Corrigendum: Michel Pascal, Olivier Lorvelec & Jean-Denis Vigne, Holocene turnover of the French vertebrate fauna. Biological Invasions (2005) 7: 99–106

Corrigendum

Corrigendum: Michel Pascal, Olivier Lorvelec &; Jean-Denis Vigne, Holocene turnover of the French vertebrate fauna. Biological Invasions (2005) 7: 99–106     

French attempts to eradicate non-indigenous mammals and their consequences for native biota

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. This revised version was published online in July 2006 with corrections to the Cover Date.     

Patterns of Biological Invasions in French Freshwater Systems by Non-Indigenous Macroinvertebrates

Freshwater biodiversity is threatened by several mechanisms, of which the introduction of non-indigenous species and habitat alteration are the two most important. Exotic species act at various levels of organisation of macroinvertebrate communities, and are involved in different processes mediating their impacts on biodiversity, such as habitat modification or negative interactions with autochthonous fauna. The present work gives a list of the 43 French freshwater non-indigenous species, which represent 1.2% of the French freshwater macroinvertebrates. We provide their geographic origins, their distributions among zoological units by comparison with the native fauna and their functional characteristics according to a recent typology based on bio/ecological traits. An exponential trend of the cumulated number of non-indigenous species was evidenced, with a clumping of invaders within crustaceans and molluscs. Donor areas of non-indigenous species are in majority European, and the Ponto-Caspian basin is identified as the principal one. This pattern could be explained by a spread along waterways but its origin lies in a process of recolonisation of defaunated areas following several episodes of glaciation/deglaciation in Western Europe during the last 80,000 years. Finally, from a functional point of view, non-indigenous species exhibit a limited diversity, with two functional groups representing 80% of them.     

Effects of anthropogenic habitat disturbance on local pollinator diversity and species turnover across a precipitation gradient

Anthropogenic habitat disturbance can have profound effects on multiple components of forest biotas including pollinator assemblages. We assessed the effect of small-scale disturbance on local richness, abundance, diversity and evenness of insect pollinator fauna; and how habitat disturbance affected species turnover across the landscape and overall diversity along a precipitation gradient in NW Patagonia (Argentina). We evaluated the effect of disturbance on overall pollinator fauna and then separately for bees (i.e. Apoidea) and non-bee pollinators. Locally, disturbed habitats had significantly higher pollinator species richness and abundances than undisturbed habitats for the whole pollinator assemblage, but not for bees or non-bees separately. However, significant differences in species richness between habitats vanished after accounting for differences in abundance between habitat types. At a local scale Shannon–Weaver diversity and evenness did not vary with disturbance. A β diversity index indicated that, across forest types, species turnover was lower between disturbed habitats than between undisturbed habitats. In addition, rarefaction curves showed that disturbed habitats as a whole accumulated fewer species than undisturbed habitats at equivalent sample sizes. We concluded that small patches of disturbed habitat have a negligible effect on local pollinator diversity; however, habitat disturbance reduced β diversity through a homogenization of the pollinator fauna (in particular of bees) across the landscape.     

Early Holocene chronology and environment of Ampasambazimba,A Malagasy subfossil lemur site

According to a traditional but hitherto untested reconstruction of Madagascar’s Holocene environment, continuous forest preceded the monotonous grassland formations that now cover most of the island’s interior. Preliminary analyses of pollen samples collected near¹⁴C-dated horizons at Ampasambazimba (central Madagascar) indicate that a mosaic of woodlands, bushlands, and savanna existed close to this important vertebrate subfossil site around 7000–8000 BP. Although most members of Madagascar’s recently extinct “subfossil” fauna are thought to have been forest dwellers, several may have preferred a more open habitat like the one inferred for the region of Ampasambazimba. Dry savanna-woodland burns more readily than does dense forest and may have been more severely affected by the forces which transformed the Malagasy environment during the later Holocene. If so, the loss of savanna-woodland, not dense forest, may have been an important factor in the subfossil extinctions.     

