Fire indirectly benefits fitness in two invasive species

Ecosystems perturbed from their natural disturbance regimes are more vulnerable to establishment and dominance of exotic plant species. Restoration efforts that reintroduce fire have achieved mixed success in reducing the abundance of exotic plants. The responses of many native species to fire are well known; fire-adapted species respond directly (heat and smoke cue germination) and indirectly (post-fire environment benefits seedling survivorship and growth) to fire. However, the direct and indirect effects of fire are unknown for most exotic plant species. We tested the direct and indirect effects of fire on two exotic invaders of Asian origin, Ailanthus altissima and Lonicera maackii, in North American woodlands. To quantify the direct effects of fire, we compared germination rates of seeds exposed to varying levels of heat and smoke in a laboratory and placed at different soil depths during a prescribed fire in the field. We examined the indirect effects of fire by comparing seedling recruitment in burned and unburned woodland plots. Results indicate that neither A. altissima nor L. maackii have germination cues associated with fire. However, both species have greater seedling recruitment in burned as compared to unburned areas in the field. Although seeds of these invasive species are not specifically adapted to fire, they still benefit from post-fire environments and pose a challenge to restoration of fire-maintained ecosystems. Future studies using our approach will allow land managers to better predict how communities will respond to restoration efforts and to understand variability observed in past restoration projects.     

Abiotic barriers limit tree invasion but do not hamper native shrub recruitment in invaded stands

The interplay between the invasion of alien plant species and re-colonization of native plant species is important for conservation. Sandy coastal plains (called restinga in Brazil) were used as a model system to explore the abiotic barriers that potentially limit the initial establishment of alien and native woody plants in invaded and non-invaded areas. The study tested the influence of light availability, soil type and litter layer on recruitment of a wind-dispersed alien tree (Casuarina equisetifolia) and two bird-dispersed native shrubs under a Casuarina stand and in the preserved restinga. The effect of soil type and the physical and allelopathic effects of Casuarina litter on seedling emergence of the three species were also evaluated under greenhouse conditions. Low dispersal associated with low seedling emergence and zero survival of young plants (mainly due to microhabitat conditions) apparently prevents the spread of Casuarina in the preserved restinga. The main cause of low recruitment of native species in the Casuarina stand was the physical barrier of the litter. However, if seeds overcome this physical barrier, the presence of litter improves seedling emergence and survival of young plants, mainly because the litter increases soil moisture. Sowing seeds below litter and planting young plants of native shrubs on litter can improve the re-colonization of native plants in invaded areas. In conclusion, Casuarina invasion on sandy coastal plains is strongly limited by abiotic barriers, but anthropogenic disturbances are altering the key processes that naturally make the restinga resistant to invasion.     

Drivers of invasive tree and shrub natural regeneration in temperate forests

We assessed drivers of ecological success along resource availability gradients for three invasive woody species: Prunus serotina Ehrh., Quercus rubra L. and Robinia pseudoacacia L. We aimed to check how much of invasion success, measured by invader biomass, is explained by propagule pressure and plant community invasibility. Using 3 years of observations from 372 study plots (100 m² each) in temperate forests of Wielkopolski National Park (Poland) we investigated the hierarchy of predictors and partial dependencies using the random forest method. Our study indicated that propagule pressure explained more variance in success of invaders than invasibility—describing availability of resources and competitors in understory vegetation. We also found different responses of seedlings and saplings, connected with dependence on stored carbohydrates, which decreased seedling responses to resource availability gradients. However, resource availability (light and leaf litter predictors) had greater influence than predictors describing understory vegetation. Based on importance and response strength the species studied may be arranged by decreasing requirements for soil fertility and acidity: P. serotina?<?Q. rubra?<?R. pseudoacacia, whereas for light requirements and competition vulnerability the order is: P. serotina?>?Q. rubra?>?R. pseudoacacia. However, low light requirements of R. pseudoacacia may be biased by high proportion of sprouts supplied by parental trees. Results provide guidelines for effective management of invasive woody species in forest ecosystems and describe complex interactions between factors studied on ecological success of invaders.     

