Biological invasions and climate change amplify each other’s effects on dryland degradation

Climate models predict that, in the coming decades, many arid regions will experience increasingly hot conditions and will be affected more frequently by drought. These regions are also experiencing rapid vegetation change, notably invasion by exotic grasses. Invasive grasses spread rapidly into native desert ecosystems due, in particular, to interannual variability in precipitation and periodic fires. The resultant destruction of non-fire-adapted native shrub and grass communities and of the inherent soil resource heterogeneity can yield invader-dominated grasslands. Moreover, recurrent droughts are expected to cause widespread physiological stress and mortality of both invasive and native plants, as well as the loss of soil resources. However, the magnitude of these effects may differ between invasive and native grasses, especially under warmer conditions, rendering the trajectory of vegetated communities uncertain. Using the Biosphere 2 facility in the Sonoran Desert, we evaluated the viability of these hypothesized relationships by simulating combinations of drought and elevated temperature (+5°C) and assessing the ecophysiological and mortality responses of both a dominant invasive grass (Pennisetum ciliare or buffelgrass) and a dominant native grass (Heteropogan contortus or tanglehead). While both grasses survived protracted drought at ambient temperatures by inducing dormancy, drought under warmed conditions exceeded the tolerance limits of the native species, resulting in greater and more rapid mortality than exhibited by the invasive. Thus, two major drivers of global environmental change, biological invasion and climate change, can be expected to synergistically accelerate ecosystem degradation unless large-scale interventions are enacted.     

Invader presence disrupts the stabilizing effect of species richness in plant community recovery after drought

Higher biodiversity can stabilize the productivity and functioning of grassland communities when subjected to extreme climatic events. The positive biodiversity–stability relationship emerges via increased resistance and/or recovery to these events. However, invader presence might disrupt this diversity–stability relationship by altering biotic interactions. Investigating such disruptions is important given that invasion by non‐native species and extreme climatic events are expected to increase in the future due to anthropogenic pressure. Here we present one of the first multisite invader × biodiversity × drought manipulation experiment to examine combined effects of biodiversity and invasion on drought resistance and recovery at three semi‐natural grassland sites across Europe. The stability of biomass production to an extreme drought manipulation (100% rainfall reduction; BE: 88 days, BG: 85 days, DE: 76 days) was quantified in field mesocosms with a richness gradient of 1, 3, and 6 species and three invasion treatments (no invader, Lupinus polyphyllus, Senecio inaequidens). Our results suggest that biodiversity stabilized community productivity by increasing the ability of native species to recover from extreme drought events. However, invader presence turned the positive and stabilizing effects of diversity on native species recovery into a neutral relationship. This effect was independent of the two invader's own capacity to recover from an extreme drought event. In summary, we found that invader presence may disrupt how native community interactions lead to stability of ecosystems in response to extreme climatic events. Consequently, the interaction of three global change drivers, climate extremes, diversity decline, and invasive species, may exacerbate their effects on ecosystem functioning.     

Leaf and stem physiological responses to summer and winter extremes of woody species across temperate ecosystems

Winter cold limits temperate plant performance, as does summer water stress in drought‐prone ecosystems. The relative impact of seasonal extremes on plant performance has received considerable attention for individual systems. An integrated study compiling the existing literature was needed to identify overall trends. First, we conducted a meta‐analysis of the impacts of summer and winter on ecophysiology for three woody plant functional types (winter deciduous angiosperms, evergreen angiosperms and conifers), including data for 210 records from 75 studies of ecosystems with and without summer drought across the temperate zone. Second, we tested predictions by conducting a case study in a drought‐prone Mediterranean ecosystem subject to winter freezing. As indicators of physiological response of leaves and xylem to seasonal stress, we focused on stomatal conductance (g_s), percent loss of stem xylem hydraulic conductivity (PLC) and photochemical efficiency of photosystem II (F_v/F_m). Our meta‐analysis showed that in ecosystems without summer drought, g_s was higher during summer than winter. By contrast, in drought‐prone ecosystems many species maintained open stomata during winter, with potential strong consequences for plant carbon gain over the year. Further, PLC tended to increase and F_v/F_m to decrease from summer to winter for most functional types and ecosystems due to low temperatures. Overall, deciduous angiosperms were most sensitive to climatic stress. Leaf gas exchange and stem xylem hydraulics showed a coordinated seasonal response at ecosystems without summer drought. In our Mediterranean site subjected to winter freezing the species showed similar responses to those typically found for ecosystems without summer drought. We conclude that winter stress is most extreme for systems without summer drought and systems with summer drought and winter freezing, and less extreme for drought‐prone systems without freezing. In all cases the evergreen species show less pronounced seasonal responses in both leaves and stems than deciduous species.     

