Soil Characteristics Drive Ficaria verna Abundance and Reproductive Output

Lesser celandine (Ficaria verna Huds.), an invasive plant from Europe, is becoming widespread in river valleys throughout the northeastern United States and the Pacific Northwest. Its high rate of asexual bulbil and tuber production creates dense infestations threatening native spring ephemerals. Ficaria verna abundance and reproductive output (seeds, bulbils, and tubers) were examined in invaded transects spanning a disturbance gradient away from a river. Site characteristics (photosynthetically active radiation [PAR], soil pH, moisture, texture, and nutrients) were quantified to examine their roles in plant abundance and reproduction. A larger-scale study examined random transects not specifically chosen based on F. verna infestations. Soil characteristics and slope were hypothesized to drive F. verna abundance and reproduction; we also hypothesized that reproductive output and biomass would be highest at intermediate distances from rivers, where disturbances are infrequent. Ficaria verna abundance and reproductive output varied considerably by site; soil characteristics, rather than landscape placement, appeared to drive plant abundance and reproduction. Lower percent sand was associated with significantly higher F. verna stem density and bulbil and tuber production. CEC was significantly negatively related to F. verna biomass and tuber counts. In the larger-scale survey, slope and PAR were significantly negatively related to F. verna presence and percent cover, respectively. Overall, these findings suggest that soil texture and slope can help explain higher abundance and reproductive outputs. However, reproductive output and biomass were not significantly greater at intermediate distances, contrary to expectations. We did not observe any seed production in any of the plots, although we did see a few plants with seeds outside our study area in the second year, demonstrating a near-complete reliance on asexual reproduction in these populations. This study expands on the current limited understanding of F. verna and can help management by identifying areas likely to support dense infestations

Influence of Microsite Disturbance on the Establishment of Two Congeneric Invasive Thistles

The successful establishment of invasive species has been shown to depend on aspects of the invaded community, such as gap characteristics. Biotic resistance may be particularly critical for stopping invaders at early life history stages, but new species can often invade following disturbances, which may create microsites with very different characteristics than are usually present. We examine the response of two invasive thistle species, Carduus nutans L. and C. acanthoides L., to three different microsite characteristics: disturbance type, size, and water availability. The two species initially responded differently to the type of disturbance: C. acanthoides had higher emergence and survival in plots with both above- and belowground disturbance, whereas C. nutans had better early performance in large microsites with above-ground disturbance only. Later in their life cycle, C. nutans performed better in plots that had been disturbed both above- and belowground, whereas C. acanthoides was largely unaffected by disturbance type. Increased emergence and survival, larger size and a higher proportion flowering were observed in larger gaps for both species throughout the life cycle. Watering had a negative impact on C. nutans emergence and fall survival and on C. acanthoides survival to the following summer. Overall, these results suggest that disturbance-generated microsite characteristics (disturbance type and size) may have large impacts on establishment of these two Carduus species, which in turn may persist well beyond the initial stages of growth. Studying invader responses to disturbance can help us to understand under what circumstances they are likely to establish and create persistent problems; avoiding or ameliorating such situations will have significant management benefits

Soil Characteristics Drive Ficaria verna Abundance and Reproductive Output

Lesser celandine (Ficaria verna Huds.), an invasive plant from Europe, is becoming widespread in river valleys throughout the northeastern United States and the Pacific Northwest. Its high rate of asexual bulbil and tuber production creates dense infestations threatening native spring ephemerals. Ficaria verna abundance and reproductive output (seeds, bulbils, and tubers) were examined in invaded transects spanning a disturbance gradient away from a river. Site characteristics (photosynthetically active radiation [PAR], soil pH, moisture, texture, and nutrients) were quantified to examine their roles in plant abundance and reproduction. A larger-scale study examined random transects not specifically chosen based on F. verna infestations. Soil characteristics and slope were hypothesized to drive F. verna abundance and reproduction; we also hypothesized that reproductive output and biomass would be highest at intermediate distances from rivers, where disturbances are infrequent. Ficaria verna abundance and reproductive output varied considerably by site; soil characteristics, rather than landscape placement, appeared to drive plant abundance and reproduction. Lower percent sand was associated with significantly higher F. verna stem density and bulbil and tuber production. CEC was significantly negatively related to F. verna biomass and tuber counts. In the larger-scale survey, slope and PAR were significantly negatively related to F. verna presence and percent cover, respectively. Overall, these findings suggest that soil texture and slope can help explain higher abundance and reproductive outputs. However, reproductive output and biomass were not significantly greater at intermediate distances, contrary to expectations. We did not observe any seed production in any of the plots, although we did see a few plants with seeds outside our study area in the second year, demonstrating a near-complete reliance on asexual reproduction in these populations. This study expands on the current limited understanding of F. verna and can help management by identifying areas likely to support dense infestations

