Dynamics and mechanisms of secondary invasion following biological control of an invasive plant.

Secondary invasions in which nontarget invaders expand following eradication of a target invader commonly occur in habitats with multiple invasive plant species and can prevent recovery of native communities. However, the dynamics and mechanisms of secondary invasion remain unclear. Here, we conducted a common garden experiment to test underlying mechanisms of secondary invasion for 14 nontarget invaders after biological control of Ambrosia artemisiifolia in two consecutive years. We found secondary invasion for all tested nontarget invaders, but secondary invasiveness (change relative to natives) varied with species and time. Specifically, secondary invasiveness depended most strongly on phylogenetic relatedness between the target and nontarget invaders in the first year with closely related nontarget invaders being most invasive. By contrast, secondary invasiveness in the second year was mostly driven by functional traits with taller nontarget invaders or those with higher specific leaf area, or smaller seeds especially invasive. Our study indicates that secondary invasion is likely to occur wherever other invasive plants co-occur with an invasive species targeted for control. Furthermore, the most problematic invaders will initially be species closely related to the target invader but then species with rapid growth and high reproduction are most likely to be more aggressive secondary invaders.

Effects of nutrient pulses on exotic species shift from positive to neutral with decreasing water availability.

Temporal fluctuation in nutrient availability generally promotes the growth of exotic plant species and has been recognized as an important driver of exotic plant invasions. However, little is known about how the impact of fluctuating nutrients on exotic species is dependent on the availability of other resources, although most ecosystems are experiencing dramatic variations in a wide variety of resources due to global change and human disturbance. Here, we explored how water availability mediates the effect of nutrient pulses on the growth of six exotic and six native plant species. We subjected individual plants of exotic and native species to well watered or water stressed conditions. For each level of water availability, we added equivalent amounts of nutrients at a constant rate, as a single large pulse, or in multiple small pulses. Under well watered conditions, nutrient pulses promoted exotic plant growth relative to nutrients supplied constantly, while they had no significant effect on natives. In contrast, under water stressed conditions, water deficiency inhibited the growth of all exotic and native species. More importantly, nutrient pulses did not increase plant growth relative to nutrients supplied constantly and these phenomena were observed for both exotic and native species. Taken together, our study shows that the impact of fluctuating nutrient availability on the growth of exotic plant species strongly depends on the variation of other resources, and that the positive effect of nutrient pulses under well watered conditions disappears under water stressed conditions. Our findings suggest that the variation in multiple resources may have complex feedback on exotic plant invasions and, therefore, it is critical to encompass multiple resources for the evaluation of fluctuating resource availability effects on exotic plant species. This will allow us to project the invasive trajectory of exotic plant species more accurately under future global change and human disturbance.

Fluctuations in resource availability shape the competitive balance among non-native plant species.

Fluctuating resource availability plays a critical role in determining non-native plant invasions through mediating the competitive balance between non-native and native species. However, the impact of fluctuating resource availability on interactions among non-native species remains largely unknown. This represents a barrier to understanding invasion mechanisms, particularly in habitats that harbor multiple non-native species with different responses to fluctuating resource availability. To examine the responses of non-native plant species to nutrient fluctuations, we compared the growth of each of 12 non-native species found to be common in local natural areas to nutrients supplied at a constant rate or supplied as a single large pulse in a pot experiment. We found that seven species produced more biomass with pulsed nutrients compared to constant nutrients (hereafter "benefitting species"), while the other five species did not differ between nutrient enrichment treatments (hereafter "non-benefitting species"). To investigate how nutrient fluctuations influence the interactions among non-native plant species, we established experimental non-native communities in the field with two benefitting and two non-benefitting non-native species. Compared with constant nutrient supply, the single large pulse of nutrient did not influence community biomass, but strongly increased the biomass and cover of the benefitting species and decreased those of the non-benefitting species. Furthermore, the benefitting species had higher leaf N content and greater plant height when nutrients were supplied as a single large pulse than at a constant rate, whereas the non-benefitting species showed no differences in leaf N content and were shorter when nutrients were supplied as a single large pulse than at a constant rate. Our results add to the growing evidence that the individual responses of non-native species to nutrient fluctuation are species-specific. More importantly, benefitting species were favored by nutrients coming in a pulse, while non-benefitting ones were favored by nutrients coming constantly when they grew together. This suggests that nutrient fluctuations can mediate the competitive balance among non-native plants and may thus determine their invasion success in a community harboring multiple non-native plant species.