A decade of emerald ash borer effects on regional woodpecker and nuthatch populations

The emerald ash borer (EAB) Agrilus planipennis, first detected in 2002 in the vicinity of Detroit, Michigan, USA, has spread throughout much of eastern and midwestern North America as of 2016, resulting in widespread mortality of ash trees in the genus Fraxinus. We investigated the effects of this newly available, exotic food source on populations of six species of largely resident insectivorous birds, including five species of woodpeckers and white-breasted nuthatch (Sitta canadensis), using North American Breeding Bird Survey data for breeding season estimates and Audubon Christmas Bird Counts for winter season estimates. We found evidence for relatively modest and variable effects of EAB invasion on the populations of these birds during the breeding season, but highly significant numerical increases during the winter that in several cases appeared to be increasing as the EAB invasion has progressed. Our results confirm that the EAB invasion is resulting in increased populations of several insectivorous birds, primarily during the winter. They also suggest that the numerical response of woodpeckers to EAB may be such that avian predation, which represents a significant, and possibly the largest, morality factor affecting some EAB populations, may continue to increase and help control the EAB epidemic as the invasion continues.     

Distribution and abundance of a keystone tree, <Emphasis Type="Italic">Schinziophyton rautanenii</Emphasis>, and factors affecting its structure in Zambia,southern Africa

Schinziophyton rautanenii is a keystone tree species whose fruits are eaten by wildlife and livestock, nuts are used to extract oil for human use and its wood is used for making curios that are sold to tourists. The species occurs in southern African countries of Angola, Namibia, Botswana, Mozambique, Zambia, Zimbabwe and the Democratic Republic of Congo but little is known about its population and conservation status. The objectives of this study were to (i) assess factors determining the distribution and abundance of the species in Zambia, (ii) evaluate its regeneration potential and population status and (iii) assess whether the main determinants of forest and species stand structures also favour S. rautanenii. The study is based on three forest surveys conducted in 2004, 2005–2008 and 2015. In Zambia S. rautanenii abundance and population structure are controlled by soil, climate and human factors. The species is most abundant in southwest Zambia where population recruitment is good but is rare and recruitment is poor elsewhere in the country. Fire appears to be a major cause of tree damage and possibly death. The elephant is a major disperser of S. rautanenii nuts and the low recruitment levels in the country may partly be attributed to the decline in the elephant population from about 250,000 in 1960 to 28,000 in 2008. Different conservation strategies will be required for areas where the species is rare and for those where the species is abundant. Whatever the strategies, it will be important to address issues of fire management, fruit harvesting and the role of wildlife in the conservation of S. rautanenii in Zambia.     

Native ecosystem engineer facilitates recruitment of invasive crab and native invertebrates

Native ecosystem engineers that add physical structure to ecosystems can facilitate invasive species. In this study we determined the effects of the native tube-forming serpulid worm, Galeolaria caespitose on the recruitment of the invasive New Zealand porcelain crab, Petrolisthes elongatus, and whether invasive crab recruitment was related to the recruitment of native species. P. elongatus is abundant beneath intertidal rocks around Tasmania, southern Australia, and the underside of these rocks is usually covered with a calcareous matrix formed by the serpulid. We used an experimental approach to investigate whether rocks, serpulids on the underside of rocks and adult P. elongatus influenced the recruitment P. elongatus and native communities. P. elongatus and native invertebrates only recruited in the presence of rocks indicating the importance of rock as primary recruitment habitat. Moreover, the presence of serpulids on the underside of rocks significantly increased the recruitment of P. elongatus and native invertebrates compared to rocks without serpulids. Rocks with higher densities of adult P. elongatus at the end of the experiment also had higher densities of P. elongatus recruits. The density of P. elongatus recruits did not influence native species richness and abundance although there was some evidence that high P. elongatus recruitment was correlated with shifts in native community structure. We have shown that a native ecosystem engineer facilitates recruitment of an invasive crab but this does not appear to influence the recruitment of native species.     

