Invasive blackberry outcompetes the endemic Galapagos tree daisy Scalesia pedunculata.

Island florae are under threat from habitat loss and competition with introduced species worldwide. In the Galapagos Islands, the endemic tree daisy Scalesia pedunculata (Asteraceae) is the dominant tree in the cloud forest of Santa Cruz Island but suffers from competition with the invasive blackberry Rubus niveus. At the site Los Gemelos, a S. pedunculata population was monitored from 2014 to 2021 following mechanical and chemical removal of R. niveus from 17 plots and compared with 17 additional plots where R. niveus remained. The aim of this study was to evaluate the impacts of the R. niveus invasion on S. pedunculata by characterizing the effects of R. niveus removal. Parameters measured in S. pedunculata were diameter at breast height (DBH, from which annual growth rates were determined), total height, survival of individual plants, and recruitment. In the presence of R. niveus, S. pedunculata trees had smaller DBH stems and shorter asymptotic maximum heights, growth rates declined for thin trees, the mortality of larger trees was elevated, and S. pedunculata recruitment was absent. R. niveus removal resulted in DBH-ratios of S. pedunculata more frequently meeting our threshold for fast growth (1.2), trees growing significantly thicker and taller, annual mortality being lower (12.5% vs. 16.2% per year), and recruitment being successful. In the presence of R. niveus, lower survival, growth, and absent recruitment suggested that S. pedunculata could reach quasi-extinction in ~20 years. Swift and decisive management action is needed to prevent the Scalesia forest on Santa Cruz Island from disappearing in less than two decades.

Global rise in emerging alien species results from increased accessibility of new source pools.

Our ability to predict the identity of future invasive alien species is largely based upon knowledge of prior invasion history. Emerging alien species-those never encountered as aliens before-therefore pose a significant challenge to biosecurity interventions worldwide. Understanding their temporal trends, origins, and the drivers of their spread is pivotal to improving prevention and risk assessment tools. Here, we use a database of 45,984 first records of 16,019 established alien species to investigate the temporal dynamics of occurrences of emerging alien species worldwide. Even after many centuries of invasions the rate of emergence of new alien species is still high: One-quarter of first records during 2000-2005 were of species that had not been previously recorded anywhere as alien, though with large variation across taxa. Model results show that the high proportion of emerging alien species cannot be solely explained by increases in well-known drivers such as the amount of imported commodities from historically important source regions. Instead, these dynamics reflect the incorporation of new regions into the pool of potential alien species, likely as a consequence of expanding trade networks and environmental change. This process compensates for the depletion of the historically important source species pool through successive invasions. We estimate that 1-16% of all species on Earth, depending on the taxonomic group, qualify as potential alien species. These results suggest that there remains a high proportion of emerging alien species we have yet to encounter, with future impacts that are difficult to predict.

Globally Distributed Arbuscular Mycorrhizal Fungi Associated With Invasive Cinchona pubescens on Santa Cruz Island, Galápagos.

The presence of arbuscular mycorrhizal fungi (AMF) is essential for the survival and establishment of most plant species in nature. The reproductive success of invasive plant species in a particular habitat could also depend on these AM fungi. Cinchona pubescens, commonly known as quinine, is highly invasive on Santa Cruz Island, Galápagos, but at the same time severely endangered in its native range on mainland Ecuador due to overexploitation in the past. In this study, we aim at determining the AMF communities associated with C. pubescens at both locations to investigate whether the successful invasion of C. pubescens on Santa Cruz is related to its association with a particular community of AMF. For this, roots of C. pubescens trees were sampled at three sites, one site on Santa Cruz and two sites in the province of Loja, on mainland Ecuador. Communities of AM fungi were determined through the molecular cloning and sequencing of the 18S nrDNA gene and through the delimitation of Operational Taxonomic Units (OTUs), associated with the plant roots. We found 36 AMF OTUs associated with C. pubescens, most of them belonging to the genus Glomus. The highest richness of AMF OTUs was detected in samples from sites located on mainland Ecuador. The AMF communities between Santa Cruz and mainland Ecuador were significantly different, and only five OTUs were shared between both locations. Two dominant OTUs in C. pubescens from Santa Cruz were detected but no dominant OTUs from mainland Ecuador. Almost two thirds of the OTUs associated with C. pubescens had a wide global distribution. Our results suggest that the successful invasion of C. pubescens on Santa Cruz could have been facilitated by local generalist AMF and not by particular AMF. The observed generalist AMF from both locations could be important for conservation plans of restoring the endangered C. pubescens in the native forests on mainland Ecuador.

Plant invasion causes alterations in Darwin's finch feeding patterns in Galápagos cloud forests.

Invasive species pose a major threat to forest biodiversity, particularly on islands such as the Galapágos. Here, invasive plants are threatening the remnants of the unique cloud forest and its iconic Darwin's finches. We posit that food web disturbances caused by invasive Rubus niveus (blackberry), have contributed to the rapid decline of the insectivourous green warbler finch (Certhidae olivacea). We compared the birds' dietary changes in long-term management, short-term management and unmanaged areas. We measured C:N ratios, and δ15N­nitrogen and δ13C­carbon values in both consumer tissues (bird-blood) and food sources (arthropods), as indicators of resource use change, and collected mass abundance, and arthropod diversity data. We characterised the birds' diets using isotope mixing models. The results revealed that finches in (blackberry-invaded) unmanaged areas foraged more on abundant, yet lower quality, arthropods present in the invaded understory. This suggests that blackberry encroachment leads to a decrease in food source quality with physiological consequences for green warbler finch chicks. Results also implied that blackberry control has a short-term impact on food source quantity, which led to a decrease in chick recruitment that we observed in our previous studies; despite this, in the long-term, these managed systems show signs of recovery within three years of restoration.