Morphological trait-matching in plant-Hymenoptera and plant-Diptera mutualisms across an elevational gradient.

Morphological trait-matching and species abundance are thought to be the main factors affecting the frequency and strength of mutualistic interactions. However, the relative importance of trait-matching and species abundance in shaping species interactions across environmental gradients remains poorly understood, especially for plant-insect mutualisms involving generalist species. Here, we characterised variation in species and trait composition and the relative importance of trait-matching and species abundance in shaping plant-Hymenoptera and plant-Diptera mutualisms in four meadows across an elevational gradient (2,725-3,910 m) in Yulong Snow Mountain, Southwest China. We also evaluated the effects of morphological traits of flower visitors and plant composition on their foraging specialisation (d' and normalised degree). There was a high degree of dissimilarity in the composition of Hymenoptera and Diptera visitors and their visited plants between communities. This variation was mainly driven by the spatial replacement of species. Both for plant-Hymenoptera and plant-Diptera networks, trait-matching between nectar tube depth and proboscis length was a stronger predictor of the interactions between temporally co-occurring plants and flower visitors than species abundance. Fourth-corner analyses revealed statistically significant trait-matching between nectar tube depth and proboscis length in plant-Hymenoptera networks at all sites, suggesting that Hymenoptera consistently foraged on plant species with nectar tube depths matching their proboscis lengths. By contrast, significant trait-matching in plant-Diptera networks was only observed at the two lower elevation sites. The species-level specialisation d' of flower visitors increased significantly as the proboscis length and the difference in nectar tube depth between the plant community and the plants visited by flower visitors increased. Our results highlight that the importance of trait-matching in shaping pairwise interactions and niche partitioning depends on the specific features (e.g. species composition and trait availability) of the plant-pollinator system. For specialised plant-Hymenoptera systems, trait-matching is an important determinant of species interactions, whereas for generalist plant-Diptera systems, trait-matching is relatively unimportant.

Addressing challenges of an ageing population by rationally forecasting travel behaviours of the elderly: a case study in China.

This research aims to investigate the difference of the travel behaviours of the elderly in different urban areas. On the basis of the results of cluster analysis, the nested logit (NL) model is adopted to investigate the travel decision of the elderly in central urban areas and non-central urban areas. Moreover, it is used to forecast the change of their travel behaviours by implementing the policies of age-friendly society. The results show that the elderly in central urban areas are inclined to decide travel modes in advanced of determining trip chains, while the trip chains are first determined in non-central urban areas. For the central urban areas, the policies mainly affect the choice of travel modes of the elderly, rather than the choice of trip chains. However, for the non-central urban area, the policies simultaneously influence the choice of travel mode and trip chains of the elderly. Based on the results of the forecast, some measures are proposed in this study to meet different travel requirements of the elderly in central urban areas and non-central urban areas.

Incomplete reproductive isolation between Rhododendron taxa enables hybrid formation and persistence.

The evolutionary consequences of hybridization ultimately depend on the magnitude of reproductive isolation between hybrids and their parents. We evaluated the relative contributions of pre- and post-zygotic barriers to reproduction for hybrid formation, hybrid persistence and potential for reproductive isolation of hybrids formed between two Rhododendron species, R. spiciferum and R. spinuliferum. Our study established that incomplete reproductive isolation promotes hybrid formation and persistence and delays hybrid speciation. All pre-zygotic barriers to reproduction leading to hybrid formation are incomplete: parental species have overlapping flowering; they share the same pollinators; reciprocal assessments of pollen tube germination and growth do not differ among parents. The absence of post-zygotic barriers between parental taxa indicates that the persistence of hybrids is likely. Reproductive isolation was incomplete between hybrids and parents in all cases studied, although asymmetric differences in reproductive fitness were prevalent and possibly explain the genetic structure of natural hybrid swarms where hybridization is known to be bidirectional but asymmetric. Introgression, rather than speciation, is a probable evolutionary outcome of hybridization between the two Rhododendron taxa. Our study provides insights into understanding the evolutionary implications of natural hybridization in woody plants.

