Johannes J. Le Roux

Super-Genotype: Global Monoclonality Defies the Odds of Nature

The ability to respond to natural selection under novel conditions is critical for the establishment and persistence of introduced alien species and their ability to become invasive. Here we correlated neutral and quantitative genetic diversity of the weed Pennisetum setaceum Forsk. Chiov. (Poaceae) with differing global (North American and African) patterns of invasiveness and compared this diversity to native range populations. Numerous molecular markers indicate complete monoclonality within and among all of these areas (FST = 0.0) and is supported by extreme low quantitative trait variance (QST = 0.00065–0.00952). The results support the general-purpose-genotype hypothesis that can tolerate all environmental variation. However, a single global genotype and widespread invasiveness under numerous environmental conditions suggests a super-genotype. The super-genotype described here likely evolved high levels of plasticity in response to fluctuating environmental conditions during the Early to Mid Holocene. During the Late Holocene, when environmental conditions were predominantly constant but extremely inclement, strong selection resulted in only a few surviving genotypes.

Long-distance dispersal maximizes evolutionary potential during rapid geographic range expansion

Conventional wisdom predicts that sequential founder events will cause genetic diversity to erode in species with expanding geographic ranges, limiting evolutionary potential at the range margin. Here, we show that invasive European starlings (Sturnus vulgaris) in South Africa preserve genetic diversity during range expansion, possibly as a result of frequent long‐distance dispersal events. We further show that unfavourable environmental conditions trigger enhanced dispersal, as indicated by signatures of selection detected across the expanding range. This brings genetic variation to the expansion front, counterbalancing the cumulative effects of sequential founding events and optimizing standing genetic diversity and thus evolutionary potential at range margins during spread. Therefore, dispersal strategies should be highlighted as key determinants of the ecological and evolutionary performances of species in novel environments and in response to global environmental change.

Introduced and invasive cactus species: a global review

Understanding which species are introduced and become invasive are central questions in invasion science. In this sense, the cactus family is an interesting case study. Only 57 of the 1922 cactus species are currently recorded as invasive. There are three invasion hotspots: South Africa, Australia, and Spain. However, we identified large areas of the world with suitable climates. The invasive taxa represent an interesting subset of the total pool: they occur in two of the three major phylogenetic clades and in 13 of the 130 cactus genera, they possess five of the 12 cactus growth forms, and they tend to have larger native ranges.

Cultivation shapes genetic novelty in a globally important invader

Acacia saligna is a species complex that has become invasive in a number of countries worldwide where it has caused substantial environmental and economic impacts. Understanding genetic and other factors contributing to its success may allow managers to limit future invasions of closely related species. We used three molecular markers to compare the introduced range (South Africa) to the native range (Western Australia). Nuclear markers showed that invasive populations are divergent from native populations and most closely related to a cultivated population in Western Australia. We also found incongruence between nuclear and chloroplast data that, together with the long history of cultivation of the species, suggest that introgressive hybridization (coupled with chloroplast capture) may have occurred within A. saligna. While we could not definitively prove introgression, the genetic distance between cultivated and native A. saligna populations was comparable to known interspecific divergences among other Acacia species. Therefore, cultivation, multiple large‐scale introductions and possibly introgressive hybridization have rapidly given rise to the divergent genetic entity present in South Africa. This may explain the known global variation in invasiveness and inaccuracy of native bioclimatic models in predicting potential distributions.

