge on Invasive Alien Species Edited by Laura Capdevila-Argu¨elles and Bernardo Zilletti G.E.I. – GrupoEspecies Invasoras,C.B.I. –Centro Invasiones Biolo´gicas,Leo´n,Spain Reprinted fromBiological Invasions, Volume 7(1),2005. 123 A C.I.P.cataloguerecordforthisbook isavailable fromtheLibrary of Congress ISBN1-4020-2902-0 Published bySpringer, P.O. 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Printed in the Netherlands Table of contents Introduction Laura Capdevila-Argu¨elles & Bernardo Zilletti 1–2 Invasion biology of Australian ectomycorrhizal fungi introduced with eucalypt plantations into the Iberian Peninsula Jesu´s D´ıez 3–15 Phenological and demographic behaviour of an exotic invasive weed in agroecosystems Jordi Recasens, V´ıctor Calvet, Alicia Cirujeda & Josep Antoni Conesa 17–27 Short-term responses to salinity of an invasive cordgrass Jesu´s M. Castillo, Alfredo E. Rubio-Casal, Susana Redondo, Antonio A. A´lvarez-Lo´pez, Teresa Luque, Carlos Luque, Francisco J. Nieva, Eloy M. Castellanos & Manuel E. Figueroa 29–35 The American brine shrimp as an exotic invasive species in the Western Mediterranean Francisco Amat, Francisco Hontoria, Olga Ruiz, Andy J. Green, Marta I. Sa´nchez, Jordi Figuerola & Francisco Hortas 37–47 Impact of an introduced Crustacean on the trophic webs of Mediterranean wetlands Walter Geiger, Paloma Alcorlo, Angel Baltana´s & Carlos Montes 49–73 Loss of diversity and degradation of wetlands as a result of introducing exotic crayfish C.F. Rodr´ıguez, E. Be´cares, M. Ferna´ndez-Ala´ez & C. Ferna´ndez-Ala´ez 75–85 Worldwideinvasionofvectormosquitoes:presentEuropeandistributionandchallengesforSpain Roger Eritja, Rau´l Escosa, Javier Lucientes, Eduard Marque`s, Ricardo Molina, David Roiz & SantiagoRuiz 87–97 Holocene turnover of the French vertebrate fauna Michel Pascal & Olivier Lorvelec 99–106 Life-history traits of invasive fish in small Mediterranean streams Anna Vila-Gispert, Carles Alcaraz & Emili Garc´ıa-Berthou 107–116 Genetic introgression on freshwater fish populations caused by restocking programmes Mar´ıa Jose´ Madeira, Benjam´ın J. Go´mez-Moliner & Annie Machordom Barbe´ 117–125 Eradications of invasive alien species in Europe: a review Piero Genovesi 127–133 Frenchattemptstoeradicatenon-indigenousmammalsandtheirconsequencesfornativebiota Olivier Lorvelec & Michel Pascal 135–140 Successful eradication of invasive rodents from a small island through pulsed baiting inside covered stations JorgeF.Orueta,YolandaAranda,Toma´sGo´mez,GerardoG.Tapia&LinoSanchez-Ma´rmol 141–147 BiologicalInvasions(2005)7:1–2 (cid:1)Springer2005 Introduction The prevention and minimization of the impact on biological invasions carried out in Spain, the of invasive alien species has become a priority on GEI (Grupo Especies Invasoras) organised the the agenda of the international community First National Conference on Invasive Alien because of their negative impact on biodiversity, Species in 2003. More than 150 people from dif- economy and public health. This has led in ferent parts of the peninsula and from other recent years to the development of tools (legal, countries belonging to different institutions (uni- technical, etc.) and programmes. It has become versities, public administration, NGOs) clear that the scale of the problem requires an responded to this ‘call for action’ and partici- interdisciplinary approach, the planning of long- pated in the conference. term actions (strategic perspective), and urgency. The present publication, which contains some Biological invasions are a global problem of the contributions of the First National Con- whose local impact can be of great magnitude. ference on Invasive Alien species, aims to Effective responses are needed, cooperation and increase global knowledge on invasive alien spe- coordination among countries and/or different cies. sectors of society are a must. In this frame, scien- For editing this issue we counted on the help tific research is fundamental to improving knowl- of a large number of people. We are grateful edge on invasive alien species and developing to the authors for their contributions and full new methodologies to fight them. response to the suggestions and requirement of Big steps to improve the understanding of bio- the editors. The committee of referees played a logical invasions have been taken in recent years very important role in the project. To all of thanks to the increase of basic research in this them, for