Responses of native plant populations on an unprotected beach to disturbance by storm-induced overwash events

Extreme climatic events disturb plant communities, altering species composition, and abundance. For herbaceous populations on coastal beaches, responses to disturbance are often spatially heterogeneous and depend on species’ life history. This study documents the effects of two overwash events, caused by Hurricane Irene in August 2011 and Superstorm Sandy in October 2012, on the abundance of native species on a low-lying, unprotected beach on Staten Island, New York, USA. Temporary plots (33 cm diameter) spaced at 3-m intervals along 30-m transects perpendicular to shoreline, were used in September 2007, 2011, 2012, and 2013 to survey lower (0–9 m), middle (12–21 m), and upper (24–30 m) regions of the same beach. The same procedure was used to quantify seedling recruitment in spring (May) after each overwash. Complex changes in density the year after each overwash were specific for distinct beach regions and depended on species’ life history. Rhizomatous perennial grasses (Ammophila breviligulata and Panicum amarum) maintained or increased ramet populations, possibly stimulated by sand deposition on the middle and upper beach. The first overwash was early enough in the growing season to permit subsequent seed production during autumn, and seedling recruitment occurred for three native annuals (Cenchrus tribuloides, Heterotheca subaxillaris, and Triplasis purpurea), and the tall perennial Solidago sempervirens the next spring. The second overwash was later in autumn and probably removed seeds from the site and/or deeply buried the seeds of two annuals (Cenchrus and Heterotheca) which did not recruit seedlings the next spring. Triplasis seedlings were abundant in a blowout on the lower beach. Given more frequent storms and sea-level rise in the future, it becomes imperative to maintain an assemblage of native plant species with diverse life histories when restoring coastal beaches.     

Allelopathic Potential of <Emphasis Type="Italic">Koelreuteria bipinnata</Emphasis> var. <Emphasis Type="Italic">integrifoliola</Emphasis> on Germination of Three Turf Grasses

Allelopathy is very important for the scientific disposition of garden plants. To understand the allelopathic potential of Koelreuteria bipinnata Franch. var. integrifoliola, the germination of Agrostis tenuis Sibth., Festuca arundinacea Schreb. and Lolium perenne L. were determined under laboratory conditions. The results showed that root, stem and leaf aqueous extracts of K. bipinnata var. integrifoliola had allelopathic effects on all three turf grasses, and the allelopathic activity varied according to extract concentrations, test species, and extract sources. Lower extract concentrations did not affect or promoted the germination and initial seedling growth of turf grasses, but the highest concentrations almost had inhibitory effect. The order of allelopathic potentials of the three organs on germination of these receptors was root < stem < leaf. And at the highest concentration of leaf extract, the most strongly inhibition was found in A. tenuis, followed by F. arundinaces and then L. perenne. In addition, according to gas chromatography–mass spectrometry (GC–MS) analysis, the allelopathic potential compounds and their abundance in root, stem and leaf were obviously different. Therefore, the allelopathic compounds may responsible for allelopathy of K. bipinnata var. integrifoliola. These findings suggested that more attention should be paid to the leaf of K. bipinnata var. integrifoliola for the relative higher allelopathic effects.     

Diversity and abundance of coccinellids (Coleoptera: Coccinellidae) on trees in parks and tree rows of Belgrade

Harmonia axyridis is an invasive coccinellid species that was first recorded in Serbia in 2008. In order to establish whether after its arrival it became the dominant coccinellid species, we investigated the diversity and abundance of coccinelids on trees in parks and tree rows of Belgrade during 2015 and 2016. Thirteen coccinellid species were recorded in these investigations. Frequently occurring among them were Adalia bipunctata, A. decempunctata, Coccinella septempunctata, Exochomus quadripustulatus, Harmonia axyridis, H. quadripunctata, Oenopia conglobata, and Popylea quatuordecimpunctata. All of them had different dynamics of appearance during the vegetation. As a result of this, the values of their dominance in the coccinellid community constantly changed. However, only in the cases of H. axyridis, O. conglobata, and C. septempunctata were those values high enough for us to be able to say that they were dominant in terms of abundance during a certain period. Although the amounts of summer precipitation in Serbia were very different during 2015 and 2016, the trends of changes in the diversity of coccinellids throughout the growing season in these two years were similar.     

