Differences in arbuscular mycorrhizal fungal communities associated with sugar maple seedlings in and outside of invaded garlic mustard forest patches

Garlic mustard (Alliaria petiolata) is a Eurasian native that has become invasive in North America. The invasive success of A. petiolata has been partly attributed to its production of allelopathic compounds that can limit the growth of arbuscular mycorrhizal fungi (AMF). Although such effects are well known, specific effects on the richness and community composition of AMF associated with woody species have not been explored. We collected sugar maple (Acer saccharum) seedlings from eight natural forest sites in Ohio and Massachusetts, containing areas either invaded or uninvaded by A. petiolata. We measured AMF root colonization of seedlings, isolated DNA from the roots and performed PCR-TRFLP analysis to assess the richness and community composition of AMF. As expected, we found reduced AMF colonization in A. petiolata invaded patches. A. petiolata did not alter the detected TRF richness, but was associated with significant changes in the composition of AMF communities in half of the sites monitored in each region. Our results suggest that although AMF colonization was reduced in A. petiolata patches, many indigenous AMF communities include AMF that are tolerant to allelopathic effects of A. petiolata.     

An exotic invader drives the evolution of plant traits that determine mycorrhizal fungal diversity in a native competitor

The symbiosis between land plants and arbuscular mycorrhizal fungi (AMF) is one of the most widespread and ancient mutualisms on the planet. However, relatively little is known about the evolution of these symbiotic plant–fungal interactions in natural communities. In this study, we investigated the symbiotic AMF communities of populations of the native plant species Pilea pumila (Urticaceae) with varying histories of coexistence with a nonmycorrhizal invasive species, Alliaria petiolata (Brassicaceae), known to affect mycorrhizal communities. We found that native populations of P. pumila with a long history of coexistence with the invasive species developed more diverse symbiotic AMF communities. This effect was strongest when A. petiolata plants were actively growing with the natives, and in soils with the longest history of A. petiolata growth. These results suggest that despite the ancient and widespread nature of the plant–AMF symbiosis, the plant traits responsible for symbiotic preferences can, nevertheless, evolve rapidly in response to environmental changes.     

Observations of herbivore attack on garlic mustard (Alliaria petiolata) in Southwestern Ontario, Canada

We have observed what appears to be the first significant report of herbivory of a population of Alliaria petiolata (M. Bieb.) Cavara & Grande (Garlic Mustard) in Canada. Ceutorhynchus erysimi Fabr., Plutella xylostella L. (Diamondback Moth), and Philaenus spumarius (Meadow Spittlebug) were identified attacking, and successfully proceeding through different stages of development on A. petiolata. From the field observations, we hypothesize that these species have recognized A. petiolata as a host and therefore may be candidates for the management of the invasive species.     

Landscape scale assessment of stream channel and riparian habitat restoration needs

Human modifications of streams and rivers have caused extensive stream channel and riparian degradation. Cost-effective, rapid assessment tools can be used to better manage such areas by identifying the status of habitats for restoration planning and protection. We used a spatially explicit, reach-scale geographic information system modeling strategy to examine stream channel and riparian condition and prioritize restoration actions. The stream channel condition index uses information on land use, road and railroad density, and sinuosity. The riparian condition index uses calculations of percent forest, patch density, and convexity based on land cover in the floodplain. Reaches were classified into restoration categories based on stream channel and riparian condition model results, land ownership, slope, position in the subwatershed, and adjacency to high-quality habitat. We compared modeled restoration priority rankings with those in the management plan for the East Credit subwatershed in Ontario, Canada. Predicted stream channel restoration priority rankings matched field-based classifications for 86% of the reaches in the East Credit subwatershed. Predicted riparian restoration priority rankings matched field-based classifications for 81% of the reaches. Our methods replicate with fairly good accuracy the results obtained using intensive field surveys and stakeholder input. Managers can use these cost-effective strategy development tools to identify candidate reaches for further study and prioritize stream channel and riparian restoration actions over large regions.     

