Issues in Bioinvasion Science

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.

An experimental work was conducted in Lleida (Spain) aiming to characterise the phenology and to quantify the demographic processes regulating the populations of Abutilon theophrasti Medicus in maize fields. Seedling emergence started a few days after crop sowing in early May and continued during two more months. The vegetative phase was very long due to the late seeding emergence; these later emerged plants showed a slower development, and many of them did not reach the fertility stage. A flowering peak was observed 12 weeks after emergence in late July, and fruit dehiscence and seed setting started in mid August, several weeks before crop harvest. Four different cohorts were identified, and two main peaks of emergence were determined 21 and 49 days after crop sowing nearest related with field irrigation. A functional logarithmic relationship between cumulative growing degree-days (GDD) and cumulative emergence was also described. The resulting demographic diagram reflects greater values relating to seedling survival for May cohorts (90.2 vs 7.9%), to fertility (100 vs 75%) and to fecundity (3774 vs 92 seeds pl⁻¹ ) than those determined for the June cohorts. The late emerged plants are subjected to a high density and are strongly affected by light competition, and their reproductive phase initiation delay is of about 10–20 days. In an assay conducted in Petri dishes, the seeds provided from plants emerged earlier were found more vigorous and germinated more than those from late emerged plants, which seem to be affected by incomplete fruit and seed ripening. Following the crop cycle without any weed control, the population rate increase was about 21.2. These values explain the high invasion capacity of this weed in the local summer irrigated fields, which consists in assuring their presence through a persistent soil seed bank and increasing the probability to spread to other fields.

Salinity is one of the main chemical factors in salt marshes. Studies focused on the analysis of salinity tolerance of salt marsh plants are very important, since they may help to relate their physiological tolerances with distribution limits in the field. Spartina densiflora is a South America cordgrass, which has started its invasion of the European coastline from the southwestern Iberian Peninsula. In this work, short-term responses in adult tussocks of S. densiflora from southwestern Spain are studied over a wide range of salinity in a greenhouse experiment. Our results point out that S. densiflora has a high tolerance to salinity, showing high growth and net photosynthesis rates from 0.5 to 20 ppt. S. densiflora showed at the lowest salinity (0.5 ppt) high levels of photoinhibition, compensated by higher levels of energy transmission between photosystems. Adaptative mechanisms, as those described previously, would allow it to live in fresh water environments. At the highest salinity (4 0 ppt), S. densiflora showed a high stress level, reflected in significant decreases in growth, net photosynthesis rate and photochemical e ciency of Photosystem II. These responses support S. densiflora invasion patterns in European estuaries, with low expansion rates along the coastline and faster colonization of brackish marshes and river banks.

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.

Based on a review and our own data, we present an overview of the ecological impacts on the trophic web of Mediterranean wetlands by an introduced Decapod Crustacean, the red swamp crayfish (Procambarus clarkii). P. clarkii lacks efficient dispersal mechanisms but is very well adapted to the ecological conditions of Mediterranean wetlands (fluctuating hydroperiods with regular intervals of drought). As an opportunistic, omnivorous species, which adapts its ecology and life history characteristics, such as timing and size at reproduction to changing environmental conditions, it became readily established in most of the Mediterranean wetland environments. High reproductive output, short development time and a flexible feeding strategy are responsible for its success as an invader. Like most crayfish, it occupies a keystone position in the trophic web of the invaded system and interacts strongly with various trophic levels. It e ciently grazes on macrophytes and is one of the main factors, besides the impact of flamingos, cattle and introduced fish, of the change of many water bodies from a macrophyte dominated, clear water equilibrium to a phytoplankton driven turbid water balance. Juveniles feed on protein rich animal food with the corresponding impact on the macroinvertebrate community in competition with other crayfish or fish species. At the same time, it serves as a prey for mammals, birds and fish. Due to its predatory and grazing activity, it efficiently canalises energy pathways reducing food web complexity and structure. Feeding also on detritus it opens, especially in marshlands, the detritic food chain to higher trophic levels which results in an increase of crayfish predators. As a vector of diseases, it has a severe impact on the preservation and reintroduction of native crayfish. P. clarkii accumulates heavy metals and other pollutants in its organs and body tissues and transmits them to higher trophic levels. Due to the long history of its presence, the complex interactions it established within the invaded ecosystems and the socio-economic benefits it provides to humans, prevention and control seem the most promising management measures to reduce the negative impact of this crayfish species.

An Asiatic mosquito species, Aedes albopictus, began to spread worldwide in the 1970s thanks to marine transport of tires and other goods, leading to colonization of many areas of the world. This species is a vector of major human diseases such as Dengue, Yellow Fever and the West Nile virus. In Europe, it was established in Albania and Italy and has been detected in other countries such as France; no records exist for Spain as yet. Colonization by Aedes albopictus is a major public health concern considering that the West Nile virus and several other viruses are known to circulate sporadically in the Mediterranean. Additionally, the parent species Aedes aegypti was the vector causing severe outbreaks of Dengue and Yellow Fever two centuries ago. Although Ae. aegypti was also introduced, it was eradicated from Spain. Both mosquitoes shared habitat types, diseases transmitted and many bionomic data. This article contains a review of the present Ae. albopictus distribution range worldwide and discusses the likelihood of an establishment in Spain in view of climatological and geographical data.