Specialist reassociation and residence time modulate the evolution of defense in invasive plants: A meta-analysis.

Invasive plants typically escape specialist herbivores but are often attacked by generalist herbivores in their introduced ranges. The shifting defense hypothesis suggests that this will cause invasive plants to evolve lower resistance against specialists, higher resistance against generalists, and greater tolerance to herbivore damage. However, the duration and direction of selective pressures can shape the evolutionary responses of resistance and tolerance for invasive plants. Two critical factors are (1) residence time (length of time that an invasive species has been in its introduced range) and (2) specialist herbivore reassociation (attack by purposely or accidentally introduced specialists). Yet, these two factors have not been considered simultaneously in previous quantitative syntheses. Here, we performed a meta-analysis with 367 effect sizes from 70 studies of 35 invasive plant species from native and invasive populations. We tested how the residence time of invasive plant species and specialist reassociation in their introduced ranges affected evolutionary responses of defenses against specialists and generalists, including herbivore resistance traits (physical barriers, digestibility reducers and toxins), resistance effects (performance of and damage caused by specialists or generalists) and tolerance to damage (from specialists or generalists). We found that residence time and specialist reassociation each significantly altered digestibility reducers, specialist performance, generalist damage, and tolerance to specialist damage. Furthermore, residence time and specialist reassociation strongly altered toxins and generalist performance, respectively. When we restricted consideration to invasive plant species with both longer residence times and no reassociation with specialists, invasive populations had lower resistance to specialists, similar resistance to generalists, and higher tolerance to damage from both herbivore types, compared with native populations. We conclude that the duration and direction of selective pressure shape the evolutionary responses of invasive plants. Under long-term (long residence time) and stable (no specialist reassociation) selective pressure, invasive plants generally decrease resistance to specialists and increase tolerance to generalist damage that provides mixed support for the shifting defense hypothesis.

Reproductive Isolation Among Three Nocturnal Moth-Pollinated Sympatric Habenaria Species (Orchidaceae).

Comparison and quantification of multiple pre- and post-pollination barriers to interspecific hybridization are important to understand the factors promoting reproductive isolation. Such isolating factors have been studied recently in many flowering plant species which seek after the general roles and relative strengths of different pre- and post-pollination barriers. In this study, we quantified six isolating factors (ecogeographic isolation, phenological isolation, pollinator isolation, pollinia-pistil interactions, fruit production, and seed development) that could possibly be acting as reproductive barriers at different stages among three sympatric Habenaria species (H. limprichtii, H. davidii, and H. delavayi). These three species overlap geographically but occupy different microhabitats varying in soil water content. They were isolated through pollinator interactions both ethologically (pollinator preference) and mechanically (pollinia attachment site), but to a variable degree for different species pairs. Interspecific crosses between H. limprichtii and H. davidii result in high fruit set, and embryo development suggested weak post-pollination barriers, whereas bidirectional crosses of H. delavayi with either of the other two species fail to produce fruits. Our results revealed that pollinators were the most important isolating barrier including both ethological and mechanical mechanisms, to maintain the boundaries among these three sympatric Habenaria species. Our study also highlights the importance of a combination of pre-and post-pollination barriers for species co-existence in Orchidaceae.

Species-specific plant-mediated effects between herbivores converge at high damage intensity.