Fish community comparisons along environmental gradients in lakes of France and north-east USA

Aim  To assess whether eight traits of fish communities (species richness, three reproductive traits and four trophic traits) respond similarly to environmental gradients, and consequently display convergence between the lakes of France and north-east USA (NEUSA).
Location  75 French and 168 north-east USA lakes.
Methods  The data encompass fish surveys, the assignment of species into reproductive and trophic guilds, and environmental variables characterizing the lakes and their catchments. The analytical procedure was adapted from the recommendations of Schluter (1986 ) [ Ecology , 67 , 1073–1085].
Results  The comparison of the regional pools of lacustrine fishes indicated that NEUSA was about twice as speciose as France, mostly due to higher species turnover across lakes, although NEUSA lakes were consistently about 20% more speciose than French lakes for a given surface area. Warmer environments were consistently inhabited by a higher proportion of phytophilous and guarder species than were colder lakes. Hence there was convergence in community reproductive traits. Conversely, there was no evidence of convergence in the trophic structure of lacustrine fish communities between regions.
Main conclusions  The influence of temperature on the availability and quality of spawning substrates appears to be a major constraint on present-day lacustrine fish communities. In parallel, phylogenetic constraints, past events such as the diversification of the North American fish fauna, and selective extinctions during Pleistocene glaciations and subsequent recolonizations contribute to explaining the dissimilarities between the communities of the two regions and differences in their relationship to the environment.     

Biological Invasions in Austria: Patterns and Case Studies

This paper provides a review of the first national inventory of non-indigenous species in Austria. In summary, 1110 vascular plant species (27 of the entire flora), 83 mycetes and at least 500 animal species (approximately 1 of the entire fauna) were documented for Austria, which are introduced intentionally or unintentionally by humans after 1492 and reported from the wild. About 25 of non-indigenous vascular plant species have become naturalized. Most non-indigenous vascular plants are native to the Palaearctic region (55%; with 33% originating from the Mediterranean subregion) and North America (20%). More than 90% of non-indigenous plant species are confined to naturally and anthropogenically disturbed (ruderal, urban, arable land, and riverine) habitats. Aquatic ecosystems are more affected and vulnerable to changes in their animal species composition. The current data demonstrate that non-indigenous species continue to invade and disperse and it also emphasize the necessity and responsibility to develop scientific strategies to minimize the impact of biological invasions and to raise public awareness of the problem.     

Obligate groundwater fauna of France: diversity patterns and conservation implications

We examined taxonomic and geographic patterns of the obligate groundwater fauna (i.e. stygobiotic fauna) by assembling in a distributional data base all species occurrences reported from France since 1805. A simulated annealing algorithm was used to identify conservation targets. Until the 60s, biological surveys were restricted to caves but the proportion of sampling sites in unconsolidated sediments increased from 1 to 16% over the last 40 years. A total of 380 species and subspecies in 40 families were collected, 70% of which being restricted to France. As observed in other temperate regions, the stygobiotic fauna was dominated by crustaceans (65% of species) and molluscs (22%). The cumulative number of species did not level off over time, clearly showing that biodiversity was underestimated. Temporal trends in the cumulative number of obligate groundwater and surface water species suggested that groundwater comprised more crustaceans than surface freshwater. Endemism was high although the geographic range size of species increased as distributional data accumulated. Of 380 species, 156 were known from a single 400-km² cell, among which 73% were located in the southern third of France. The distribution map of species richness changed dramatically over time, indicating that the location of richness hotspots was sensitive to sampling effort. Less than 2% of the French landscape was needed to capture 60% of known species. Thus, a large proportion of species could be protected by focusing habitat conservation efforts on a few complementary species-rich aquifers located in distinct regions.     

Late quaternary turnover of mammals in Borneo: the zooarchaeological record

The Quaternary has been a period of repeated, oscillating patterns of climate change. Global fluctuations in sea level affected the island status of Borneo, which was probably joined to continental Asia for more than half of the last 250,000 years. Alternating connection and isolation, coupled with the ecological barrier of a savanna corridor running from the Malay Peninsula to Java during periods of marine recession, are reflected in the present mammal fauna of Borneo. 38% of mammal species (excluding bats) are endemic, and some distinctive species or subspecies are confined to the north of the island. No known sites in Borneo match the Early and Middle Pleistocene regional sources in eastern Java. However, caves at Niah, Sireh and Jambusan, Sarawak, and Madai, Sabah, provide a zooarchaeological record covering the past 50,000 years. The Late Pleistocene mammals of Borneo included ten species also present among a Javan Middle Pleistocene savanna-adapted assemblage. Of these, four are categorised as ‘megafuana’: a giant pangolin, Javan rhinoceros, Malay tapir and tiger; the Sumatran rhinoceros can be added. In addition, there are less secure Pleistocene records of Asian elephant from Sarawak and Brunei. Holocene canid remains from Madai could either be the dhole or an early domestic dog. Palynological data combined with the mammal fauna confirm that around 45,000 years ago the vicinity of Niah was vegetated by closed forest. The continuous presence of a suite of arboreal specialists, including large primates, indicates that forest cover persisted through the terminal Pleistocene. Among local extinctions, the giant pangolin apparently disappeared early in this period, but tiger, Javan rhinoceros and tapir probably survived into the last millennium. Human predation of juveniles may account for the loss of the large ungulates, but the disappearance of tiger needs another explanation. Despite hunting pressure throughout the terminal Pleistocene and Holocene, a population of orangutan survived at Niah until perhaps the last millennium. Size diminution observed among large, medium and small mammal species is interpreted as the selective impact of environmental change. Once more is known about their ecology, changes in the bat fauna of Niah cave may provide indicators of environmental impacts affecting the wider mammal community during the later Holocene. In conclusion, it is recommended that the three nations, Brunei Darussalam, Malaysia and Indonesia, should support the WWF sponsored ‘Heart of Borneo’ as the most hopeful project to provide sustainable management of the rare and threatened forest-adapted wild mammals of the island.     

