Margherita Gioria

Resource competition in plant invasions: emerging patterns and research needs

Invasions by alien plants provide a unique opportunity to examine competitive interactions among plants. While resource competition has long been regarded as a major mechanism responsible for successful invasions, given a well-known capacity for many invaders to become dominant and reduce plant diversity in the invaded communities, few studies have measured resource competition directly or have assessed its importance relative to that of other mechanisms, at different stages of an invasion process. Here, we review evidence comparing the competitive ability of invasive species vs. that of co-occurring native plants, along a range of environmental gradients, showing that many invasive species have a superior competitive ability over native species, although invasive congeners are not necessarily competitively superior over native congeners, nor are alien dominants are better competitors than native dominants. We discuss how the outcomes of competition depend on a number of factors, such as the heterogeneous distribution of resources, the stage of the invasion process, as well as phenotypic plasticity and evolutionary adaptation, which may result in increased or decreased competitive ability in both invasive and native species. Competitive advantages of invasive species over natives are often transient and only important at the early stages of an invasion process. It remains unclear how important resource competition is relative to other mechanisms (competition avoidance via phenological differences, niche differentiation in space associated with phylogenetic distance, recruitment and dispersal limitation, indirect competition, and allelopathy). Finally, we identify the conceptual and methodological issues characterizing competition studies in plant invasions, and we discuss future research needs, including examination of resource competition dynamics and the impact of global environmental change on competitive interactions between invasive and native species.

Early bird catches the worm: germination as a critical step in plant invasion

The germination behavior of a plant influences its fitness, persistence, and evolutionary potential, as well as its biotic environment. This can have major effects on the invasive potential of a species. We review the findings of four types of experimental studies comparing basic germination characteristics of invasive versus non-invasive congeners, in their non-native or native distribution range; invasive alien versus native species; and invasive species in their native versus non-native distribution range. Early and/or rapid germination is typical of invasive species rather than their non-invasive congeners, and represents a pre-adaptation from which many invasive and naturalized species benefit. It also occurs more often in invasive than native species, suggesting that competition mitigation or avoidance in the early stages of a plant’s life, via the exploitation of vacant germination niches, might be more useful than a superior competitive ability in novel environments. This is further supported by a tendency of invasive species to germinate earlier and/or faster and have broader germination cues in their non-native than in their native range. It is also supported by broader germination requirements being reported for invasive species than their non-invasive or native congeners. In contrast, high percentage germination is not a consistent predictor of invasiveness, suggesting that the incorporation of a larger fraction of seed production into the soil seed bank rather than high germination is a better (or safer) strategy in novel environments. These patterns indicate that differences in the germination behavior of alien and native species contribute to the invasiveness of many species, although evidence under natural conditions is needed. The role of such differences in the establishment and spread of invasive species in novel environments and their long-term impact on community dynamics requires further study.

Timing is everything: does early and late germination favor invasions by herbaceous alien plants?

Plant invasions represent a unique opportunity to study the mechanisms underlying community assembly rules and species distribution patterns. While a superior competitive ability has often been proposed as a major driver of successful plant invasions, its significance depends crucially on the timing of any competitive interaction. We assess whether a mismatch in germination phenology can favor the establishment of alien species, allowing them to exploit vacant niches where competition is low. As well as having important effects on the survival, growth and fitness of a species, asymmetric competition and potential soil legacies resulting from early or late germination can also impact on species recruitment. However, early or late germination comes at a cost, increases the risks of exposure to unfavorable conditions and requires an enhanced abiotic resistance if it is to lead to successful establishment. While there are several anecdotal accounts of early and late germination for invasive species, there are limited comparative data with resident species growing under natural conditions. Available evidence from grassland communities indicates that a short-term germination advantage or priority (few days/weeks) provides invasive species with a strong competitive advantage over native species and is a critical factor in many invasions. While the exploitation of periods of low competition is a plausible mechanism for the successful establishment of many invasive plants, direct evidence for this strategy is still scarce. This is particularly true with regard to the exploitation of late germination niches. Consequently, long-term comparative monitoring of the germination phenology of invasive and native plants in situ is needed to assess its significance in a range of ecosystems and its impact on community dynamics.