Functionally Similar Species Confer Greater Resistance to Invasion: Implications for Grassland Restoration

Plant community functional composition can be manipulated in restored ecosystems to reduce the establishment potential of invading species. This study was designed to compare invasion resistance among communities with species functionally similar or dissimilar to yellow starthistle (Centaurea solstitialis), a late‐season annual. A field experiment was conducted in the Central Valley of California with six experimental plant communities that included (1) six early‐season native annual forbs (AF); (2) five late‐season native perennials and one summer annual forb (NP); (3) a combination of three early‐season native annual forbs and three late‐season native perennials (FP); (4) six early‐season non‐native annual grasses (AG); (5) monoculture of the late‐season native perennial grass Elymus glaucus (EG); and (6) monoculture of the late‐season native perennial Grindelia camporum (GC). Following establishment, C. solstitialis seed was added to half of the plots, and a monoculture of C. solstitialis (CS) was established as a control. Over a 5‐year period, the AF and AG communities were ineffective at preventing C. solstitialis invasion. Centaurea solstitialis cover remained less than 10% in the FP and NP communities, except in year 1. By the fourth year, E. glaucus cover was greater than 50% in NP and FP communities and had spread to all other communities (e.g., 27% cover in CS in year 5). Communities containing E. glaucus, which is functionally similar to C. solstitialis, better resisted invasion than communities lacking a functional analog. In contrast, G. camporum, which is also functionally similar to C. solstitialis, failed to survive. Consequently, species selection for restored communities must consider not only functional similarity to the invader but also establishment success, competitiveness, and survivorship.     

Effects of extreme weather events and legume presence on mycorrhization of Plantago lanceolata and Holcus lanatus in the field

Little is known about direct and indirect effects of extreme weather events on arbuscular mycorrhizal fungi (AMF) under field conditions. In a field experiment, we investigated the response of mycorrhization to drought and heavy rain in grassland communities. We quantified AMF biomass in soil, mycorrhization of roots of the grass Holcus lanatus and the forb Plantago lanceolata, as well as plant performance. Plants were grown in four‐species communities with or without a legume. We hypothesised that drought increases and heavy rain decreases mycorrhization, and that higher mycorrhization will be linked to improved stress resistance and higher biomass production. Soil AMF biomass increased under both weather extremes. Heavy rain generally benefitted plants and increased arbuscules in P. lanceolata. Drought neither reduced plant performance nor root mycorrhization. Arbuscules increased in H. lanatus several weeks after drought, and in P. lanceolata several weeks after heavy rain spells. These long‐lasting effects of weather events on mycorrhization highlight the indirect influence of climate on AMF via their host plant. Legume presence increased plant community biomass, but had only minor effects on mycorrhization. Arbuscule colonisation was negatively correlated with senescence during the dry summer. Mycorrhization and biomass production in P. lanceolata were positively related. However, increased mycorrhization was related to less biomass in the grass. AMF mycelium in soil might generally increase under extreme events, root colonisation, however, is host species specific. This might amplify community shifts in grassland under climate change by further increasing stress resistance of species that already benefit from changed precipitation.     