Factors influencing the dispersal of a native parasitoid, <Emphasis Type="Italic">Phasgonophora sulcata</Emphasis>, attacking the emerald ash borer: implications for biological control

High parasitism by a native parasitoid, Phasgonophora sulcata Westwood (Hymenoptera: Chalcididae), has been reported on emerald ash borer (hereafter EAB), Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), in North America. Use of this parasitoid in an augmentative biological control program has been proposed to slow the spread of EAB, yet information is lacking on key aspects of this parasitoid’s dispersal. We document the flight capacity and walking activity of P. sulcata, its potential fecundity, and describe how age, body size, temperature, and time of day affect these parameters. Wasp flight capacity, measured using flight mills, increased with temperature and decreased with age. Unexpectedly, age and body size did not affect wasp walking activity, and we saw no relationship between walking activity and flight capacity. Older wasps had lower potential fecundity than younger wasps. These results suggest that P. sulcata should be released as pupae near EAB-infested ash trees to improve efficacy and potential biological control success.     

Sexual reproduction of clonal dwarf shrubs in a forest–tundra ecotone

Successful sexual reproduction may be more important for regeneration of clonal species in high-latitude and -altitude areas than has been previously suggested. We investigated the potential of Vaccinium myrtillus, V. vitis-idaea and Empetrum nigrum ssp. hermaphroditum (E. hermaphroditum) for sexual reproduction at three sites in the forest–tundra ecotone in Finnish Lapland. We studied whether the potential differs between plant communities, whether disturbance enhances germination, and whether seedling emergence is limited by seed availability. We established a field experiment with disturbance and sowing treatments, and monitored seed and seedling numbers and survival rates for two years. The number of mature seeds of V. myrtillus was higher in plants from the tundra heath than in those from the coniferous and mountain birch forests. The number of mature seeds and seedlings emerging from the seed bank of E. hermaphroditum tended also to be higher in the tundra heath. Disturbance marginally increased the seedling emergence of V. myrtillus and E. hermaphroditum, whereas sowing generally increased the seedling numbers. The seedling number of V. myrtillus was lower in the tundra heath and that of E. hermaphroditum was lower in the coniferous forest than at the other sites. Seedling survival was equal for all plant species at all sites. We conclude that the capacity for sexual reproduction varies among plant communities, and seed availability is a stronger constraint than microsite availability for the studied species. Once the crucial early phase of seedling establishment is overcome, seedling survival enables successful recruitment of V. myrtillus, V. vitis-idaea and E. hermaphroditum in the subarctic area.     

Size matters: predation of fish eggs and larvae by native and invasive amphipods

Invasive predators can have dramatic impacts on invaded communities. Extreme declines in macroinvertebrate populations often follow killer shrimp (Dikerogammarus villosus) invasions. There are concerns over similar impacts on fish through predation of eggs and larvae, but these remain poorly quantified. We compare the predatory impact of invasive and native amphipods (D. villosus and Gammarus pulex) on fish eggs and larvae (ghost carp Cyprinus carpio and brown trout Salmo trutta) in the laboratory. We use size-matched amphipods, as well as larger D. villosus reflecting natural sizes. We quantify functional responses, and electivity amongst eggs or larvae and alternative food items (invertebrate, plant and decaying leaf). D. villosus, especially large individuals, were more likely than G. pulex to kill trout larvae. However, the magnitude of predation was low (seldom more than one larva killed over 48 h). Trout eggs were very rarely killed. In contrast, carp eggs and larvae were readily killed and consumed by all amphipod groups. Large D. villosus had maximum feeding rates 1.6–2.0 times higher than the smaller amphipods, whose functional responses did not differ. In electivity experiments with carp eggs, large D. villosus consumed the most eggs and the most food in total. However, in experiments with larvae, consumption did not differ between amphipod groups. Overall, our data suggest D. villosus will have a greater predatory impact on fish populations than G. pulex, primarily due to its larger size. Higher invader abundance could amplify this difference. The additional predatory pressure could reduce recruitment into fish populations.     