Pre-release monitoring of Alliaria petiolata (garlic mustard) invasions and the impacts of extant natural enemies in southern Michigan forests

Monitoring of populations of a target weed species prior to releasing natural enemies has the potential to improve the rigor and safety of biological control and to determine the invader’s impacts on native communities while creating a reference point for evaluating the efficacy of subsequent biocontrol agent releases. Eight populations of garlic mustard, Alliaria petiolata (M. Bieb) Cavara and Grande (Brassicaceae), an invasive weed in southern Michigan, were monitored in anticipation of releases of classical biological control agents. The A. petiolata populations were shown to be expanding with 44.4% of initially uninvaded sampling quadrats becoming invaded after four years. While 88.2% of quadrats with A. petiolata showed evidence of foliar damage from pathogens and browsing by mammals, insects and other invertebrates, levels of damage were low and had little impact on rosette or seedling survival. Contrary to expectations, damage was positively correlated with A. petiolata fecundity (P = 0.0465). Given the continued expansion of A. petiolata and the lack of significant herbivore damage by acquired natural enemies, a biological control program should be considered against this invasive plant. If biological control agents are released, the results of this study will provide a benchmark for evaluating their performance.     

Effects of Herbicide on the Invasive Biennial Alliaria petiolata (Garlic Mustard) and Initial Responses of Native Plants in a Southwestern Ohio Forest

Restoration often includes control of invasive plants, but little is known about how native plant communities respond to this control. The biennial Alliaria petiolata (M. Bieb.) Cavara and Grande (garlic mustard) is one of the most prevalent invasive plants in forests of eastern North America. We investigated the effects of the herbicide Round‐up (glyphosate) on Alliaria and the response of the forest floor plant community to the herbicide and the subsequent decline of Alliaria. In an old‐growth Acer–Fagus stand and a second‐growth Liriodendron‐dominated stand in Hueston Woods State Nature Preserve, Ohio, United States, we spot applied Round‐up in November 2000 and 2001 in 25 1 × 1–m plots and maintained 25 control plots. Herbicide decreased Alliaria density in both stands and reduced the density of other species in leaf during treatment (mostly exotic winter annuals) in the old‐growth stand. Treatment did not affect the initial density of the Alliaria cohort that germinated in the spring of 2001, but decreased the 2002 cohort. Community differences were found in the old‐growth stand after Alliaria reduction, specifically greater cover of spring ephemerals in the herbicide treatment. In the second‐growth stand, herbicide treatment increased reproduction of the late‐summer perennial, Phryma leptostachya. These results indicate that glyphosate reduces Alliaria without negatively impacting native species and that some native species respond positively to a single‐year reduction in this invasive biennial.     

Exotic Slugs Pose a Previously Unrecognized Threat to the Herbaceous Layer in a Midwestern Woodland

Developing effective restoration strategies requires first identifying the underlying factors limiting native plant recovery. The slug Deroceras reticulatum is an important herbivore in Europe, a global agricultural pest, and is introduced and abundant throughout eastern North America, but little information is available on the effect of this exotic herbivore on the forest herbaceous layer. Here, we test the palatability of 12 forest herbs to the introduced slug D. reticulatum and use field surveys to determine the degree to which slugs are damaging plants in the field. In laboratory feeding trials, slugs readily consumed most plants, but avoided the grass Elymus virginicus, the invasive forb Alliaria petiolata (garlic mustard), and thicker leaved plants. In the field, we documented significant slug damage, with close to 50% or more of plant leaves damaged by slugs on five of the six native species tested. Slug damage in the field was predicted by laboratory‐determined acceptability, but was significantly greater on short‐statured rosette species than on erect plants for a given acceptability value. Our results identify introduced slugs as an important, but overlooked obstacle to forest herb restoration and potential drivers of larger scale understory compositional change. The relaxed herbivore pressure on A. petiolata, relative to native competitors, suggests that invasive plant removal alone may not result in the recovery of native flora. Rather, restoration of unpalatable native species should accompany invasive plant control in slug invaded areas. Erect forbs, thick‐leaved plants, and graminoids should have the greatest success where introduced slugs are abundant.     