Methods to test the interactive effects of drought and plant invasion on ecosystem structure and function using complementary common garden and field experiments

Abiotic global change drivers affect ecosystem structure and function, but how they interact with biotic factors such as invasive plants is understudied. Such interactions may be additive, synergistic, or offsetting, and difficult to predict. We present methods to test the individual and interactive effects of drought and plant invasion on native ecosystems. We coupled a factorial common garden experiment containing resident communities exposed to drought (imposed with rainout shelters) and invasion with a field experiment where the invader was removed from sites spanning a natural soil moisture gradient. We detail treatments and their effects on abiotic conditions, including soil moisture, light, temperature, and humidity, which shape community and ecosystem responses. Ambient precipitation during the garden experiment exceeded historic norms despite severe drought in prior years. Soil moisture was 48% lower in drought than ambient plots, but the invader largely offset drought effects. Additionally, temperature and light were lower and humidity higher in invaded plots. Field sites spanned up to a 10‐fold range in soil moisture and up to a 2.5‐fold range in light availability. Invaded and resident vegetation did not differentially mediate soil moisture, unlike in the garden experiment. Herbicide effectively removed invaded and resident vegetation, with removal having site‐specific effects on soil moisture and light availability. However, light was generally higher in invader‐removal than control plots, whereas resident removal had less effect on light, similar to the garden experiment. Invasion mitigated a constellation of abiotic conditions associated with drought stress in the garden experiment. In the field, where other factors co‐varied, these patterns did not emerge. Still, neither experiment suggested that drought and invasion will have synergistic negative effects on ecosystems, although invasion can limit light availability. Coupling factorial garden experiments with field experiments across environmental gradients will be effective for predicting how multiple stressors interact in natural systems.     

Contrasting physiological responsiveness of establishing trees and a C4 grass to rainfall events,intensified summer drought,and warming in oak savanna

Climate warming and drought may alter tree establishment in savannas through differential responses of tree seedlings and grass to intermittent rainfall events. We investigated leaf gas exchange responses of dominant post oak savanna tree (Quercus stellata and Juniperus virginiana) and grass (Schizachyrium scoparium, C₄ grass) species to summer rainfall events under an ambient and intensified summer drought scenario in factorial combination with warming (ambient, +1.5 °C) in both monoculture and tree‐grass mixtures. The three species differed in drought resistance and response of leaf gas exchange to rainfall events throughout the summer. S. scoparium experienced the greatest decrease in A_area (?56% and ?66% under normal and intensified drought, respectively) over the summer, followed by Q. stellata (?44%, ?64%), while J. virginiana showed increased A_area under normal drought (+13%) and a small decrease in A_area when exposed to intensified summer drought (?10%). Following individual rainfall events, mean increases in A_area were 90% for S. scoparium, 26% for J. virginiana and 22% for Q. stellata. The responsiveness of A_area of S. scoparium to rainfall events initially increased with the onset of drought, but decreased dramatically as summer drought progressed. For Q. stellata, A_area recovery decreased as drought progressed and with warming. In contrast, J. virginiana showed minimal fluctuations in A_area following rainfall events, in spite of declining water potential, and warming enhanced recovery. J. virginiana will likely gain an advantage over Q. stellata during establishment under future climatic scenarios. Additionally, the competitive advantage of C₄ grasses may be reduced relative to trees, because grasses will likely exist below a critical water stress threshold more often in a warmer, drier climate. Recognition of unique species responses to critical global change drivers in the presence of competition will improve predictions of grass–tree interactions and tree establishment in savannas in response to climate change.     