Invasional Interference Due to Similar Inter- and Intraspecific Competition Between Invaders May Affect Management

As the number of biological invasions increases, the potential for invader– invader interactions also rises. The effect of multiple invaders can be superadditive (invasional meltdown), additive, or subadditive (invasional interference); which of these situations occurs has critical implications for prioritization of management efforts. Carduus nutans and C. acanthoides, two congeneric invasive weeds, have a striking, segregated distribution in central Pennsylvania, USA. Possible hypotheses for this pattern include invasion history and chance, direct competition, or negative interactions mediated by other species, such as shared pollinators. To explore the role of resource competition in generating this pattern, we conducted three related experiments using a response-surface design throughout the life cycles of two cohorts. Although these species have similar niche requirements, we found no differential response to competition between conspecifics vs. congeners. The response to combined density was relatively weak for both species. While direct competitive interactions do not explain the segregated distributional patterns of these two species, we predict that invasions of either species singly, or both species together, would have similar impacts. When prioritizing which areas to target to prevent the spread of one of the species, it is better to focus on areas as yet unaffected by its congener; where the congener is already present, invasional interference makes it unlikely that the net effect will change

Moving from Pattern to Process: Coexistence Mechanisms Under Intermediate Disturbance Regimes

Coexistence mechanisms that require environmental variation to operate contribute importantly to the maintenance of biodiversity. One famous hypothesis of diversity maintenance under disturbance is the intermediate disturbance hypothesis (IDH). The IDH proposes patterns of peaked diversity under intermediate disturbance regimes, based on a tension between competitively superior species and species which can rapidly colonize following disturbance. We review the literature, and describe recent research that suggests that more than one underlying mechanism can generate this unimodal diversity pattern in disturbed environments. Several exciting emerging research areas are identified, including interactions between disturbance types, operation of the IDH in multi-trophic systems, and changes in disturbance regimes. However, empirical work is still focussed on describing the IDH pattern, with little emphasis on identifying its mechanistic basis. We discuss how to extend methods for identifying different coexistence mechanisms, developed in the theoretical literature, to experimental research. In an attempt to operationalize these various ideas we outline a hypothetical IDH research programme. A solid understanding of the life history attributes of the component species and their responses to disturbance will facilitate identification of the coexistence mechanism(s) underlying the IDH pattern, and provide a framework by which empirical and theoretical results can be more fully integrated

Moving from Pattern to Process: Coexistence Mechanisms Under Intermediate Disturbance Regimes

Coexistence mechanisms that require environmental variation to operate contribute importantly to the maintenance of biodiversity. One famous hypothesis of diversity maintenance under disturbance is the intermediate disturbance hypothesis (IDH). The IDH proposes patterns of peaked diversity under intermediate disturbance regimes, based on a tension between competitively superior species and species which can rapidly colonize following disturbance. We review the literature, and describe recent research that suggests that more than one underlying mechanism can generate this unimodal diversity pattern in disturbed environments. Several exciting emerging research areas are identified, including interactions between disturbance types, operation of the IDH in multi-trophic systems, and changes in disturbance regimes. However, empirical work is still focussed on describing the IDH pattern, with little emphasis on identifying its mechanistic basis. We discuss how to extend methods for identifying different coexistence mechanisms, developed in the theoretical literature, to experimental research. In an attempt to operationalize these various ideas we outline a hypothetical IDH research programme. A solid understanding of the life history attributes of the component species and their responses to disturbance will facilitate identification of the coexistence mechanism(s) underlying the IDH pattern, and provide a framework by which empirical and theoretical results can be more fully integrated