Limited natural regeneration of unique Scalesia forest following invasive plant removal in Galapagos.

More than 60% of the flora of the Galapagos Islands is introduced and some of these species have become invasive, severely altering ecosystems. An example of an affected ecosystem is the Scalesia forest, originally dominated by the endemic giant daisy tree Scalesia pedunculata (Asteraceae). The remnant patches of this unique forest are increasingly being invaded by introduced plants, mainly by Rubus niveus (blackberry, Rosaceae). To help large-scale restoration of this ecologically important forest, we seek to better understand the natural regeneration of S. pedunculata after invasive plant control. We monitored naturally recruited S. pedunculata saplings and young trees over five years in an area where invasive plant species are continuously being removed by manual means. We measured survival, height and growth of S. pedunculata saplings and young trees along permanent transects. Percent cover of surrounding plant species and of canopy shade directly above each S. pedunculata individual were determined, as well as distance to the next mature S. pedunculata tree. We identified potential factors influencing initial sapling survival and growth by applying generalized linear models. Results showed a rapid growth of saplings and young trees of up to 0.45 cm per day and a high mortality rate, as is typical for pioneer species like S. pedunculata. Sapling survival, growth and mortality seemed to be influenced by light availability, surrounding vegetation and distance to the next adult S. pedunculata tree. We concluded that natural regeneration of S. pedunculata was high only five months after the last herbicide application but that 95% of these recruits had died over the 5-year period. Further studies are needed to corroborate whether the number of surviving trees is sufficient to replace the aging adult trees and this way maintain remnants of the Scalesia forest. Urgent action is needed to help improve future restoration strategies to prevent further degradation of this rapidly shrinking threatened forest ecosystem.

The impact of invasive plant management on the foraging ecology of the Warbler Finch (Certhidea olivacea) and the Small Tree Finch (Camarhynchus parvulus) on Galápagos.

In recent decades, arboreal Darwin's Finches have suffered from a dramatic population decline, which has been attributed to parasitism by the invasive botfly Philornis downsi. However, changes to their primary habitat caused by invasive plant species may have additionally contributed to the observed population decline. The humid cloud forest on Santa Cruz Island is a stronghold of arboreal Darwin's Finches but has been invaded by blackberry (Rubus niveus). In some areas, manual control and herbicide application are used to combat this invasion, both causing a temporary removal of the entire understory. We hypothesized that the removal of the understory reduces the availability of arthropods, which are a main food source during chick rearing. We compared the foraging behaviour of Warbler Finches (Certhidea olivacea) and Small Tree Finches (Camarhynchus parvulus) at three study sites that varied in the degree of R. niveus invasion and the length of time since the last herbicide application. We used prey attack rate and foraging success as an index for food availability and predicted a lower attack rate and foraging success in areas that had recently been sprayed with herbicides. We found that both the invasion and the management of R. niveus influenced microhabitat use, foraging substrate and prey choice in both species. Contrary to our hypothesis, we did not find a lower attack rate or foraging success in the area with recent herbicide application. This may be explained by the finding that both species mainly foraged in the canopy but also used dead plant structures of the understory of the recently controlled area that resulted from the invasive plant management.

Alien species pathways to the Galapagos Islands, Ecuador.

Alien species, one of the biggest threats to natural ecosystems worldwide, are of particular concern for oceanic archipelagos such as Galápagos. To enable more effective management of alien species, we reviewed, collated and analysed all available records of alien species for Galápagos. We also assembled a comprehensive dataset on pathways to and among the Galápagos Islands, including tourist and resident numbers, tourist vessels, their itineraries and visitation sites, aircraft capacity and occupancy, air and sea cargo and biosecurity interceptions. So far, 1,579 alien terrestrial and marine species have been introduced to Galápagos by humans. Of these, 1,476 have become established. Almost half of these were intentional introductions, mostly of plants. Most unintentional introductions arrived on plants and plant associated material, followed by transport vehicles, and commodities (in particular fruit and vegetables). The number, frequency and geographic origin of pathways for the arrival and dispersal of alien species to and within Galápagos have increased over time, tracking closely the increase in human population (residents and tourists) on the islands. Intentional introductions of alien species should decline as biosecurity is strengthened but there is a danger that unintentional introductions will increase further as tourism on Galápagos expands. This unique world heritage site will only retain its biodiversity values if the pathways for invasion are managed effectively.

No saturation in the accumulation of alien species worldwide.

Although research on human-mediated exchanges of species has substantially intensified during the last centuries, we know surprisingly little about temporal dynamics of alien species accumulations across regions and taxa. Using a novel database of 45,813 first records of 16,926 established alien species, we show that the annual rate of first records worldwide has increased during the last 200 years, with 37% of all first records reported most recently (1970-2014). Inter-continental and inter-taxonomic variation can be largely attributed to the diaspora of European settlers in the nineteenth century and to the acceleration in trade in the twentieth century. For all taxonomic groups, the increase in numbers of alien species does not show any sign of saturation and most taxa even show increases in the rate of first records over time. This highlights that past efforts to mitigate invasions have not been effective enough to keep up with increasing globalization.