Disentangling water sources in a gypsum plant community. Gypsum crystallization water is a key source of water for shallow-rooted plants

Background and AimsGypsum drylands are widespread worldwide. In these arid ecosystems, the ability of different species to access different water sources during drought is a key determining factor of the composition of plant communities. Gypsum crystallization water could be a relevant source of water for shallow-rooted plants, but the segregation in the use of this source of water among plants remains unexplored. We analysed the principal water sources used by 20 species living in a gypsum hilltop, the effect of rooting depth and gypsum affinity, and the interaction of the plants with the soil beneath them.MethodsWe characterized the water stable isotope composition, δ ²H and δ ¹⁸O, of plant xylem water and related it to the free and gypsum crystallization water extracted from different depths throughout the soil profile and the groundwater, in both spring and summer. Bayesian isotope mixing models were used to estimate the contribution of water sources to plant xylem sap.Key ResultsIn spring, all species used free water from the top soil as the main source. In summer, there was segregation in water sources used by different species depending on their rooting depth, but not on their gypsum affinity. Gypsum crystallization water was the main source for most shallow-rooted species, whereas free water from 50 to 100 cm depth was the main source for deep-rooted species. We detected plant–soil interactions in spring, and indirect evidence of possible hydraulic lift by deep-rooted species in summer.ConclusionsPlants coexisting in gypsum communities segregate their hydrological niches according to their rooting depth. Crystallization water of gypsum represents an unaccounted for, vital source for most of the shallow-rooted species growing on gypsum drylands. Thus, crystallization water helps shallow-rooted species to endure arid conditions, which eventually accounts for the maintenance of high biodiversity in these specialized ecosystems.     

Shrub persistence and increased grass mortality in response to drought in dryland systems

Droughts in the southwest United States have led to major forest and grassland die‐off events in recent decades, suggesting plant community and ecosystem shifts are imminent as native perennial grass populations are replaced by shrub‐ and invasive plant‐dominated systems. These patterns are similar to those observed in arid and semiarid systems around the globe, but our ability to predict which species will experience increased drought‐induced mortality in response to climate change remains limited. We investigated meteorological drought‐induced mortality of nine dominant plant species in the Colorado Plateau Desert by experimentally imposing a year‐round 35% precipitation reduction for eight continuous years. We distributed experimental plots across numerous plant, soil, and parent material types, resulting in 40 distinct sites across a 4,500 km² region of the Colorado Plateau Desert. For all 8 years, we tracked c. 400 individual plants and evaluated mortality responses to treatments within and across species, and through time. We also examined the influence of abiotic and biotic site factors in driving mortality responses. Overall, high mortality trends were driven by dominant grass species, including Achnatherum hymenoides, Pleuraphis jamesii, and Sporobolus cryptandrus. Responses varied widely from year to year and dominant shrub species were generally resistant to meteorological drought, likely due to their ability to access deeper soil water. Importantly, mortality increased in the presence of invasive species regardless of treatment, and native plant die‐off occurred even under ambient conditions, suggesting that recent climate changes are already negatively impacting dominant species in these systems. Results from this long‐term drought experiment suggest major shifts in community composition and, as a result, ecosystem function. Patterns also show that, across multiple soil and plant community types, native perennial grass species may be replaced by shrubs and invasive annuals in the Colorado Plateau Desert.     

Invasion windows for a global legume invader are revealed after joint examination of abiotic and biotic filters

• Successful alien plant invasion is influenced by both climate change and plant–plant interactions. We estimate the single and interactive effects of competition and extreme weather events on the performance of the global legume invader Lupinus polyphyllus (Lindl.).
• In three experimental studies we assessed (i) the stress tolerance of seedling and adult L. polyphyllus plants against extreme weather events (drought, fluctuating precipitation, late frost), (ii) the competitive effects of L. polyphyllus on native grassland species and vice versa, and (iii) the interactive effects of extreme weather events and competition on the performance of L. polyphyllus.
• Drought reduced growth and led to early senescence of L. polyphyllus but did not reduce adult survival. Fluctuating precipitation events and late frost reduced the length of inflorescences. Under control conditions, interspecific competition reduced photosynthetic activity and growth of L. polyphyllus. When subjected to competition during drought, L. polyphyllus conserved water while simultaneously maintaining high assimilation rates, demonstrating increased water use efficiency. Meanwhile, native species had reduced performance under drought.
• In summary, the invader gained an advantage under drought conditions through a smaller reduction in performance relative to its native competitors but was competitively inferior under control conditions. This provides evidence for a possible invasion window for this species. While regions of high elevation or latitude with regular severe late frost events might remain inaccessible for L. polyphyllus, further spread across Europe seems probable as the predicted increase in drought events may favour this non‐native legume over native species.