Animal xenodiversity in Italian inland waters: distribution, modes of arrival, and pathways

The paper provides a list of the non-indigenous animal species occurring today in Italian inland waters. Xenodiversity was found to amount to 112 species (64 invertebrates and 48 vertebrates), which contribute for about 2% to the inland-water fauna in Italy. Northern and central regions are most affected, and Asia, North America, and the rest of Europe are the main donor continents. The large majority of non-indigenous species entered Italy as a direct or indirect effect of human intervention. A difference between invertebrates and vertebrates was found for their mode of arrival (unintentional for invertebrates and intentional for vertebrates). Accidental transport, in association with both fish (for aquaculture or stock enhancement) and crops, has been the main vector of invertebrate introductions, whereas vertebrates were mostly released for stocking purposes. Overall stock enhancement (47.92%) and culture (37.5%) prevailed over the other pathways. Seventeen and 7 species of our list are included among the 100 worst invasive species of Europe (DAISIE) and of the world (IUCN), respectively. For some (but not all) non-indigenous species recorded in Italy the multilevel impact exerted on the recipient communities and ecosystems is known, even if rarely quantified, but knowledge on their chronic impact is still missing. Additional research is needed to provide criteria for prioritizing intervention against well established invaders and identify which new potential invader should be targeted as “unwanted”.     

Invasion success of non-indigenous aquatic and semi-aquatic plants in their native and introduced ranges. A comparison between their invasiveness in North America and in France

Aquatic and semi-aquatic plants comprise few species worldwide, yet the introduction of non-indigenous plants represents one of the most severe examples of biological invasions. My goal is to compare the distribution and the biology of aquatic and semi-aquatic plants in their introduced ranges and in their native ranges. The primary objective of this study is to test the hypothesis that invasive species have evolved traits likely to increase their success in the new range. I made two reciprocal comparisons, i.e. I compared European species in France and in North America, and North American species in France and in North America. Twenty-seven species were classified according to their invasiveness in their introduced area. I␣found six invasive macrophyte species in France native to North America and 17 invasive species in North America native to Europe. Four species are invasive in both areas. There is no general tendency for macrophytes to be more vigorous in their introduced ranges. Most non-indigenous aquatic and semi-aquatic species are potentially invasive or widespread and well-established in their introduced country, while few species seem to be restricted in their distribution.     

Basic features of the nemoral curculionid-beetle fauna (Coleoptera,Curculionoidea) from eastern Europe and western Siberia

There are 132 curculionids associated with deciduous trees in eastern Europe and western Siberia. The weevils and bark beetles make up the basis of the fauna, forming almost 80%. The fauna of oak is the richest (64 species). The species composition on the ash-tree (50% are monophages) is highly original. Impoverishment of the species composition from the west to the east is revealed. The fauna declines by 18% from 45 to 55° E. The species composition reduces two times beyond the Urals. The faunal boundaries are revealed from west to east and from east to west. The most significant boundary (47% of the fauna do not pass it) for distribution of species to the east is situated between 40 and 45° E. Two significant faunal boundaries are revealed from south to north. The fauna of curculionids of deciduous trees is formed by four groups of species. The southern European species (54%) and western palaearctic species (40%) dominate. The species composition of the studied territory is similar to the fauna of Middle Europe. Formation of the modern fauna of curculionids of deciduous trees in eastern Europe probably occurred in the Holocene. The western Palaearctic and Transpalaearctic species could have occupied the considered territory since the Late Pleistocene. Southern European species appeared in the considered territory after the Valdai glaciation during the warming in the Atlantic period of the Holocene. Penetration of East Palaearctic species began rather recently (last few decades), probably due to climate warming and economic activities.     