Plants are often exposed to multiple herbivores and densities of these attackers (or corresponding damage intensities) often fluctuate greatly in the field. Plant-mediated interactions vary among herbivore species and with changing feeding intensity, but little is known about how herbivore identity and density interact to determine plant responses and herbivore fitness. Here, we investigated this question using Triadica sebifera (tallow) and two common and abundant specialist insect herbivores, Bikasha collaris (flea beetle) and Heterapoderopsis bicallosicollis (weevil). By manipulating densities of leaf-feeding adults of these two herbivore species, we tested how variations in the intensity of leaf damage caused by flea beetle or weevil adults affected the performance of root-feeding flea beetle larvae and evaluated the potential of induced tallow root traits to predict flea beetle larval performance. We found that weevil adults consistently decreased the survival of flea beetle larvae with increasing leaf damage intensities. In contrast, conspecific flea beetle adults increased their larval survival at low damage then decreased larval survival at high damage, resulting in a unimodal pattern. Chemical analyses showed that increasing leaf damage from weevil adults linearly decreased root carbohydrates and increased root tannin, whereas flea beetle adults had opposite effects as weevil adults at low damage and similar effects as them at high damage. Furthermore, across all feeding treatments, flea beetle larval survival correlated positively with concentrations of carbohydrates and negatively with concentration of tannin, suggesting that root primary and secondary metabolism might underlie the observed effects on flea beetle larvae. Our study demonstrates that herbivore identity and density interact to determine systemic plant responses and plant-mediated effects on herbivores. In particular, effects are species-specific at low densities, but converge at high densities. These findings emphasize the importance of considering herbivore identity and density simultaneously when investigating factors driving plant-mediated interactions between herbivores, which advances our understanding of the structure and composition of herbivore communities and terrestrial food webs.

Pollination ecology in China from 1977 to 2017.

China is one of most biodiverse countries in the world, containing at least 10% of all angiosperm species. Therefore, we should anticipate a diverse, pollinator fauna. China also has a long history of applied ethnobiology, including a sustainable agriculture based on apiculture and plant-pollinator interactions. However, the science of pollination ecology is a far younger sub-discipline in China, compared to in the West. Chinese studies in pollination ecology began in the 1970s. For this review, we compiled a complete reference database (>600 publications) of pollination studies in China. Using this database, we identified and analyzed gaps and limitations in research on the pollination systems of native and naturalized species. Specifically, we asked the following questions: 1) What do we know about the pollination systems of native, Chinese species? 2) How does Chinese pollination ecology compare with the development of pollination research abroad and which aspects of research should be pursued by Chinese anthecologists in the near future? 3) What research on pollination in China will advance our understanding and contribute to our ongoing analyses of endemism and conservation? Subsequently, we segregated and identified prospective lines of future research that are unique to China and can only be done in China. This requires discussing priorities within a systematic approach.

Does reproductive isolation reflect the segregation of color forms in Spiranthes sinensis (Pers.) Ames complex (Orchidaceae) in the Chinese Himalayas?

Isolation between species, or taxa sharing a common lineage, depends primarily on the relative strengths of various reproductive barriers. Previous studies on reproductive isolation between orchids emphasized mechanical and ethological barriers in flowers of species showing food and/or sexual mimicry. In this study, we investigated and quantified a series of prepollination and postpollination barriers between pink and white forms of Spiranthes sinensis sl, a nectar-secreting complex. We generated ML trees based on trnS-G and matK to explore phylogenetic relationships in this species complex. Spiranthes sinensis sl segregated from some other congeners, but the white form constituted a distinct clade in relation to the pink form. The white form secreted 2-Phenylethanol as it is a single-scent compound and was pollinated almost exclusively by native, large-bodied Apis cerana and Bombus species (Apidae). Apis cerana showed a high floral constancy to this form. The scentless, pink form was pollinated primarily by smaller bees in the genera Ceratina (Apidae), and members of the family Halictidae, with infrequent visits by A. cerana and Bombus species. Fruit set and the production of large embryos following interform pollination treatments were significantly lower compared to intraform pollination results for the white form. Our results suggested that pollinator isolation, based on color and scent cues, may result in greater floral constancy in white populations when both forms are sympatric as two different, guilds of pollinators forage selectively preventing or reducing prospective gene flow. Postpollination barriers appear weaker than prepollination barriers but they also play a role in interform isolation, especially in the white form. Our findings suggest that floral color forms in S. sinensis do not represent an unbalanced polymorphism. Interpretations of the evolutionary status of these forms are discussed.