Biological invasions are a population-level rather than a species-level phenomenon.

Biological invasions pose a rapidly expanding threat to the persistence, functioning and service provisioning of ecosystems globally, and to socio-economic interests. The stages of successful invasions are driven by the same mechanism that underlies adaptive changes across species in general-via natural selection on intraspecific variation in traits that influence survival and reproductive performance (i.e., fitness). Surprisingly, however, the rapid progress in the field of invasion science has resulted in a predominance of species-level approaches (such as deny lists), often irrespective of natural selection theory, local adaptation and other population-level processes that govern successful invasions. To address these issues, we analyse non-native species dynamics at the population level by employing a database of European freshwater macroinvertebrate time series, to investigate spreading speed, abundance dynamics and impact assessments among populations. Our findings reveal substantial variability in spreading speed and abundance trends within and between macroinvertebrate species across biogeographic regions, indicating that levels of invasiveness and impact differ markedly. Discrepancies and inconsistencies among species-level risk screenings and real population-level data were also identified, highlighting the inherent challenges in accurately assessing population-level effects through species-level assessments. In recognition of the importance of population-level assessments, we urge a shift in invasive species management frameworks, which should account for the dynamics of different populations and their environmental context. Adopting an adaptive, region-specific and population-focused approach is imperative, considering the diverse ecological contexts and varying degrees of susceptibility. Such an approach could improve and refine risk assessments while promoting mechanistic understandings of risks and impacts, thereby enabling the development of more effective conservation and management strategies.

Taming the terminological tempest in invasion science.

Standardised terminology in science is important for clarity of interpretation and communication. In invasion science - a dynamic and rapidly evolving discipline - the proliferation of technical terminology has lacked a standardised framework for its development. The result is a convoluted and inconsistent usage of terminology, with various discrepancies in descriptions of damage and interventions. A standardised framework is therefore needed for a clear, universally applicable, and consistent terminology to promote more effective communication across researchers, stakeholders, and policymakers. Inconsistencies in terminology stem from the exponential increase in scientific publications on the patterns and processes of biological invasions authored by experts from various disciplines and countries since the 1990s, as well as publications by legislators and policymakers focusing on practical applications, regulations, and management of resources. Aligning and standardising terminology across stakeholders remains a challenge in invasion science. Here, we review and evaluate the multiple terms used in invasion science (e.g. 'non-native', 'alien', 'invasive' or 'invader', 'exotic', 'non-indigenous', 'naturalised', 'pest') to propose a more simplified and standardised terminology. The streamlined framework we propose and translate into 28 other languages is based on the terms (i) 'non-native', denoting species transported beyond their natural biogeographic range, (ii) 'established non-native', i.e. those non-native species that have established self-sustaining populations in their new location(s) in the wild, and (iii) 'invasive non-native' - populations of established non-native species that have recently spread or are spreading rapidly in their invaded range actively or passively with or without human mediation. We also highlight the importance of conceptualising 'spread' for classifying invasiveness and 'impact' for management. Finally, we propose a protocol for classifying populations based on (i) dispersal mechanism, (ii) species origin, (iii) population status, and (iv) impact. Collectively and without introducing new terminology, the framework that we present aims to facilitate effective communication and collaboration in invasion science and management of non-native species.

What's the effectiveness of stocking actions in small creeks? The role of water discharge behind hatchery trout downstream movement.

Fish stocking to enhance freshwater fisheries or to improve the conservation status of endangered fish species is a common practice in many countries. Little is known, however, of the effectiveness of these practices in spite of the high efforts and investments required. The movement of subadult/adult hatchery-released brown trout Salmo trutta L. was studied by passive telemetry in a small tributary of Lake Lugano (i.e., Laveggio Creek, Canton Ticino, Switzerland). Hatchery fish, together with some resident wild individuals sampled during electrofishing surveys, were tagged with Passive Integrated Transponders (PIT) tags. Hatchery fish were released upstream and downstream a submersible monitoring antenna, which was anchored to the streambed in a pass-over orientation. The number of hatchery fish detected daily by the antenna (divided between fish released upstream and downstream the antenna) was analyzed in relation to the daily water discharge, to search for similar patterns in their fluctuation over time. Only the movement of fish released upstream the antenna displayed a significant relationship with water discharge, with the highest number of fish detected during periods of high-water flow, occurring after heavy rains. High-water discharge events had a significant role in hatchery trout downstream movement in our study site, likely acting as a driver for the downstream migration to Lake Lugano. Such events contributed to the poor effectiveness of stocking actions in this small tributary, providing further evidence against stocking strategies based on subadult/adult fish.

Biological and trophic consequences of genetic introgression between endemic and invasive Barbus fishes.

Genetic introgression with native species is recognized as a detrimental impact resulting from biological invasions involving taxonomically similar invaders. Whilst the underlying genetic mechanisms are increasingly understood, the ecological consequences of introgression are relatively less studied, despite their utility for increasing knowledge on how invasion impacts can manifest. Here, the ecological consequences of genetic introgression from an invasive congener were tested using the endemic barbel populations of central Italy, where the invader was the European barbel Barbus barbus. Four populations of native Barbus species (B. plebejus and B. tyberinus) were studied: two purebred and two completely introgressed with alien B. barbus. Across the four populations, differences in their biological traits (growth, body condition and population demographic structure) and trophic ecology (gut content analysis and stable isotope analysis) were tested. While all populations had similar body condition and were dominated by fish up to 2 years of age, the introgressed fish had substantially greater lengths at the same age, with maximum lengths 410-460 mm in hybrids versus 340-360 mm in native purebred barbel. The population characterized by the highest number of introgressed B. barbus alleles (81 %) had the largest trophic niche and a substantially lower trophic position than the other populations through its exploitation of a wider range of resources (e.g. small fishes and plants). These results attest that the genetic introgression of an invasive congener with native species can result in substantial ecological consequences, including the potential for cascading effects. SUPPLEMENTARY INFORMATION: The online version contains supplementary material available at 10.1007/s10530-021-02577-6.