their patient work, invaluable intellec- field. However, in spite of the successes obtained tual contribution and support, our most sincere and enlightening information gained on ‘the bio- thanks. Our gratitude goes to James T. Carlton logical invasions phenomenon’, there are still and James Drake, who have succeeded each many gaps to be filled. The implementation of other in the scientific direction of Biological models, the improvement of mitigation methods Invasions, for having granted us the opportu- (more effective and more environmentally nity to publish the present issue and for super- friendly) and prevention techniques (e.g. risk vising its scientific quality. We owe thanks to analysis) are a priority. However given that inva- Springer and, in particular, to Suzanne Mek- sive alien species represent a global threat, it is king and Ellen Girmscheid who have helped necessary to make the best use of the current bring this project to completion. information available. We are deeply indebted to all the institutions It is absolutely necessary to optimise the who sponsored the conference, especially to Fun- efforts in scientific research, establishing mecha- dacio´n Biodiversidad, which funded, among nisms to develop the exchange of information other things, the present edition. on invasive alien species, keeping in mind that Thanks are also due to the GISP (Global Inva- in the case of biological invasions solutions to a sive Species Programme), ISSG/IUCN (Invasive problem may be found in other parts of the Species Specialists Group/World Conservation world. Union) and COE (Council of Europe), who have In this spirit and with the objective, among supported our work and participated in the con- others, of combining and sharing the experiences ference. 2 Last, but not least, we extend our warmest Laura Capdevila-Argu¨elles & Bernardo Zilletti thanks to Maj de Poorter, Piero Genovesi, G.E.I. – Grupo Especies Invasoras Michel Pascal, Steve Raaymakers, Eladio Fern- c/ Moise´s de Le´on N. 22 Bajo a´ndez-Galiano Daniel Simberloff, Richard N. 24006 Le´on Mack and Ana Isabel Queiroz for their constant Spain and ever present help and support. E-mail: [email protected] Funded by Other sponsors Scientific patronage BiologicalInvasions(2005)7:3–15 (cid:1)Springer2005 Invasion biology of Australian ectomycorrhizal fungi introduced with eucalypt plantations into the Iberian Peninsula Jesu´ s Dı´ez Departamento Biologı´a Vegetal, Universidad de Alcala´, 28871 Alcala´ de Henares (Madrid), Spain (e-mail: [email protected]; fax: +34-91-8855066) Received4June2003;acceptedinrevisedform30March2004 Key words: ectomycorrhizal fungi, Eucalyptus, exotic fungi, forest plantations, invasion ecology Abstract 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 atten- tion. During the past years, we have investigated the biology of ectomycorrhizal fungi in eucalypt plan- tations 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 fre- quent 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 nat- uralization. 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 obli- gate 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 frater- na, 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 environ- mental risks associated with the introduction of Australasian ectomycorrhizal fungi into the Iberian Peninsula. Introduction Mycorrhizal fungi are essential in plant nutri- tion in terrestrial ecosystems, and terrestrial Alien plants often require mutualistic partners to plants present different types of obligate mycor- overcome barriers to establishment in foreign rhizal symbioses. Herbs and shrubs form arbuscu- environments. Mutualisms that facilitate inva- lar mycorrhizas (AM) with glomales (Smith and sions occur at several phases of the life cycle of Read 1997). The low specificity of AM relation- alien invading plants. However, even when wind ships and the easy acquisition of mutualistic sym- and native generalist animals mediate flower pol- bionts by herbs and shrubs in any ecosystems are lination and seed spread, the lack of compatible important reasons for so many ecosystems being mycorrhizal symbionts can limit the spread of susceptible to invasion by alien plants. As a con- alien plants (Richardson et al. 2000a). sequence, the introduction of AM fungi does not 4 seem toplayamajor role inmediating plant inva- terranean forests. To the best of our knowledge, sions, except on some remote islands that are this is the