Biological Flora of the British Isles: Gunnera tinctoria

Summary 1. This account presents information on all aspects of the biology of Gunnera tinctoria (Molina) Mirb. (G. chilensis Lam.; G. scabra Ruix & Pav.; G. pilosa Kunth) that are relevant to understanding its ecological characteristics and behaviour. The main topics are presented within the standard framework of the Biological Flora of the British Isles: distribution, habitat, communities, responses to biotic factors, responses to environment, structure and physiology, phenology, floral and seed characters, herbivores and disease, history and conservation. 2. Gunnera tinctoria is a gynomonoecious, clonal, perennial herb that is naturalized in parts of Britain, becoming invasive in parts of Ireland and, more recently, Scotland. It occurs where winter temperatures are mild, and precipitation and humidity are high. Gunnera tinctoria is native to South America, predominantly in the Andean region of Chile and Colombia, and probably in parts of Argentina, ranging from sea level to c. 2000 m a.s.l. 3. Typical habitats in Britain and Ireland include stream and river banks, lake and pond margins, coastal cliffs, as well as disturbed areas, such as roadsides, quarries and ditches. In its native range it occurs predominantly on the banks of rivers and streams, on coastal cliffs and within canopy gaps or at the margins of temperate-humid rain forests. 4. Gunnera tinctoria occurs on a variety of substrates, mainly on alluvial or colluvial soils derived from volcanic material or on thin gley soils of marine origin. In Ireland, it occurs naturally on soils with a pH ranging from 4.6 to 6.2 and has been cultivated in soils with a pH up to 7. Soil moisture content and soil organic matter vary greatly, although it rarely colonizes highly organic soils such as peat. It is susceptible to even mild water deficits at all stages of development, and its seedlings are also sensitive to waterlogged conditions. 5. Gunnera tinctoria produces large numbers of seeds and also spreads clonally, by a horizontal rhizome system. It is wind pollinated, although insect pollination has been reported in New Zealand. Seeds are likely to be predominantly water and/or bird dispersed. In its invasive range, it can form a large and persistent soil seed bank. Recruitment from seeds seems to be important for its initial establishment, while vegetative propagation is the main means of expansion, leading to dense clonal stands. Long-distance seed dispersal seems to be central to the colonization of new areas, although the transport of vegetative propagules may also be important. 6. Gunnera tinctoria is a strong competitor in its invasive range, particularly in wet, humid environments. Its competitive ability arises from its large stature, the persistence of its seeds and rhizomes and a capacity for fixing nitrogen through a unique intracellular symbiosis with cyanobacteria (Nostoc) that may be particularly important for supporting the rapid growth of established plants early in the spring.

Distributional patterns of endemic, native and alien species along a roadside elevation gradient in Tenerife, Canary Islands

Invasion by alien plant species may be rapid and aggressive, causing erosion of local biodiversity. This is particularly true for islands, where natural and anthropogenic corridors promote the rapid spread of invasive plants. Although evidence shows that corridors may facilitate plant invasions, the question of how their importance in the spread of alien species varies along environmental gradients deserves more attention. Here, we addressed this issue by examining diversity patterns (species richness of endemic, native and alien species) along and across roads, along an elevation gradient from sea-level up to 2050 m a.s.l. in Tenerife (Canary Islands, Spain), at multiple spatial scales. Species richness was assessed using a multi-scale sampling design consisting of 59 T-transects of 150 m × 2 m, along three major roads each placed over the whole elevation gradient. Each transect was composed of three sections of five plots each: Section 1 was located on the road edges, Section 2 at intermediate distance, and Section 3 far from the road edge, the latter representing the “native community” less affected by road-specific disturbance. The effect of elevation and distance from roadsides was evaluated for the three groups of species (endemic, native and alien species), using parametric and non-parametric regression analyses as well as additive diversity partitioning. Differences among roads explained the majority of the variation in alien species richness and composition. Patterns in alien species richness were also affected by elevation, with a decline in richness with increasing elevation and no alien species recorded at high elevations. Elevation was the most important factor determining patterns in endemic and native species. These findings confirm that climate filtering reflected in varying patterns along elevational gradients is an important determinant of the richness of alien species (which are not adapted to high elevations), while anthropogenic pressures may explain the richness of alien species at low elevation.

Testing for differences in beta diversity from plot-to-plot dissimilarities

The central role of beta diversity in describing biodiversity patterns has been investigated in many fields of ecology and biogeography. While a variety of measures of beta diversity have been proposed over the past five decades, the question of how to test for differences in beta diversity among different sets of sampling plots has been addressed only rarely. Here, we describe a simple analytical procedure to test for differences in beta diversity among distinct sets of plots. The advantage of this approach compared to methods that have been previously proposed lies in its randomization procedure. Such a procedure creates a distribution of null values of the test statistic that is compatible with the null hypothesis of no significant difference in multivariate dispersion between the groups. The proposed test was illustrated using a large dataset of plant and water beetle (Coleoptera) assemblages collected from 45 farmland ponds in Ireland.