Increasing functional modularity with residence time in the co-distribution of native and introduced vascular plants

Species gain membership of regional assemblages by passing through multiple ecological and environmental filters. To capture the potential trajectory of structural changes in regional meta-communities driven by biological invasions, one can categorize species pools into assemblages of different residence times. Older assemblages, having passed through more environmental filters, should become more functionally ordered and structured. Here we calculate the level of compartmentalization (modularity) for three different-aged assemblages (neophytes, introduced after 1500 AD; archaeophytes, introduced before 1500 AD, and natives), including 2,054 species of vascular plants in 302 reserves in central Europe. Older assemblages are more compartmentalized than younger ones, with species composition, phylogenetic structure and habitat characteristics of the modules becoming increasingly distinctive. This sheds light on two mechanisms of how alien species are functionally incorporated into regional species pools: the settling-down hypothesis of diminishing stochasticity with residence time, and the niche-mosaic hypothesis of inlaid neutral modules in regional meta-communities.

Spatial sorting drives morphological variation in the invasive bird, Acridotheris tristis.

The speed of range expansion in many invasive species is often accelerating because individuals with stronger dispersal abilities are more likely to be found at the range front. This ‘spatial sorting’ of strong dispersers will drive the acceleration of range expansion. In this study, we test whether the process of spatial sorting is at work in an invasive bird population (Common myna, Acridotheris tristis) in South Africa. Specifically, we sampled individuals across its invasive range and compared morphometric measurements relevant and non-relevant to the dispersal ability. Besides testing for signals of spatial sorting, we further examined the effect of environmental factors on morphological variations. Our results showed that dispersal-relevant traits are significantly correlated with distance from the range core, with strong sexual dimorphism, indicative of sex-biased dispersal. Morphological variations were significant in wing and head traits of females, suggesting females as the primary dispersing sex. In contrast, traits not related to dispersal such as those associated with foraging showed no signs of spatial sorting but were significantly affected by environmental variables such as the vegetation and the intensity of urbanisation. When taken together, our results support the role of spatial sorting in facilitating the expansion of Common myna in South Africa despite its low propensity to disperse in the native range.

Australian acacias as invasive species: lessons to be learnt from regions with long planting histories

Problems associated with invasiveness of non-native tree species used in forestry are increasing rapidly worldwide and are most severe in areas with a long history of plantings. Lessons learnt in areas with long histories of plantings and invasions may be applicable to areas with shorter planting histories. Most research towards understanding such tree invasions has focused on Pinus species, though all groups of trees that have been widely used in forestry are invasive to some extent. This paper explores the experience of Australian Acacia species (wattles). Unlike some other groups of trees, no particular set of traits clearly separates highly invasive from less- or non-invasive wattles. All species that have been widely planted over a long period have become invasive; the extent of invasions is largely a function of human usage. These findings imply that propagule pressure in concert with residence times are the main drivers of invasiveness in wattles (many factors mediate these drivers, including fire, forest clearance and soil disturbance). The massive extent of recent plantings of Australian Acacia species in South-east Asia is therefore likely to result in large-scale invasions unless proactive management is implemented. The history of wattles in South Africa highlights the need for such proactive management. Wattles were of considerable net value to the South African economy immediately after introduction. However, the costs of wattle invasions increased over time to such an extent that (certainly over the last few decades) these costs exceed the benefits derived from the forestry industry. Wattles now dominate many natural ecosystems. We recommend several interventions to prevent a similar pattern in South-east Asia and to ensure the sustainability of plantation forestry based on wattles in the region. A spatially explicit assessment of invasion risk is required, and a monitoring system should be implemented. Cost–benefit analyses (that consider the full suite of perspectives relating to costs and benefits) need to be applied to determine the need for sustainable mitigation methods. Options for reducing potential invasiveness should be implemented; these include biological control targeting seed production (very good success has been achieved in South Africa) and the use of sterile cultivars.

Elucidating the native sources of an invasive tree species, Acacia pycnantha, reveals unexpected native range diversity and structure.