     

The combined effects of a long‐term experimental drought and an extreme drought on the use of plant‐water sources in a Mediterranean forest

Vegetation in water‐limited ecosystems relies strongly on access to deep water reserves to withstand dry periods. Most of these ecosystems have shallow soils over deep groundwater reserves. Understanding the functioning and functional plasticity of species‐specific root systems and the patterns of or differences in the use of water sources under more frequent or intense droughts is therefore necessary to properly predict the responses of seasonally dry ecosystems to future climate. We used stable isotopes to investigate the seasonal patterns of water uptake by a sclerophyll forest on sloped terrain with shallow soils. We assessed the effect of a long‐term experimental drought (12 years) and the added impact of an extreme natural drought that produced widespread tree mortality and crown defoliation. The dominant species, Quercus ilex, Arbutus unedo and Phillyrea latifolia, all have dimorphic root systems enabling them to access different water sources in space and time. The plants extracted water mainly from the soil in the cold and wet seasons but increased their use of groundwater during the summer drought. Interestingly, the plants subjected to the long‐term experimental drought shifted water uptake toward deeper (10–35 cm) soil layers during the wet season and reduced groundwater uptake in summer, indicating plasticity in the functional distribution of fine roots that dampened the effect of our experimental drought over the long term. An extreme drought in 2011, however, further reduced the contribution of deep soil layers and groundwater to transpiration, which resulted in greater crown defoliation in the drought‐affected plants. This study suggests that extreme droughts aggravate moderate but persistent drier conditions (simulated by our manipulation) and may lead to the depletion of water from groundwater reservoirs and weathered bedrock, threatening the preservation of these Mediterranean ecosystems in their current structures and compositions.     

Effects of climate‐induced increases in summer drought on riparian plant species: a meta‐analysis

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Secondary invasion: When invasion success is contingent on other invaders altering the properties of recipient ecosystems

Positive interactions between exotic species may increase ecosystem‐level impacts and potentially facilitate the entry and spread of other exotic species. Invader‐facilitated invasion success—”secondary invasion”—is a key conceptual aspect of the well‐known invasional meltdown hypothesis, but remains poorly defined and empirically underexplored. Drawing from heuristic models and published empirical studies, we explore this form of “secondary invasion” and discuss the phenomenon within the recognized conceptual framework of the determinants of invasion success. The term “secondary invasion” has been used haphazardly in the literature to refer to multiple invasion phenomena, most of which have other more accepted titles. Our usage of the term secondary invasion is akin to “invader‐facilitated invasion,” which we define as the phenomenon in which the invasion success of one exotic species is contingent on the presence, influence, and impacts of one or more other exotic species. We present case studies of secondary invasion whereby primary invaders facilitate the entry or establishment of exotic species into communities where they were previously excluded from becoming invasive. Our synthesis, discussion, and conceptual framework of this type of secondary invasion provides a useful reference to better explain how invasive species can alter key properties of recipient ecosystems that can ultimately determine the invasion success of other species. This study increases our appreciation for complex interactions following invasion and highlights the impacts of invasive species themselves as possible determinants of invasion success. We anticipate that highlighting “secondary invasion” in this way will enable studies reporting similar phenomena to be identified and linked through consistent terminology.     