first investigation dealing with the inva- poor in AM fungi (Richardson et al. 2000a). sion ecology of exoticECM fungi. Mostforesttrees,however,associate withagroup of basidiomycetes and ascomycetes forming ecto- mycorrhizas (ECM) (Newman and Reddell 1987). Exotic forestry: invasions of alien trees Ectomycorrhizal symbioses present a range of host–fungus specificities. For this reason, exotic Origin of exotic forestry forestry has often needed the introduction of compatible ectomycorrhizal fungal symbionts There is evidence of large-scale forestation in the (Grove and Le Tacon 1993). For many non- ancient Mediterranean basin, where timber- and native trees, notably for pines and eucalypts, the crop-producing trees were planted as long ago as lack of symbionts was a major barrier to estab- 255 B.C. (Zobel et al. 1987). Inspite of its long lishmentandinvasioninthesouthernhemisphere, history, the scale of forestry remained small until before the build-up of inoculums through human recently. Large-scale forestry was not widespread activity (Armstrong and Hensbergen 1996; Davis until the second half of the 20th century (Zobel et al. 1996; Richardson et al. 2000a). The role of et al. 1987), when pines and eucalypts were ECM fungi in facilitating the establishment and widely planted outside their natural ranges. invasion of alien trees was claimed by Richardson Pines, which comprise only Holarctic species, et al. (1994) to explain the invasion patterns of were planted in South America, South Africa pines in South Africa. Little investigation, how- and Australia. The Australasian eucalypts were ever, has been done on the role played by the planted worldwide. In particular, the need for ECM fungi in the naturalization of eucalypts increased wood production to improve living beyond their natural range. conditions made the genus Eucalyptus one of the This paper provides a framework for thinking most widely planted silvicultural crops. In addi- about the effects of the introduction of exotic tion, in many damaged ecosystems, afforestation fungi with plantations of exotic forest trees. Dif- with alien eucalypts was driven by the belief that ferent sections will deal with aspects of the inva- such plantings were beneficial to the environment sion biology of eucalypts and their ECM fungi. (Zobel et al. 1987). We first lay the groundwork for our paper by briefly introducing the history of the exotic for- Naturalization and invasion of pines and eucalypts estry and several terms used in studies of biologi- cal invasions. The paper will be illustrated with a In this paper, we will use the following three con- study conducted on exotic plantations of eucalyp- cepts: introduction, naturalization and invasion, ts in the Iberian Peninsula. We will describe ecto- as defined by Richardson et al. (2000b). A tree mycorrhizal communities of Australasian fungi introduction takes place when humans transport present in these plantations. We will next analyze a tree across a geographical barrier to a new area. the role that ECM fungi plays in promoting (or Naturalization refers to the species establishing limiting) invasion rates of the eucalypts intro- new self-perpetuating populations and becoming duced. The central core of the paper will try to incorporated within the native flora. The natural- explain the lags between the introduction and the izedtreesregeneratefreely,butmainlyundertheir spread of the eucalypts, as a factor that depends own canopies. In contrast, invasive species recruit on the dispersion of propagules of introduced seedlings,ofteninverylargenumbers,atlongdis- ECM fungi. We report host shifts of Australian tances from parent plants (often more than fungi to native ectomycorrhizal plants detected to 100 m). Only some of the naturalized plants date. We discuss whether the introduced fungi become invasive, producing important environ- could threaten natural communities of ECM mentalor economical damages. fungi by out-competing the native ECM fungi All trees that are widely planted in alien envi- from their natural hosts. This work includes an ronments can become invasive and spread under analysis of the potential effects of the invasion of certain conditions (Richardson 1998). Conse- these exotic fungi in the nutrient cycling of Medi- quently, the use of exotic trees has often caused 5 environmental damages in different parts of the ing mycorrhizal symbioses. In most mycorrhizal world (Binggeli 1996). The species