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  1. Abstract

    Plant traits can be helpful for understanding grassland ecosystem responses to climate extremes, such as severe drought. However, intercontinental comparisons of how drought affects plant functional traits and ecosystem functioning are rare. The Extreme Drought in Grasslands experiment (EDGE) was established across the major grassland types in East Asia and North America (six sites on each continent) to measure variability in grassland ecosystem sensitivity to extreme, prolonged drought. At all sites, we quantified community‐weighted mean functional composition and functional diversity of two leaf economic traits, specific leaf area and leaf nitrogen content, in response to drought. We found that experimental drought significantly increased community‐weighted means of specific leaf area and leaf nitrogen content at all North American sites and at the wetter East Asian sites, but drought decreased community‐weighted means of these traits at moderate to dry East Asian sites. Drought significantly decreased functional richness but increased functional evenness and dispersion at most East Asian and North American sites. Ecosystem drought sensitivity (percentage reduction in aboveground net primary productivity) positively correlated with community‐weighted means of specific leaf area and leaf nitrogen content and negatively correlated with functional diversity (i.e., richness) on an intercontinental scale, but results differed within regions. These findings highlight both broad generalities but also unique responses to drought of community‐weighted trait means as well as their functional diversity across grassland ecosystems.

     
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    Free, publicly-accessible full text available February 1, 2025
  2. Abstract

    Plant traits are useful proxies of plant strategies and can influence community and ecosystem responses to climate extremes, such as severe drought. Few studies, however, have investigated both the immediate and lagged effects of drought on community‐weighted mean (CWM) plant traits, with even less research on the relative roles of interspecific vs. intraspecific trait variability in such responses.

    We experimentally reduced growing season precipitation by 66% in two cold‐semi‐arid grassland sites in northern China for four consecutive years to explore the drought resistance of CWM traits as well as their recovery 2 years following the drought. In addition, we isolated the effects of both interspecific and intraspecific trait variability on shifts in CWM traits.

    At both sites, we observed significant effects of drought on interspecific and intraspecific trait variability which, in some cases, led to significant changes in CWM traits. For example, drought led to reduced CWM plant height and leaf phosphorous content, but increased leaf carbon content at both sites, with responses primarily due to intraspecific trait shifts. Surprisingly, these CWM traits recovered completely 2 years after the extreme drought. Intraspecific trait variability influenced CWM traits via both positive and negative covariation with interspecific trait variability during drought and recovery phases.

    These findings highlight the important role of interspecific and intraspecific trait variability in driving the response and recovery of CWM traits following extreme, prolonged drought.

    Read the freePlain Language Summaryfor this article on the Journal blog.

     
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  3. Hector, Andrew (Ed.)
  4. Abstract

    Asexual reproduction plays a fundamental role in the structure, dynamics and persistence of perennial grasslands. Thus, assessing how asexual reproductive traits of plant communities respond to drought may be key for understanding grassland resistance to drought and recovery following drought.

    Here, we quantified three asexual reproductive traits (i.e. above‐ground tiller abundance, below‐ground bud abundance and the ratio of tillers to buds) during a 4‐year severe drought and a 2‐year drought recovery period in four grasslands that spanned an aridity gradient in northern China. We also assessed the relationship between these traits and the resistance and recovery of above‐ground net primary productivity (ANPP).

    We found that drought had limited and largely inconsistent effects on asexual reproduction among drought and recovery years and grasslands overall. Drought increased tiller abundance in the first treatment year and reduced bud banks by the fourth treatment year across grasslands. However, neither of the three asexual reproductive traits were correlated with drought resistance of ANPP. Drought legacies differed among the four grasslands with positive, negative and no legacies evident for the three asexual reproductive traits, and no clear relationship with aridity. Bud banks and tiller to bud ratio decreased and increased, respectively, in the first recovery year, but not in the second recovery year. In contrast to drought resistance, community bud abundance was strongly related to recovery, such that communities with higher bud abundance had greater ANPP recovery following drought.

    Synthesis. These results suggest that asexual reproductive traits may be important drivers of ecosystem recovery after drought, but that variable responses of these asexual reproduction traits during drought complicates predictions of overall grassland responses.