BACKGROUND AND AIMS Understanding the introduction history of invasive plant species is important for their management and identifying effective host-specific biological control agents. However, uncertain taxonomy, intra- and interspecific hybridization, and cryptic speciation may obscure introduction histories, making it difficult to identify native regions to explore for host-specific agents. The overall aim of this study was to identify the native source populations of Acacia pycnantha, a tree native to south-eastern Australia and invasive in South Africa, Western Australia and Portugal. Using a phylogeographical approach also allowed an exploration of the historical processes that have shaped the genetic structure of A. pycnantha in its native range. METHODS Nuclear (nDNA) and plastid DNA sequence data were used in network and tree-building analyses to reconstruct phylogeographical relationships between native and invasive A. pycnantha populations. In addition, mismatch distributions, relative rates and Bayesian analyses were used to infer recent demographic processes and timing of events in Australia that led to population structure and diversification. KEY RESULTS The plastid network indicated that Australian populations of A. pycnantha are geographically structured into two informally recognized lineages, the wetland and dryland forms, whereas the nuclear phylogeny showed little geographical structure between these two forms. Moreover, the dryland form of A. pycnantha showed close genetic similarity to the wetland form based on nDNA sequence data. Hybrid zones may explain these findings, supported here by incongruent phylogenetic placement of some of these taxa between nuclear and plastid genealogies. CONCLUSIONS It is hypothesized that habitat fragmentation due to cycles of aridity inter-dispersed with periods of abundant rainfall during the Pleistocene (approx. 100 kya) probably gave rise to native dryland and wetland forms of A. pycnantha. Although the different lineages were confined to different ecological regions, we also found evidence for intraspecific hybridization in Victoria. The invasive populations in Portugal and South Africa represent wetland forms, whereas some South African populations resemble the Victorian dryland form. The success of the biological control programme for A. pycnantha in South Africa may therefore be attributed to the fact that the gall-forming wasp Trichilogaster signiventris was sourced from South Australian populations, which closely match most of the invasive populations in South Africa.

The structure of legume–rhizobium interaction networks and their response to tree invasions

We provide data on how legume-rhizobia interaction webs react to invasions by exotic legumes. This is the first study of its kind and found that general hypotheses derived from above-ground mutualistic webs may not hold for below-ground counterparts. Specifically, we found that legume-rhizobia interactions at the community level are highly specialised resulting in strongly modular webs, which are not nested, and that invasive legumes do not infiltrate existing native webs but rather form unique and novel modules in webs.

No consistent association between changes in genetic diversity and adaptive responses of Australian acacias in novel ranges

Common garden studies comparing trait differences of exotic species between native and introduced ranges rarely incorporate an analysis of genetic variation, but simply infer that trait shifts between ranges are genetically determined. We compared four growth-related traits (total biomass, relative growth rate RGR, specific leaf area SLA, and root to shoot ratio R:S) of five invasive Fabaceae species (Acacia cyclops, A. longifolia, A. melanoxylon, A. saligna, Paraserianthes lophantha), grown in a common garden experiment using seeds from introduced and native ranges across Australia. Chloroplast microsatellite loci were used to compare genetic diversity of native and introduced populations to determine standing genetic diversity and infer introduction history. We asked whether shifts in traits associated with faster growth due to enemy release in the introduced range were associated with levels of genetic diversity associated with introduction history. We found differences in traits between ranges, although these traits varied among the species. Compared to native-range populations, introduced-range Acacia longifolia had greater biomass and larger SLA; A. cyclops had greater RGR; and A. melanoxylon displayed lower R:S. Genetic diversity in the introduced range was lower for one of those species, A. longifolia, and two others that did not show differences in traits, A. saligna and P. lophantha. Diversity was higher in the introduced range for A. melanoxylon and did not differ among ranges for A. cyclops. These patterns of genetic diversity suggest that a genetic bottleneck may have occurred following the introduction of A. longifolia, A. saligna and P. lophantha. In contrast greater or comparable genetic diversity in the introduced range for A. melanoxylon and A. cyclops suggests introductions from multiple sources. This study has shown that a reduction in genetic diversity in the introduced range is not necessarily associated with a reduced capacity for adaptive responses or invasion potential in the novel range.