Distribution and spread of environmental weeds along New?Zealand roadsides

Most non-native weeds and other naturalised plants are in the early stages of invasion into New Zealand landscapes. For this invasion to be controlled, even partially, it is important to understand the dominant routes, mechanisms, and rates of weed spread across landscapes. With their linear corridors of disturbed habitats, roadsides are thought to play a large role in the spread of some weeds. We used both new surveys and existing data to assess which of the 328 environmental weeds listed by the Department of Conservation are most frequently found and where on roadsides, and whether distribution patterns are consistent with linear dispersal. We also analysed historical survey data for relationships between reserve weediness and proximity to roads. We surveyed 340 plots of 100-m-long stretches of roadside across four regions and found between 2 and 19 environmental weeds per plot; 128 species in total (Chao estimate 148). Especially abundant were agricultural (weeds and cultivated) species, species that have been naturalised for well over 50 years, and species that disperse externally attached to vertebrates. While we purposefully sampled within 10 km of town limits, we found no strong effect of distance from town on roadside weed richness, including richness of just ornamentally sourced weeds. Instead, number of houses within 250?m and presence of an adjacent house or other residential structure were both important, as was presence of woody vegetation on and adjacent to roadsides. Reserves adjacent to roads had significantly higher weed richness than reserves further from roads, although the causal mechanisms are unclear. Our results suggest that while roadsides include suitable habitats for most environmental weeds, distributions are patchy and roads show little sign of acting as linear dispersal corridors, instead they largely reflect neighbouring land uses. As such, roadside weeds should best be managed as part of the wider landscape.     

How novel are the chemical weapons of garlic mustard in North American forest understories?

The Novel Weapons Hypothesis predicts that invasive plants excel in their new ranges because they produce novel metabolites to which native species possess little resistance. We examined the novelty of the phytochemistry of the Eurasian invader, Alliaria petiolata, in North America by comparing its phytochemical profile with those of closely related Brassicaceae native to North America. We examined the profile and/or concentrations of glucosinolates, alliarinoside, flavonoids, cyanide, and trypsin inhibitors in cauline leaves of field-collected A. petiolata, Arabis laevigata, Cardamine concatenata, C. bulbosa, and C. douglassii. Cyanide and the glucosinolates and flavonoids produced by A. petiolata were detected only in A. petiolata. Trypsin inhibitor activity was highest in A. laevigata, intermediate in the Cardamine species, and lowest in A. petiolata. The phytochemical profile of A. petiolata was distinct from those of four closely related and/or abundant Brassicaceaeous species native to North America, providing support for the Novel Weapons Hypothesis.     

Concurrent Management of an Exotic Species and Initial Restoration Efforts in Forests

One of the proximate results of forest fragmentation, and a cause of continued microenvironmental change and exacerbation of ecological problems, is increased invasions by weedy plant species. One such example is Alliaria petiolata (Brassicaceae), a serious pest threatening much of eastern North America. Alliaria petiolata impedes mitigation of fragmentation and restoration efforts because it tends to outcompete and possibly extirpate much of the native understory species on localized scales. As part of a strategy to address the problems of fragmented habitats, an experiment was conducted to determine whether Sanguinaria canadensis (Papaveraceae) could outcompete A. petiolata. Using an additive design, I transplanted S. canadensis at densities of 0, 1, 2, 3, 5, 7, 9, 11, 15, and 20 ramets/m² in 1997 and allowed them to interact with initial A. petiolata densities of 128 seedlings and 31 rosettes/m². As of 2000, multivariate analyses of variance with repeated measures and simple analyses of variance indicated that initial S. canadensis densities of as little as 5 ramets/m² suppressed A. petiolata. Initial S. canadensis densities of 9 and 11 ramets/m² resulted in the lowest numbers of late‐spring seedlings, numbers and sizes of year 1 and 2 rosettes, numbers and gross areas of stem leaves, numbers of flowering individuals, number of flowers, number of fruits (siliques), and height at flowering. While it remains to be tested whether this will continue and if the reestablishment of S. canadensis will help reassemble forest ecosystems, the experiments indicated that transplanting S. canadensis was effective at mitigating the spread of A. petiolata.     

Does understory vegetation reflect the history of fluvial disturbance in a riparian forest?