Disturbance governs dominance of an invasive forb in a temporary wetland

Dominance of invasive species is often assumed to be due to a superior ability to acquire resources. However, dominance in plant communities can arise through multiple interacting mechanisms, including disturbance. Inter-specific competition can be strongly affected by abiotic conditions, which can determine the outcome of competitive interactions. We evaluated competition and disturbance as mechanisms governing dominance of Phyla canescens (hereafter lippia), an invasive perennial forb from South America, in Paspalum distichum (perennial grass, hereafter water couch) meadows in floodplain wetlands of eastern Australia. Water couch meadows (in the study area) are listed under the Ramsar Convention due to their significance as habitat for migratory waterbirds. In the field, we monitored patterns of vegetation boundaries between the two species in response to flooding. Under controlled glasshouse conditions, we explored competitive interactions between the native water couch and lippia subject to different soil moisture/inundation regimes. We did this using a pairwise factorial glasshouse experiment that manipulated neighbor density (9 treatments) and soil moisture/inundation (4 treatments). In the field trial, inundation increased the cover of water couch. Under more controlled conditions, the invader had a competitive effect on the native species only under dry soil conditions, and was strongly inhibited by inundation. This suggests that dry conditions favor the growth of the invader and wetter (more historical) conditions favor the native grass. In this system, invader dominance is governed by altered disturbance regimes which give the invader a competitive advantage over the native species.     

Tolerance to drought and flooding events provides a competitive advantage for an invasive over a native plant species

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Ecosystem functioning impacts of the invasive seaweed Sargassum muticum (Fucales,Phaeophyceae)

Ongoing changes in natural diversity due to anthropogenic activities can alter ecosystem functioning. Particular attention has been given to research on biodiversity loss and how those changes can affect the functioning of ecosystems, and, by extension, human welfare. Few studies, however, have addressed how increased diversity due to establishment of nonindigenous species (NIS) may affect ecosystem function in the recipient communities. Marine algae have a highly important role in sustaining nearshore marine ecosystems and are considered a significant component of marine bioinvasions. Here, we examined the patterns of respiration and light‐use efficiency across macroalgal assemblages with different levels of species richness and evenness. Additionally, we compared our results between native and invaded macroalgal assemblages, using the invasive brown macroalga Sargassum muticum (Yendo) Fensholt as a model species. Results showed that the presence of the invader increased the rates of respiration and production, most likely as a result of the high biomass of the invader. This effect disappeared when S. muticum lost most of its biomass after senescence. Moreover, predictability–diversity relationships of macroalgal assemblages varied between native and invaded assemblages. Hence, the introduction of high‐impact invasive species may trigger major changes in ecosystem functioning. The impact of S. muticum may be related to its greater biomass in the invaded assemblages, although species interactions and seasonality influenced the magnitude of the impact.     

High competitiveness of a resource demanding invasive acacia under low resource supply

Mechanisms controlling the successful invasion of resource demanding species into low-resource environments are still poorly understood. Well-adapted native species are often considered superior competitors under stressful conditions. Here we investigate the competitive ability of the resource demanding alien Acacia longifolia, which invades nutrient-poor Mediterranean sand dunes such as in coastal areas of Portugal. We explore the hypothesis that drought may limit invasion in a factorial competition experiment of the alien invasive versus two native species of different functional groups (Halimium halimifolium, Pinus pinea), under well-watered and drought conditions. Changes in biomass, allocation pattern, and N-uptake-efficiency (via ¹⁵N-labeling) indicated a marked drought sensitivity of the invader. However, highly efficient drought adaptations of the native species did not provide a competitive advantage under water limiting conditions. The competitive strength of H. halimifolium towards the alien invader under well-watered conditions turned into a positive interaction between both species under drought. Further, low resource utilization by native species benefited A. longifolia by permitting continued high nitrogen uptake under drought. Hence, the N-fixing invader expresses low plasticity by continuous high resource utilization, even under low resource conditions. The introduction of novel traits into a community like N-fixation and high resource use may promote A. longifolia invasiveness through changes in the physical environment, i.e., the water and nutrient cycle of the invaded sand dune system, thereby potentially disrupting the co-evolved interactions within the native plant community.     