Cryptic diversity within endemic Italian barbels: revalidation and description of new Barbus species (Teleostei: Cyprinidae).

Two fluviolacustrine lineages (SI1 Barbus and SI2 Barbus) of the Barbus genus have been recently detected in the Apulia-Campania ichthyogeographic district (southern Italy). The aim of this study was to determine the taxonomic status of these lineages by comparing them with the two already-established Italian fluviolacustrine species Barbus plebejus and Barbus tyberinus through a more in-depth molecular and morphological investigation. Genetic analyses were performed on both mitochondrial (cytochrome b) and nuclear (growth hormone paralog 2) DNA markers, and morphological analyses were carried out on specific age classes and purebred populations. Molecular analyses detected four evolutionary lineages at the mitochondrial level, whereas the nuclear data set highlighted the strict evolutionary relation between B. plebejus sensu stricto and the new lineages, converged in the B. plebejus complex clade. The morphological analyses allowed us to discriminate SI1 Barbus and SI2 Barbus from both B. plebejus and B. tyberinus. The new taxa could be discriminated by the greatest maximum body height and the longest pre-orbital distance, respectively. Both the new lineages have longer ventral and pectoral fins than B. plebejus and B. tyberinus, a larger caudal fin than B. tyberinus and a lower number of scales along the lateral line than B. plebejus. Both molecular and morphological results suggested the two southern Italian lineages could be considered as distinct endemic species: the formal description of the new species Barbus samniticus sp. nov. (i.e., SI1 Barbus) and the revalidation of Barbus fucini Costa, 1853 (i.e., SI2 Barbus) were thus proposed, and, for both species, molecular and morphological diagnosis were provided.

Genetic and morphological analyses reveal a complex biogeographic pattern in the endemic barbel populations of the southern Italian peninsula.

The Italian peninsula is a biodiversity hotspot, with its freshwater fish fauna characterized by high levels of local endemism. Two endemic fluvio-lacustrine fishes of the genus Barbus (barbel, family Cyprinidae) have allopatric distributions in the Tyrrhenian and Adriatic basins of Italy. Barbus plebejus inhabits the mid- to northern Adriatic basins, while B. tyberinus is widespread in all central-northern basins draining into the Tyrrhenian Sea. For basins in Southern Italy draining into the southern parts of these seas, there remains a knowledge gap on their barbel populations due to no previous genetic and morphological studies, despite their apparent biogeographic isolation. Correspondingly, this study quantified the presence and distribution of barbels in the Adriatic and Tyrrhenian basins of Southern Italy through genetic and morphological analyses of 197 fish sampled across eight populations. Testing of how local isolation has influenced the evolution and persistence of these populations was completed by examining sequence variation at two mitochondrial loci (cytochrome b and D-loop) and performing geometric morphometric analyses of body shape, plus measuring 11 morphometric and meristic characters. Phylogenetic and morphological analyses revealed the presence of two genetically distinct lineages that differed significantly from adjacent B. tyberinus and B. plebejus populations. These two new taxa, here described as SI1 and SI2 Barbus lineages, are highly structured and reflect a complex mosaic biogeographic pattern that is strongly associated with the underlying hydrographical scenarios of the basins. The geographic isolation of these basins thus has high evolutionary importance that has to be considered for maintaining endemism.

Predicting the ecological impacts of an alien invader: Experimental approaches reveal the trophic consequences of competition.

Ecological theory on the trophic impacts of invasive fauna on native competitors is equivocal. Whilst increased interspecific competition can result in coexisting species having constricted and diverged trophic niches, the competing species might instead increase their niche sizes to maintain energy intakes. Empirical experiments can test invasion theory on competitive interactions and niche sizes across different spatial scales and complexity. The consequences of increased interspecific competition from a model alien fish Leuciscus idus were tested on two taxonomically and trophically similar native fishes, Squalius cephalus and Barbus barbus. Competitive interactions were tested in tank aquaria using comparative functional responses (CFRs) and cohabitation trials. The consequences of these competitive interactions for the trophic niche sizes and positions of the fishes were tested in pond mesocosms. Comparative functional responses revealed that compared to B. barbus, L. idus had significantly higher attack and consumption rates; cohabitation trials revealed B. barbus growth rates were depressed in sympatry with L. idus. For L. idus and S. cephalus, differences in their functional response parameters and growth rates were not significant. Pond mesocosms used stable isotope metrics to quantify shifts in the trophic niche sizes of the fishes between allopatry and sympatry using a substitutive experimental design. Isotopic niches were smaller and more divergent in sympatric paired species than predicted by their allopatric treatments, suggesting trophic impacts from interspecific competition. However, an all-species sympatric treatment revealed similar niche sizes with allopatry. This maintenance of niche sizes in the presence of all species potentially resulted from the buffering of direct competitive effects of the species pairs by indirect effects. Experimental predictions from tank aquaria assisted the interpretation of the constricted and diverged trophic niches detected in the paired-species sympatric treatments of the pond mesocosms. However, the all-species sympatric treatment of this experiment revealed greater complexity in the outcomes of the competitive interactions within and between the species. These results have important implications for understanding how alien species integrate into food webs and influence the trophic relationships between native species.