Changes in functional biodiversity in an invaded freshwater ecosystem: the Moselle River

The spread of non-indigenous species and the decline of autochthonous ones are leading to a homogenization of freshwater fauna in terms of systematic units, but the functional consequences are poorly documented. We studied the peculiar case of the lower, French section of the Moselle River where 20 invertebrate species have been introduced since 1854, with a rate increasing exponentially with time. Dredge sampling performed in 1994, 1996, 2000 and 2001 at four sampling stations allowed for an evaluation of faunal changes in terms of composition, structure and function. During this period, no structural changes were recorded in spite of multiple, new and successful introductions. The evaluation of functional modifications was based on a typology of taxa exhibiting homogeneous biological/ecological traits. Functional diversity, measured as the diversity of taxa distribution among functional groupings, revealed a significant increase between 1994/1996 and 2000/2001 because those species that were over-represented during the former period reached more equilibrated densities during the latter. The major, indirect implications of these functional changes are discussed.     

Molecular phylogeny,taxonomy, and distribution of French Unio species (Bivalvia,Unionidae)

A plethora of unionid names was established in the nineteenth century by the “Nouvelle Ecole”. Although naiad morphological plasticity is well documented, the currently recognized fauna, with 17 species and subspecies included in the French checklist for the Unio genus, is still based upon morphological characters only. Insights have been provided from molecular data elsewhere in Europe and North Africa, but the French fauna remains unstudied. We present a molecular phylogeny of the Unio genus in France based on COI, 16S and 28S genes; taking up all available data in Europe plus 273 specimens collected in all main French drainages. The results show that there are either three valid species in France, with U. pictorum and U. mancus synonymized, or five, with the subspecies U. crassus courtillieri elevated to species level. Subspecies were generally not recovered, which questions the evolutionary units tacitly implied by subspecific names. Although sampling topotypes is the most reliable way to evaluate the status of a nominal subspecies, major human-induced changes in aquatic hydrosystems challenge the method. Nevertheless, operational taxonomy has to rely on ground-truthed data and we propose to reduce the actual number of valid taxa in France to the seven observed operational taxonomic units.     

When east met west: the sub-fossil footprints of the west European hedgehog and the northern white-breasted hedgehog during the Late Quaternary in Europe

The distinct distribution of the west European hedgehog Erinaceus europaeus and the northern white-breasted hedgehog Erinaceus roumanicus and their separate refugial origins after the Pleistocene is a well-known example in the zoogeography of the Holarctic. Among the Late Quaternary faunal assemblages, the west European hedgehog is recorded at 269 sites whereas the northern white-breasted hedgehog is recorded only at 52 sites in Europe. The distribution patterns of the temporal and spatial Glacial records of the west European hedgehog show a general trend: a strong restriction to glacial refugia (the Iberian and Italian Peninsulas) during the Weichselian Glacial until the end of the Last Glacial Maximum, and a colonization of southern France during the early Late Glacial between 14 000 and 125 00 ¹⁴C years BP (15 000–12 800 cal. BC). Whereas the British Isles could have already been colonised by the end of the Pre-Boreal, in the rest of Central Europe E. europaeus was clearly distributed there in the Boreal for the first time. The west European hedgehog is an absolute Holocene faunal element in Central Europe. It appears in most parts of Central Europe during the Early Holocene, when the west European hedgehog met its eastern relative, which probably was similarly sensitive. After meeting each other, the distribution limit of both Erinaceus species in Central Europe seems to have been relatively constant in its geographic extent. Because of the clear climatic correlation, E. europaeus should be considered as an indicator species for temperate climatic conditions of the Holocene fauna. This should be considered during the reconstruction of climatic conditions with the help of the analysis of quaternary faunal material.     

Non-native Species and Rates of Spread: Lessons from the Brackish Baltic Sea

The Baltic Sea, a semi-enclosed brackish water region, has been inoculated by non-indigenous species for centuries. Today, much of its biological diversity is of foreign origin (i.e. xenodiversity), intentionally or unintentionally moved by humans over ecological and geographical barriers. As many as 98 introduced species have been recorded in the Baltic Sea and Kattegat. The role and abundance of much of the unique native brackish water fauna of the Baltic Sea are threatened by these non-indigenous species. The rate of primary introductions into the Baltic has increased since the 1950s; the secondary rate of spread of non-indigenous species within the basin varies from 30–480 km/year. We review here the invasion histories of the brown alga Sargassum muticum (introduced in the early 1990s), the mud snail Potamopyrgus antipodarum (1887), the barnacle Balanus improvisus (1844), the polychaetes Marenzelleria viridis (1985) and Polydora redeki (1963), the cladoceran Cercopagis pengoi (1992) and the mysid shrimp Hemimysis anomala (1962). This revised version was published online in July 2006 with corrections to the Cover Date.     