Spatial patterns of shrub encroachment in neighbouring grassland communities in the Pyrenees: floristic composition heterogeneity drives shrub proliferation rates

We studied the role of floristic composition and associational resistance in shrub dynamics by comparing spatial patterns of shrub cover after prescribed burning in neighbouring grassland communities with different palatability. The study focused on the shrub Cytisus balansae ssp. europaeus (G. López and Jarvis) Muñoz Garmendia. Seven two-dimensional transects (20 × 0.5 m) were established to monitor shrub cover for at least 10 years after prescribed burning. Shrub cover and spatial patterns were assessed in each transect. Floristic similarity between transects and Cytisus associations with different species were estimated. Over an entire transect, shrub cover and shrub scale of pattern and patch size were lowest in the unpalatable Festuca eskia grasslands and highest in F. paniculata grasslands. At short distances, we found negative associations between Cytisus and most of the grasses, except for F. nigrescens and Agrostis capillaris, which showed positive associations with Cytisus. Thus, the effects of associational resistance on shrub encroachment were not as marked as expected, F. eskia grasslands showing the lowest shrub encroachment rates after fire. By contrast, Cytisus was positively associated with the most palatable grasses in the site, namely F. nigrescens and A. capillaris. We conclude that differences in floristic composition drive shrub encroachment rates in these spatially heterogeneous communities.     

Contrasting impacts of highly invasive plant species on flower-visiting insect communities

Invasive alien plants threaten biodiversity, ecosystems and service provision worldwide. They can have positive and negative direct and indirect effects on herbivorous insects, including those that provide pollination services. Here, we quantify how three highly invasive plant species (Heracleum mantegazzianum, Impatiens glandulifera and Fallopia japonica) influence the availability of floral resources and flower-visiting insect communities. We compared invaded with comparable uninvaded areas to assess floral resources and used pan-trapping to quantify insect communities. Only F. japonica influenced floral resource availability: sites invaded by this species had a higher flowering plant species richness and abundance of open floral units than uninvaded sites, probably due to its late flowering and the paucity of other flowering species at this time of year. Fallopia japonica was also associated with higher abundances of bumblebees, higher overall insect diversity and higher hoverfly diversity than uninvaded areas. Differences in pollinator communities were also associated with I. glandulifera and H. mantegazzianum, despite there being no detectable differences in floral resources at these sites. Specifically, there were more bumblebees and solitary bees in I. glandulifera sites, and a higher overall diversity of insects, particularly hoverflies. By contrast, H. mantegazzianum sites had a lower abundance of solitary bees and hoverflies. These findings confirm that invasive plant species have a range of species-specific effects on ecological communities. This supports the emerging view that control of invasive species, as required under international obligations, is not simple and that potential losses and gains for biodiversity must be carefully evaluated on a case-by-case basis.     