Environmentally‐contingent behaviour of invasive plants as drivers or passengers

Invasive plants are often regarded as drivers that actively reduce diversity and alter ecosystem processes such as succession. Alternatively, invaders may be passengers that simply colonize openings produced by anthropogenic disturbance and are present only temporarily. Here we test whether the behaviour of invasive species as drivers or passengers is contingent on disturbance and nutrient availability. We created twelve experimental environments (three levels of annual disturbance × four levels of nitrogen availability) for 18 years in a grassland at the northern edge of the North American Great Plains. Out of 19 invasive species initially present, two perennials (Bromus inermis, a grass, and Cirsium arvense, a forb) acted as drivers, maintaining or increasing dominance, maintaining low species richness, and forming an invader‐ dominated successional sequence. Behaviour as drivers was environmentally contingent: Bromus behaved as a driver only in less disturbed environments, and the tendency of Cirsium to behave as a driver increased significantly with both disturbance and nitrogen availability. Most invasive species (90%), however, consistently behaved as passengers, disappearing or becoming rare. The importance of disturbance and fertility for starting invasions is well‐known, but our study shows that these factors also contribute to the behaviour of some invaders as drivers. The emergence of drivers and invader‐dominated successional sequences suggests that, as rates of invasion, disturbance and eutrophication continue to increase with human activity, invasive species that act as drivers may form low‐diversity communities that persist for decades.     

The Influence of Abiotic Factors and Temporal Variation on Local Invasion Patterns of the Argentine Ant (Linepithema humile)

The Argentine ant is a common pest of mediterranean-type ecosystems worldwide, causing widespread extirpation of many native ant species. This study examines spatial and temporal patterns of invasion at a local scale, investigates the effects of this invader on native ant communities and assesses causal mechanisms for these spatial, temporal and numerical patterns, particularly in relation to local climate. Argentine ant workers were more active in the drier months of summer and fall, and abundance was correlated with cumulative precipitation the previous winter. Argentine ant worker abundance greatly increased in conjunction with EI Niño events. No similar relationship existed for native ants. This invasive ant affected native ant abundances differently by species: some species were able to tolerate its presence temporarily. Overall, this study provides a framework of factors to consider when devising control strategies for the invasive Argentine ant.     

Competition between Native Hawaiian Plants and the Invasive Grass Megathyrsus maximus: Implications of Functional Diversity for Ecological Restoration

Biodiversity loss is a global crisis, due primarily to habitat destruction and widespread nonnative invasions. Invasive grasses are particularly problematic in many tropical ecosystems, where they possess traits that promote their persistence and can drastically alter native plant communities. We explored the ecophysiological basis for restoring native Hawaiian dryland ecosystems currently dominated by the nonnative invasive grass Megathyrsus maximus (guinea grass) in a garden experiment. Three native species—Myoporum sandwicense (naio; canopy tree), Dodonaea viscosa (aalii; shrub), and Plumbago zeylanica (iliee; groundcover)—were grown with M. maximus at three levels of native functional diversity (one, two, or three species) while holding overall plant density constant. We tested which individual and functional combinations of native species were more productive and best suppressed M. maximus growth and reproduction. Megathyrsus maximus had 39–94% higher maximum photosynthetic rates (A_max) than native species and increasing native functional diversity did not affect M. maximus A_max. Aboveground, belowground, and total biomass of M. maximus varied with functional diversity, although intraspecific competition reduced growth as much as interspecific competition. Reproductive tiller production by M. maximus decreased significantly when planted with any of the native species and with increasing native functional diversity. These results indicate that high native functional diversity in an ecological restoration setting may aid in the control of a dominant invasive grass and the reintroduction of diverse native species. Recommendations for restoring degraded nonnative grasslands in Hawaii and throughout the tropics include selection of native species that are ecophysiologically competitive and have high functional diversity.