Holocene mammal extinctions in the Carpathian Basin: a review

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Prevalence of the Crayfish Plague Pathogen Aphanomyces astaci in Populations of the Signal Crayfish Pacifastacus leniusculus in France: Evaluating the Threat to Native Crayfish

Aphanomyces astaci, the crayfish plague pathogen, first appeared in Europe in the mid-19^th century and is still responsible for mass mortalities of native European crayfish. The spread of this parasite across the continent is especially facilitated by invasive North American crayfish species that serve as its reservoir. In France, multiple cases of native crayfish mortalities have been suggested to be connected with the presence of the signal crayfish Pacifastacus leniusculus, which is highly abundant in the country. It shares similar habitats as the native white-clawed crayfish Austropotamobius pallipes and, when infected, the signal crayfish might therefore easily transmit the pathogen to the native species. We investigated the prevalence of A. astaci in French signal crayfish populations to evaluate the danger they represent to local populations of native crayfish. Over 500 individuals of Pacifastacus leniusculus from 45 French populations were analysed, plus several additional individuals of other non-indigenous crayfish species Orconectes limosus, O. immunis and Procambarus clarkii. Altogether, 20% of analysed signal crayfish tested positive for Aphanomyces astaci, and the pathogen was detected in more than half of the studied populations. Local prevalence varied significantly, ranging from 0% up to 80%, but wide confidence intervals suggest that the number of populations infected by A. astaci may be even higher than our results show. Analysis of several individuals of other introduced species revealed infections among two of these, O. immunis and P. clarkii. Our results confirm that the widespread signal crayfish serves as a key reservoir of Aphanomyces astaci in France and therefore represents a serious danger to native crayfish species, especially the white-clawed crayfish. The prevalence in other non-indigenous crayfish should also be investigated as they likely contribute to pathogen transmission in the country.     

The river Rhine: a global highway for dispersal of aquatic invasive species

The river Rhine is heavily influenced by human activities and suffers from a series of environmental constraints which hamper a complete recovery of biodiversity. These constraints comprise intensive navigation and habitat modification by hydraulic engineering. Improving water quality while these constraints remain in place has led to increased colonization by aquatic invasive species. This tendency has been accelerated by the construction of canals connecting river basins. Over the last two centuries, the total surface area of river catchments connected to the river Rhine via inland waterways has been increased by a factor 21.6. Six principal invasion corridors for aquatic species to the river Rhine are discerned. The extensive network of inland waterways has allowed macroinvertebrate species from different bio-geographical regions to mix, changing communities, affecting the food webs and forming new constraints on the recovery of the native biodiversity. From the eighteenth century onward, in the freshwater sections of the river Rhine, a total of 45 non-indigenous macroinvertebrate species have been recorded. The average number of invasions per decade shows a sharp increase from <1 to 13 species. Currently, the contribution of non-indigenous species to the total species richness of macroinvertebrates in the river Rhine is 11.3%. The Delta Rhine and Upper Rhine exhibit higher numbers of non-indigenous species than other river sections, because the sea ports in the Delta Rhine and the Main-Danube canal function as invasion gateways. Important donor areas are the Ponto-Caspian area and North America (44.4 and 26.7% of the non-indigenous macroinvertebrate species, respectively). Transport via shipping and dispersal via man made waterways are the most important dispersal vectors. Intentional and unintentional introductions are highest for the period 1950–1992. The cumulative number of non-indigenous species in time is significantly correlated with the increase in total surface area of other river catchments connected to the river Rhine by means of networks of canals. The species richness of non-indigenous macroinvertebrates is strongly dominated by crustaceans and molluscs. Invasive species often tolerate higher salt content, temperature, organic pollution and current flow than native species. Spatiotemporal analyses of distribution patterns reveal that average and maximum dispersal rates of six invasive species vary between 44–112 and 137–461 km year⁻¹, respectively. Species arriving in upstream sections first show a shorter time lag between colonisation of the Delta and Upper Rhine than species initially arriving in downstream areas. Temporal analyses of macroinvertebrate assemblages in the littoral zones indicate that native species are displaced by non-indigenous species. However, established non-indigenous species are also displaced by more recent mass invaders.