Comparing available paleontological, archaeological, historical, and former distributional data with current natural history and distributions demonstrated a turnover in the French vertebrate fauna during the Holocene (subdivided into seven sub-periods). To this end, a network of 53 specialists gleaned information from more than 1300 documents, the majority never cited before in the academic literature. The designation of 699 species as native, vanished, or non-indigenous in France or in one or more of its biogeographical entities during the Holocene period was investigated. Among these 699 species, 585 were found to belong to one or more of these categories. Among the 154 species that fit the definition of non- indigenous, 86 species were new species for France during the Holocene. Fifty-one that were autochthonous vanished from France during this period. Among these 51 species, 10 (two birds and eight mammals) are now globally extinct. During the last 11 millennia, the turnover in the French vertebrate fauna yielded a net gain of 35 species. On a taxon-by-taxon basis, there was a gain in the sizes of the ichthyofauna (19 : 27%), the avifauna (10 : 3%) and the herpetofauna (7 : 9%) and a loss in the mammalian fauna (−1 : 1%). Values of a per-century invasion index were less than 1 between 9200 BC and 1600 AD but increased dramatically after this date. An exponential model fits the trajectory of this index well, reaching the value of 132 invasions per century for the last sub- period, which encompasses 1945–2002. Currently, the local ecological and economic impacts of populations of 116 species (75% of the 154 that satisfied the criteria for non-indigenous) are undocumented, and the non-indigenous populations of 107 vertebrate species (69%) are unmanaged. The delay in assessing the ecological and economical impact of non-indigenous species, which is related to a lack of interest of French academic scientists in the Science and Action programmes, prevents the public from becoming informed and hinders the debates needed to construct a global strategy. For such a strategy to be effective, it will have to be elaborated at a more global scale than in just France — definitely at least in Europe.

We compared the life-history traits of native and invasive fish species from Catalan streams in order to identify the characters of successful invasive fish species. Most of the exotic fish species were characterized by large size, long longevity, late maturity, high fecundity, few spawnings per year, and short reproductive span, whereas Iberian native species exhibited predominantly the opposite suite of traits. Species native to the southeastern Pyrenees watershed were also significantly different from species native to the rest of the Iberian Peninsula but not native to this watershed. Iberian exotic species come predominantly from large river basins, whereas Catalan streams (and other small, coastal river basins) correspond to basins and streams of a smaller size and different hydrology, with differences in species composition and life-history traits of fish. The occurrence and spread of invasive species was not significantly related to life-history traits but to introduction date. The successful predi ction of future invasive species is limited due to small differences in life-history and ecological traits between native and exotic species. Fecundity, age at maturity, water quality flexibility, tolerance to pollution and habitat seem the most discriminating life-history variables.

Eradication of alien species is a key conservation tool to mitigate the impacts caused by biologic invasions. The aim of the present paper is to review the eradications successfully completed in Europe and to discuss the main limits to a wider application of this management option in the region. On the basis of the available literature — including conference proceedings, national reports to the Bern Convention, etc. — a total of 37 eradication programmes have been recorded. Thirty-three eradications were carried out on islands and four on the mainland. The rat (Rattus spp.) has been the most common target (n = 25, 67%), followed by the rabbit (n = 4). In many cases, these eradications determined a significant recovery of native biodiversity. Differently to other regions of the world, no eradications of alien inverte-brates and marine organisms have been recorded; regarding invasive alien plants, it appears that only some very localized removals have been completed so far in Europe. The limited number of eradications carried out in Europe so far is probably due to the limited awareness of the public and the decision makers, the inadequacy of the legal framework, and the scarcity of resources. Synthetic guidelines for improving the ability of European states to respond to aliens incursions are presented.

Many European politicians, managers, and scientists believe that non-indigenous species cannot be eradicated and that attempts to do so are hazardous because of frequent undesirable results. This notion seems to be based on the view that successful eradications undertaken in many other parts of the world cannot be generalised. To allow reasoned consideration of this argument, the eradication of non-indigenous vertebrate species performed in the French territories (European and overseas) and their recorded consequences on native fauna and flora are synthesised. Nineteen vertebrate eradication attempts were recorded, with seven mammal species as the targets. Of these attempts four failed for technical reasons and one for reasons undetermined as yet. These operations took place on islands of four biogeographical areas (West-European, Mediterranean, West Indies and Indian Ocean subantarctic) except a continental one (West-European continent). Among these 19 attempts, 13 were conducted according to a global strategy that provided data on the impact of the disappearance of the non-indigenous species on several native species. This impact, never detrimental, was determined for 14 species (one mammal, nine birds, one marine turtle, one crab, one beetle, one plant). Unexpected consequences of the disappearance of the invader were recorded for four native species (29%). This result highlights the poverty of natural historical information for several taxa and the flimsiness of the empty niche concept that is often used to argue for the delay of or to prevent any action again a non-indigenous species. If French territories can be taken as an example, eradications of non-indigenous species are not impossible; a good risk assessment prevents undesirable long-term consequences for native species and several native species benefited from the disappearance of the invader. Furthermore, eradication constitutes a powerful experimental tool for ecology and natural history studies if conceived as both a management and research operation.