Plant Community Associations of Two Invasive Thistles

In order to combat the growing problems associated with biological invasions, many researchers have focused on identifying which communities are most vulnerable to invasion by exotic species. However, once established, invasive species can significantly change the composition of the communities that they invade. The first step to disentangling the direction of causality is to discern whether a relationship with other vegetation exists at all. Carduus nutans and C. acanthoides are similar invasive thistles, which have caused substantial economic damage worldwide. We assessed the associations between the thistles and the standing flora in four sites in central Pennsylvania in which they co-occur. After sampling nearly 2000 plots of 1 m2, we used partial Mantel tests to assess the differences in vegetation between thistle and non-thistle plots after accounting for location, and non-metric multidimensional scaling to visualize differences among plots and sites. We found significant differences in community composition in plots with and without Carduus thistles. The non-native species Sisymbrium officinale and Coronilla varia were consistently associated with the presence of Carduus thistles. Several species were associated with areas that were free of Carduus thistles, including an important non-native pasture species (Trifolium repens). We found no evidence for differences in composition between plots with C. nutans versus C. acanthoides, suggesting that they have similar associations with the vegetation community. We conclude that even at the within-field scale, areas invaded by Carduus thistles have different vegetation associations than uninvaded areas, allowing us to target future research about the role of vegetation structure in resisting and responding to invasion

Plant Community Associations of Two Invasive Thistles

In order to combat the growing problems associated with biological invasions, many researchers have focused on identifying which communities are most vulnerable to invasion by exotic species. However, once established, invasive species can significantly change the composition of the communities that they invade. The first step to disentangling the direction of causality is to discern whether a relationship with other vegetation exists at all. Carduus nutans and C. acanthoides are similar invasive thistles, which have caused substantial economic damage worldwide. We assessed the associations between the thistles and the standing flora in four sites in central Pennsylvania in which they co-occur. After sampling nearly 2000 plots of 1 m2, we used partial Mantel tests to assess the differences in vegetation between thistle and non-thistle plots after accounting for location, and non-metric multidimensional scaling to visualize differences among plots and sites. We found significant differences in community composition in plots with and without Carduus thistles. The non-native species Sisymbrium officinale and Coronilla varia were consistently associated with the presence of Carduus thistles. Several species were associated with areas that were free of Carduus thistles, including an important non-native pasture species (Trifolium repens). We found no evidence for differences in composition between plots with C. nutans versus C. acanthoides, suggesting that they have similar associations with the vegetation community. We conclude that even at the within-field scale, areas invaded by Carduus thistles have different vegetation associations than uninvaded areas, allowing us to target future research about the role of vegetation structure in resisting and responding to invasion

Coexistence Patterns of Two Invasive Thistle Species, Carduus nutans and C. acanthoides, at Three Spatial Scales

To better understand the competitive processes involved in invasion by congeners, we examine coexistence patterns of two invasive species, Carduus nutans and C. acanthoides, at three spatial scales. A roadside survey of 5 × 5 km blocks in a previously identified overlap zone provided information about the regional scale. At smaller scales, we surveyed four fields of natural co-occurrence, quantifying the spatial patterns at the field scale by randomly placed 1 × 1 m quadrats and at the smallest scale by detailing plant position within the quadrats. The patterns observed are strikingly different at the different scales. At the regional scale, there is positive local autocorrelation in both species but negative cross-correlation between them, consistent with previous surveys. However, at the field scale, there is positive local autocorrelation in both species, and we generally see a positive association between the two species. At the plot scale, when excluding areas of joint absence, there is again a negative association between the two species. This pattern can also be seen at the field scale when excluding plots with joint absence. These results suggest that, at the scale of a field, the strongest factor determining location is aggregation in favorable habitats, which is a stronger force than the competition-induced segregation evidenced at small scales. Lottery competition for spatially aggregated safe sites thus appears to drive the patterns observed at the field scale, while the regional scale pattern may be a result of restricted natural dispersal and invasion history