     
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  5. Abstract

    Recurrent droughts are an inevitable consequence of climate change, yet how grasslands respond to such events is unclear. We conducted a 6‐year rainfall manipulation experiment in a semiarid grassland that consisted of an initial 2‐year drought (2015–2016), followed by a recovery period (2017–2018) and, finally, a second 2‐year drought (2019–2020). In each year, we estimated aboveground net primary productivity (ANPP), species richness, community‐weighted mean (CWM) plant traits, and several indices of functional diversity. The initial drought led to reduced ANPP, which was primarily driven by limited growth of forbs in the first year and grasses in the second year. Total ANPP completely recovered as the rapid recovery of grass productivity compensated for the slow recovery of forb productivity. The subsequent drought led to a greater reduction in total ANPP than the initial drought due to the greater decline of both grass and forb productivity. The structural equation models revealed that soil moisture influenced ANPP responses directly during the initial drought, and indirectly during the subsequent drought by lowering functional diversity, which resulted in reduced total ANPP. Additionally, ANPP was positively influenced by CWM plant height and leaf nitrogen during the recovery period and recurrent drought, respectively. Overall, the greater impact of the second drought on ecosystem function than the initial drought, as well as the underlying differential mechanism, underscores the need for an understanding of how increased drought frequency may alter semiarid grassland functioning.

     
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  6. Abstract

    Grasslands are expected to experience droughts of unprecedented frequency and magnitude in the future. Characterizing grassland responses and recovery from drought is therefore critical to predict the vulnerability of grassland ecosystems to climate change. Most previous studies have focused on ecosystem responses during drought while investigations of post‐drought recovery are rare. Few studies have used functional traits, and in particular bud or clonal traits, to explore the mechanisms underlying grassland responses to and recovery from drought.

    To address this issue, we experimentally imposed a four‐year drought in a C3‐dominated grassland in northeastern China and monitored recovery for 3 years post‐drought. We investigated the immediate and legacy effects of drought on total above‐ground net primary productivity (ANPP), ANPP of functional groups (rhizomatous grasses, bunch grasses and forbs), and how the legacy effects were driven by plant species diversity, clonal traits and vegetative traits.

    We found that drought progressively reduced total ANPP over the 4‐year period. The reductions in total ANPP in the first and third drought years were caused by the decrease in ANPP of bunch grasses only, while that of the second year was caused by declines in ANPP of bunch grasses and forbs, and the fourth year decline was linked to all three functional groups. The post‐drought recovery of ANPP, which occurred despite the continued loss of plant species diversity, was mainly driven by rapid recovery of rhizomatous and bunch grasses, which compensated for the slow response by forbs. The rapid post‐drought recovery of these grasses can be attributed to their relatively large, intact bud and shoot densities post‐drought, as well as the recovery of plant height and specific leaf area. The rapid recovery of grasses possibly restricted the growth and distribution of forbs, resulting in reduced forb ANPP and, consequently, lower species diversity during the recovery period.

    Synthesis. These results highlight the potential for positive legacy effects of drought on ANPP as well as the important and complementary roles of plant reproductive and vegetative traits in mediating ecosystem recovery from drought in a C3‐dominated grassland.

     
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  7. Abstract

    Nutrient enrichment can simultaneously increase and destabilise plant biomass production, with co‐limitation by multiple nutrients potentially intensifying these effects. Here, we test how factorial additions of nitrogen (N), phosphorus (P) and potassium with essential nutrients (K+) affect the stability (mean/standard deviation) of aboveground biomass in 34 grasslands over 7 years. Destabilisation with fertilisation was prevalent but was driven by single nutrients, not synergistic nutrient interactions. On average, N‐based treatments increased mean biomass production by 21–51% but increased its standard deviation by 40–68% and so consistently reduced stability. Adding P increased interannual variability and reduced stability without altering mean biomass, while K+ had no general effects. Declines in stability were largest in the most nutrient‐limited grasslands, or where nutrients reduced species richness or intensified species synchrony. We show that nutrients can differentially impact the stability of biomass production, with N and P in particular disproportionately increasing its interannual variability.

     
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