We have examined the relationship between the history of fluvial disturbance and understory vegetation in a riparian forest. The study site was divided into three sites, by use of aerial photographs and topographical maps, with different histories of fluvial disturbance: (1) Fagus-type on land that has not been flooded for the last 39 years, at least; (2) Populus-type on land that has not been flooded since debris flow occurred 34 years ago; and (3) Salix-type on land that has been flooded periodically from an abandoned channel since debris flow occurred 34 years ago. Species richness in the Salix-type was significantly higher than in the other types. Detrended correspondence analysis revealed obvious floristic differences among the three canopy types. Canonical correspondence analysis showed that herbaceous species were mainly found on lower plots with high moss cover, implying that moss layers may capture seeds transported by the stream. Tall herbs occurred in less shaded plots and on higher plots, suggesting that their rapid growth prevented the occurrence of other species. Fagus-type was dominated by species with ingested fruits which depended on animals for their dispersal. Populus and Salix-types were dominated by species with wind dispersal or no dispersal mechanism, which depended on physical phenomena for dispersal. Attributes of current understory vegetation were connected with historical events, suggesting that riparian vegetation reflects the history of fluvial disturbance.     

Reduced competitive ability in an invasive plant

One explanation for successful plant invaders is that they evolved to be more competitive. An intuitive prediction of this Evolution of Increased Competitive Ability (EICA) hypothesis never previously tested is that invasive populations should outcompete their native ‘ancestors’ in a common environment. We tested this idea in a diallel competition experiment with Alliaria petiolata where offspring from native and invasive populations were grown alone or in all pairwise combinations. While without competition, there were no differences between native and invasive populations, native populations outperformed invasive ones when competing against each other. Our results contradict the EICA hypothesis and we conclude that it does not not hold for Alliaria petiolata. Instead, we formulate a new ERCA (Evolutionary Reduced Competitive Ability) hypothesis: if there is less competition in the invasive range and competitive ability involves traits that have a fitness cost, then selection might act against it, thereby reducing intraspecific interactions too.     

Estimating the abundance of Long‐billed Curlews in Nebraska

ABSTRACT Long‐billed Curlews (Numenius americanus) are shorebirds of high conservation concern in North America. Populations have declined in the last 150 yr primarily due to habitat loss and conversion. We conducted a 2‐yr study to estimate the density and statewide abundance of breeding Long‐billed Curlews in Nebraska during 2008 and 2009. Surveys were conducted during the prenesting period in April when Long‐billed Curlews were likely to be detected. We used a simple random sample of roadside survey routes (N= 39), each consisting of 40 5‐min point‐counts at 800‐m intervals. We modeled detection probability and found that wind speed negatively affected detectability, but found no evidence of either year effects or observer bias. We estimated there were 0.0038 Long‐billed Curlews per hectare (0.38 Long‐billed Curlews per km²) along survey routes and, by extrapolation, estimated there were 23,909 (SE = 1685; 95% CI: 20,810—27,471) Long‐billed Curlews in Nebraska. Our population estimate suggests that ～15 to 22% of the United States population of Long‐billed Curlews is found in Nebraska. Curlews were not evenly distributed within our survey area, with the highest densities in the central Sandhills, an area dominated by grass‐stabilized sand dunes and shallow wetlands, and the lowest densities in predominantly agricultural areas. Because Long‐billed Curlews in Nebraska face many potential threats resulting from land‐use changes, monitoring of the breeding population should be continued. Our survey method was efficient and yielded interpretable results; other states or regions should consider adopting this approach for estimating curlew abundance.     