that cause the symbioses, the fungal partner supplies nutrients greatest problems are generally those planted to the host plant in exchange for photosynthetic most widely and for the longest time (Pryor carbon, and may offer protection against patho- 1991; Rejma´nek and Richardson 1996). Accord- gens, toxins and drought (Smith and Read 1997). ing to Higgins and Richardson (1998), at least 19 The main types of mycorrhizal symbioses differ Pinus species are invaders of natural ecosystems in the anatomy of the mycorrhiza, which is a in the southern hemisphere; four of the most mixed root-fungus structure at which nutrient widespread invasive pines are P. halepensis, P. pa- interchanges take place (Smith and Read 1997). tula, P. pinaster and P. radiata (Higgins and Most Mediterranean plants are mycorrhizal with Richardson 1998; Richardson et al. 1990). different types of fungi. Herbs and shrubs mainly ThegenusEucalyptusL’He´rit(Myrtaceae)com- associate with glomales to form arbuscular prises evergreen woody plants, including shrubs mycorrhizas (AM), which is by far the most com- and forest trees (nearly 600 species), which are mon mycorrhizal symbiosis in the Mediterranean confinedinnaturaloccurrenceentirelytotheAus- terrestrial ecosystems (Dı´ez 1998). A restricted tralasian region, Papua, New Guinea and Timor group of plants form particular types of mycor- (Pryor 1976). More than 43 species of eucalypt rhizas with a range of particular ascomycetes and trees are planted outside their natural geographic basidiomycetes, such as the Ericales and Orchi- distribution.Eucalyptsareplantedonalargescale dales. However, most forest trees form ectomy- to provide a short rotation crop yielding wood corrhizas (EM) with a polyphyletic group of and paper pulp for industrial use (Eldridge et al. basidiomycetes and ascomycetes (Smith and 1994). Although less invasive than pines, several Read 1997; Hibbett et al. 2000). species of eucalypts already caused problems as invaders in South Africa (Richardson 1998; Forest trees are obligate ectomycorrhizal plants Richardson et al. 2000a). Eucalypts are repre- sented on many weed lists from other parts of the Pines and eucalypts are obligate ectomycorrhizal world, including California (Warner 1999) and trees, which depend on these mutualistic symbio- peninsular Spain(Sanz-Elorzaet al.2001).Signifi- ses for nutrient uptake in natural conditions cantimpactsmightresultfromtheintroductionof (Smith and Read 1997). Ectomycorrhizal symbio- eucalypts into Spain and some transformation on ses are essential in the mobilization of nutrients in various ecosystem properties; specially changes in soil forests (Fahey 1992; Read and Pe´rez-Moreno grasslands and scrubland habitats. Ecology and 2003).Innaturalconditions,theseforesttreesrely environmental politics dictate the desirability of on the ectomycorrhizal fungi, which colonize the maintaining Mediterranean grasslands and scrub- root cortex and form a nutrient-gathering ‘organ’ lands due to their high diversity in endemic plants calledectomycorrhiza(SmithandRead1997).Ec- andbecausetheseecosystemsarethenaturalhabi- tomycorrhizas are almost exclusive to forest trees. tats for many local wildlife. Eucalypt invasions In the Mediterranean, only a restricted group of can cause shifts in life-form dominance, reduced shrubs in the Cistaceae (Cistus spp., called rock- diversity, disruption of prevailing vegetation roses)formsEM,inwhichareinvolvedarangeof dynamics, and changing nutrient cycling patterns. endemic ascomycetes and basidiomycetes (Dı´ez Hence, eucalypt plantations are increasingly caus- 1998). Ectomycorrhizas are characterized struc- ing major conflicts betweenSpanish foresters, pol- turally by the presence of a dense mass of fungal iticiansandconservationists. mycelium surrounding the short lateral roots (the mantle). The mantle originates from the attach- ment of the fungal hyphae onto epidermal cells, Role of ectomycorrhizal fungi in exotic forestry and the multiplication of hyphae to form a series of hyphal layers. The fungal mycelium also grows Mycorrhizas of herbs, shrubs and forest trees among the cortical cells, forming the Hartig net. The Hartig net is the structural and functional Mutualistic interactions between fungi and plant interface between fungal and roots cells. The fun- roots are common in the plant kingdom, includ- gal mantle is connected with a highly extended
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