Many notorious invasive plants are clonal, spreading mainly by vegetative propagules. Propagule pressure (the number of propagules) may affect the establishment, growth and thus invasion success of these clonal plants, and such effects may also depend on habitat conditions. To understand how propagule pressure, habitat conditions and clonal integration affect the establishment and growth of the invasive clonal plants, an 8-week greenhouse with an invasive clonal plant, Alternanthera philoxeroides was conducted. High (five fragments) or low (one fragment) propagule pressure was established either in bare soil (open habitat) or dense native vegetation of Jussiaea repens (vegetative habitat), with the stolon connections either severed from or connected to the relatively older ramets. High propagule pressure greatly increased the establishment and growth of A. philoxeroides, especially when it grew in vegetative habitats. Surprisingly, high propagule pressure significantly reduced the growth of individual plants of A. philoxeroides in open habitats, whereas it did not affect the individual growth in vegetative habitats. A shift in the intraspecific interaction on A. philoxeroides from competition in open habitats to facilitation in vegetative habitats may be the main reason. Moreover, clonal integration significantly improved the growth of A. philoxeroides only in open habitats, especially with low propagule pressure, whereas it had no effects on the growth and competitive ability of A. philoxeroides in vegetative habitats, suggesting that clonal integration may be of most important for A. philoxeroides to explore new open space and spread. These findings suggest that propagule pressure may be crucial for the invasion success of A. philoxeroides, and such an effect also depends on habitat conditions.

Clonal traits can contribute to plant invasiveness, but little is known about the roles of division of labor (a key clonal trait) in homogeneous habitats. The hypothesis tested is that clonal integration allows division of labor and increases the overall performance of an invasive clonal plant, especially under higher soil nutrients. Clonal fragment pairs of aquatic invader Alternanthera philoxeroides (each with four ramets and a stolon apex) were grown in two homogenous habitats with high or low soil nutrient supply, and with stolon connections being either severed (clonal integration prevented) or kept intact (clonal integration allowed). Results showed that stolon connection allowed the division of labor within the clonal fragment, with basal ramets specializing in acquisition of belowground resources and apical ramets specializing in acquisition of aboveground expansion. Moreover, the capacity for division of labor was greater, which brought the clonal fragments of A. philoxeroides stronger clonal propagation and better performance in high nutrient habitats than in low nutrient habitats. The results supported our hypotheses that the developmentally programmed division of labor may facilitate the clonal expansion of this aggressive invader in some homogeneous habitats with high resource availability.

Different connected parts of clonal plants often grow in different patches and the resource contrast between patches has an important effect on the material transfer between the connected ramets. However, it is unclear whether the effect of clonal integration differs between the invasive clonal plant and the related native species in response to patch contrast. To explore this, we grew the clonal fragment pairs of plant invader Alternanthera philoxeroides and its co-genus native species A. sessilis under high contrast, low contrast, and no contrast (control) nutrient patch environments, respectively, and with stolon connections either severed or kept intact. The results showed that, at the ramet level, clonal integration (stolon connection) significantly improved the growth of apical ramets of both species, and such positive effects were significantly greater in A. philoxeroides than in A. sessilis. Moreover, clonal integration greatly increased the chlorophyll content index of apical ramets and the growth of basal ramets in A. philoxeroides but not in A. sessilis under low and high contrast. At the whole fragment level, the benefits of clonal integration increased with increasing patch contrast, and such a positive effect was more pronounced in A. philoxeroides than in A. sessilis. This study demonstrated that A. philoxeroides possesses a stronger ability of clonal integration than A. sessilis, especially in patchy environments with a higher degree of heterogeneity, suggesting that clonal integration may give some invasive clonal plants a competitive advantage over native species, thus facilitating their invasion in patchy habitats.