Modelling heterogeneity in detection probabilities in land and aerial abundance surveys in humpback whales (<Emphasis Type="Italic">Megaptera novaeangliae</Emphasis>)

The effective management and conservation of animal populations relies on statistically-sound and replicable surveys to obtain estimates of abundance and assess trends. Surveys of cetaceans, such as humpback whales Megaptera novaeangliae, are difficult to conduct and are particularly affected by bias in detection probability. For example, the probability of detection of whales from land decreases substantially with increased distance from the platform. This distance effect is also true for aerial surveys, combined with the problem that animals are unavailable for detection (underwater) whilst in the field of view. We present a novel approach that combines corrected double-platform land surveys with corrected aerial surveys to obtain a robust estimate of g(0), the probability of detection on the survey line, for aerial surveys of migrating humpback whales. Several sources of heterogeneity in detection probabilities were identified within the land and aerial surveys (including group composition, bearing of first sighting, number of groups being tracked simultaneously and cloud cover). After including these into our estimate of ĝ(0), we found that only 29% of available whales are being detected on the survey line (ĝ(0) = 0.288), which is a considerably smaller estimate than many available for humpback whales using other methods. Incorporating heterogeneity into the population surveys shows that we are likely to be underestimating the population size of whales on the east coast of Australia. The implications of this result for their conservation and management in light of increased whale-human conflict is discussed.     

PCR–RFLP assays for discerning three weevil stem feeders (Ceutorhynchus spp.) (Col.: Curculionidae) on garlic mustard (Alliaria petiolata)

Polymerase chain reaction–restriction fragment length polymorphism (PCR–RFLP) assays were developed to rapidly and cost-effectively differentiate the morphologically indistinguishable larval stages of three species, Ceutorhynchus alliariae, C. roberti and C. scrobicollis, proposed for the biological control of Alliaria petiolata. A PCR–RFLP using TaqαI restriction enzyme can differentiate the three target species only, while a PCR–RFLP double digestion using the restriction enzymes AluI and TaqαI should enable, based on virtual digestion, distinguishing them from 12 other Ceutorhynchus species associated with A. petiolata.     

Intraspecific variation in allelochemistry determines an invasive species’ impact on soil microbial communities

Invasive species can benefit from altered species interactions in their new range, and by interfering with species interactions among native competitors. Since exotic invasions are generally studied at the species level, relatively little is known about intraspecific variation in the traits that determine an invader’s effect on native species. Alliaria petiolata is a widespread and aggressive invader of forest understories that succeeds in part by interfering with mutualistic interactions between native plants and soil fungi. Here, I show that the impact of A. petiolata on soil microbial communities varied among individuals due to variation in their allelochemical concentrations. The differential impacts translated into varied effects on native tree growth, partly because A. petiolata’s allelochemicals preferentially affected the most mutualistic fungal taxa. These results highlight the importance of considering the spatial and temporal variation in an invasive species’ impacts for understanding and managing the invasion process.     

Estimating abundance and spatial distribution patterns of the bubble crab Dotilla fenestrata (Crustacea: Brachyura)

Abstract The bubble crab Dotilla fenestrata forms very dense populations on the sand flats of the eastern coast of Inhaca Island, Mozambique, making it an interesting biological model to examine spatial distribution patterns and test the relative efficiency of common sampling methods. Due to its apparent ecological importance within the sandy intertidal community, understanding the factors ruling the dynamics of Dotilla populations is also a key issue. In this study, different techniques of estimating crab density are described, and the trends of spatial distribution of the different population categories are shown. The studied populations are arranged in discrete patches located at the well‐drained crests of nearly parallel mega sand ripples. For a given sample size, there was an obvious gain in precision by using a stratified random sampling technique, considering discrete patches as strata, compared to the simple random design. Density average and variance differed considerably among patches since juveniles and ovigerous females were found clumped, with higher densities at the lower and upper shore levels, respectively. Burrow counting was found to be an adequate method for large‐scale sampling, although consistently underestimating actual crab density by nearly half. Regression analyses suggested that crabs smaller than 2.9 mm carapace width tend to be undetected in visual burrow counts. A visual survey of sampling plots over several patches of a large Dotilla population showed that crab density varied in an interesting oscillating pattern, apparently following the topography of the sand flat. Patches extending to the lower shore contained higher densities than those mostly covering the higher shore. Within‐patch density variability also pointed to the same trend, but the density increment towards the lowest shore level varied greatly among the patches compared.     