Competitive effects of the invasive shrub,Lonicera maackii (Rupr.) Herder (Caprifoliaceae), on the growth and survival of native tree seedlings

Invasive plants are often associated with reduced cover of native plants, but rarely has competition between invasives and natives been assessed experimentally. The shrub Lonicera maackii, native to northeastern Asia, has invaded forests and old fields in numerous parts of eastern North America, and is associated with reduced tree seedling density in Ohio forests. A field experiment was conducted to test the effects of established L. maackii on the survival and growth of transplanted native tree species. The experiment examined above-ground competition (by removing L. maackii shoots) and below-ground competition (by trenching around transplanted seedlings). The effects of above-ground competition with L. maackii were generally more important than below-ground competition, though both were detected. Shoot treatment was the key determinant for the survival of all species except P. serotina, whereas trenching only enhanced survival for A. saccharum caged and P. serotina, and only in the shoot removal treatment. For the surviving seedlings, L. maackii shoot removal increased growth of A. saccharum seedlings protected with cages, but actually reduced the growth of unprotected Q. rubra and A. saccharum seedlings, indicating that L. maackii shoots confer some protection from deer browsing. Significant interactions between root and shoot treatment on Q. rubra growth parameters, specifically greatest growth in the shoot present & trenched treatment, is attributed to protection from deer browsing combined with release from below-ground competition. Despite this protective function of L. maackii shoots, the overall effect of this invasive shrub is increased mortality of native tree seedlings, suggesting it impacts the natural regeneration of secondary forests.     

Interaction between invasive and potentially invasive shrub species does not influence relationships between their ecological success and distance from propagule sources

Although interactions between alien and native plant species are well studied, data on interactions between two co-existing alien species with respect to their invasibility are scarce. The aims of this study were (1) to evaluate three factors shaping abundance of the alien shrub species Cornus alternifolia: abundance of another alien (invasive) shrub species (Prunus serotina), type of tree stand (coniferous vs. broadleaved) and distance to propagule sources and (2) to assess the potential dispersal distance of the species studied. Densities of both species were assessed within 194 experimental plots (located in experimental plantations of trees) in Rogów Arboretum (Central Poland). P. serotina occurred on 79 and C. alternifolia on 33 of the 194 plots. The furthest distance of C. alternifolia from the propagule source was 338 m. C. alternifolia reached higher densities in coniferous than broadleaved tree stands. Density of C. alternifolia depended on tree stand type and distance from the propagule source, but did not depend on density of P. serotina. Density of C. alternifolia decreased with increasing distance from the propagule source; however, this relationship was modified by the type of tree stand: densities were lower in broadleaved than in coniferous stands. The presence of the invasive species seems to neither facilitate nor limit the dispersal distance of C. alternifolia, as these two species differ in shade tolerance. The study also provided the first information about C. alternifolia potential invasiveness, because earlier this species was noticed only as casually escaping from cultivation in Slovakia.     

The effects of hydramethylnon on the tropical fire ant, <Emphasis Type="Italic">Solenopsis geminata</Emphasis> (Hymenoptera: Formicidae), and non-target arthropods on Spit Island,Midway Atoll,Hawaii

Invasive ants can cause major disruptions in native ecosystems. Ant eradication methods without significant non-target effects are needed to stop incipient invasions and to aid in ecosystem restoration. Successful ant eradications are rare and there is very little understanding of the effects of ant eradication methods, such as the use of formicides, on non-target species. Here we attempted to control and possibly eradicate the invasive tropical fire ant, Solenopsis geminata, from a small islet using the formicide Maxforce^® (active ingredient: hydramethylnon), and to quantify the non-target effects on an almost exclusively alien ground-dwelling arthropod community. S. geminata abundance was reduced and the species was not detected on bait cards for 12 months post-treatment. The abundance of another non-target invasive ant that was primarily detected in pitfall traps, Tetramoruim bicarinatum, declined in pitfall traps following treatment, but seemed to be excluded from bait cards by S. geminata. Total ant abundance did not return to original levels until more than 12 months post-treatment. Populations of alien cockroaches (Order Blattaria) and crickets (Orthoptera: Gryllidae) were negatively affected by the treatment. We conclude that Maxforce^® can be used to control small infestations of S. geminata and T. bicarinatum effectively; however we recommend it be used cautiously due to the potential ecological cost to non-target species. Use in areas where infestations are small and isolated will maximize the likelihood of success while minimizing non-target effects.