Monitoring and assessing old‐growth forest stands by plot sampling

Abstract

Forest inventories are evolving towards multipurpose resource surveys, broadening their scope by including additional topics such as biodiversity issues. Surprisingly, few quantitative surveys have been devoted to old‐growth forests, even if they constitute the most acknowledged forest biodiversity icons. In this framework, the use of probabilistic sampling may provide an effective as well as rigorous support for monitoring and assessing old‐growth forests. To this purpose, the present paper proposes a two‐phase sampling scheme. In the first phase, a coarse survey of few floristic and stand structural attributes is carried out by means of small plots systematically placed on the study area. Subsequently, in the second phase, a fine assessment of a large number of ecological attributes is performed on a subset of enlarged plots selected among the first‐phase ones by means of simple random sampling without replacement. The proposed sampling scheme is implemented for monitoring and assessing the old forests of Cilento National Park (southern Italy). Results and comments are provided as an exemplicative case study.     

The effects of arbuscular mycorrhizal (AM) fungal and garlic mustard introductions on native AM fungal diversity

Introduced, non-native organisms are of global concern, because biological invasions can negatively affect local communities. Arbuscular mycorrhizal (AM) fungal communities have not been well studied in this context. AM fungi are abundant in most soils, forming symbiotic root-associations with many plant species. Commercial AM fungal inocula are increasingly spread worldwide, because of potentially beneficial effects on plant growth. In contrast, some invasive plant species, such as the non-mycorrhizal Alliaria petiolata, can negatively influence AM fungi. In a greenhouse study we examined changes in the structure of a local Canadian AM fungal community in response to inoculation by foreign AM fungi and the manipulated presence/absence of A. petiolata. We expected A. petiolata to have a stronger effect on the local AM fungal community than the addition of foreign AM fungal isolates. Molecular analyses indicated that inoculated foreign AM fungi successfully established and decreased molecular diversity of the local AM fungal community in host roots. A. petiolata did not affect molecular diversity, but reduced AM fungal growth in the greenhouse study and in a in vitro assay. Our findings suggest that both introduced plants and exotic AM fungi can have negative impacts on local AM fungi.     

Multi-scale species density model for conserving an endangered songbird

Habitat modeling across a landscape that has gradients of habitat conditions requires potential predictor data that can be quantified at biologically relevant scales. We used remotely sensed data to develop a multi-scale density model in 2018 for the golden-cheeked warbler (Setophaga chrysoparia; warbler), a species that breeds in Ashe juniper (Juniperus ashei)-oak (Quercus spp.) woodlands in central Texas, USA. We first classified Ashe juniper and broadleaf tree cover at a 1-m resolution and used this to map potential habitat across the warbler's >67,000-km² breeding range. We then designed a survey for estimating warbler density based on hierarchical distance sampling. We used stratified random sampling to survey for male warblers at 1,804 points across the continuum of tree canopy cover and composition and detected 810 warblers during our surveys. We developed a suite of potential predictor variables for modeling warbler density that reflected vegetation, topography, climate, and anthropogenic land use conditions across the breeding range and developed these at 3 scales representing the territory, site, and landscape. We modeled warbler density and used the best fit model to produce a spatially explicit estimate. Predicted warbler density was influenced by tree canopy cover and canopy height at the territory scale (100-m radius); tree canopy cover, percent of the canopy comprised of juniper, and an interaction between canopy cover and compound topographic index at the site scale (1-km radius); and annual temperature range at the landscape scale (5-km radius). We estimated a population size of 217,444 male warblers (95% CI = 153,917–311,965) and >3,000 males in each recovery unit. After controlling for the duration of point count surveys, our estimate of population size was similar to that reported from the only previous breeding range survey conducted in 2008–2009. Our model results indicated that management activities to increase warbler density should promote woodlands with high tree canopy cover, approximately 60–80% Ashe juniper composition, and tree heights >3 m. In contrast to a patch-based approach, our treatment of habitat variables as continuous helped to credibly map the warbler distribution across areas with broad transitions from woodlands to shrublands. By measuring these predictor variables at biologically relevant scales, we allowed the warbler survey data to define habitat relationships instead of using anthropogenically defined habitat patches. Outcomes from our study show the benefits of developing spatial products tailored to individual species of interest for conservation and management decisions.