Gradual changes in range size accompany long‐term trends in species richness

Species richness has long been used as an indicator of ecosystem functioning and health. Global richness is declining, but it is unclear whether sub‐global trends differ. Regional trends are especially understudied, with most focused on island regions where richness is strongly impacted by novel colonisations. We addressed this knowledge gap by testing for multi‐decade trends in species richness in nine open marine regions around North America (197 region‐years) while accounting for imperfect observations and grounding our findings in species‐level range dynamics. We found positive richness trends in eight of nine regions, four of which were statistically significant. Species' range sizes generally contracted pre‐extinction and expanded post‐colonisation, but the ranges of transient species expanded over the long‐term, slowly increasing their regional retention and driving increasing richness. These results provide more evidence that sub‐global richness trends are stable or increasing, and highlight the utility of range size for understanding richness dynamics.     

Species colonisation,not competitive exclusion,drives community overdispersion over long‐term succession

Ecological communities often transition from phylogenetic and functional clustering to overdispersion over succession as judged by space‐for‐time substitution studies. Such a pattern has been generally attributed to the increase in competitive exclusion of closely related species with similar traits through time, although colonisation and extinction have rarely been examined. Using 44 years of uninterrupted old‐field succession in New Jersey, USA, we confirmed that phylogenetic and functional clustering decreased as succession unfolded, but the transition was largely driven by colonisation. Early colonists were closely related and functionally similar to residents, while later colonists became less similar to the species present. Extirpated species were generally more distantly related to residents than by chance, or exhibited random phylogenetic/functional patterns, and their relatedness to residents was not associated with time. These results provide direct evidence that the colonisation of distant relatives, rather than extinction of close relatives, drives phylogenetic and functional overdispersion over succession.     

Top‐down control of rare species abundances by native ungulates in a grassland restoration

Ecological restorations are predicted to increase in species diversity over time until they reach reference levels. However, chronosequence studies in grasslands often show that diversity peaks after the first few years and then declines over time as grasses become more dominant. We addressed whether bison grazing and seed additions could prevent this decline in diversity. Exclosures that prevented bison grazing were compared with grazed plots over 4 years, and seed additions were conducted inside and outside exclosures to test for seed and microsite limitations. A previous study conducted 4‐months post seeding found that local species richness was primarily seed limited, but that grazing could sometimes increase seedling emergence. Here, we tested whether increased seedling emergence led to longer‐term increases in the species diversity of the plant community. We found that the seed addition effect grew smaller and the grazing effect grew stronger over time, and that seed additions affected the abundance of added species only when plots were grazed. Grazed plots had higher species diversity and lower biomass and litter buildup compared to non‐grazed plots. Our results suggest that moderate grazing by bison or management that mimics grazing can maintain diversity in grass‐dominated situations. Our results also emphasize the need to follow seed additions over several years to assess correctly whether seed limitation exists.     

Links between soil microbial communities and plant traits in a species‐rich grassland under long‐term climate change

Climate change can influence soil microorganisms directly by altering their growth and activity but also indirectly via effects on the vegetation, which modifies the availability of resources. Direct impacts of climate change on soil microorganisms can occur rapidly, whereas indirect effects mediated by shifts in plant community composition are not immediately apparent and likely to increase over time. We used molecular fingerprinting of bacterial and fungal communities in the soil to investigate the effects of 17 years of temperature and rainfall manipulations in a species‐rich grassland near Buxton, UK. We compared shifts in microbial community structure to changes in plant species composition and key plant traits across 78 microsites within plots subjected to winter heating, rainfall supplementation, or summer drought. We observed marked shifts in soil fungal and bacterial community structure in response to chronic summer drought. Importantly, although dominant microbial taxa were largely unaffected by drought, there were substantial changes in the abundances of subordinate fungal and bacterial taxa. In contrast to short‐term studies that report high resistance of soil fungi to drought, we observed substantial losses of fungal taxa in the summer drought treatments. There was moderate concordance between soil microbial communities and plant species composition within microsites. Vector fitting of community‐weighted mean plant traits to ordinations of soil bacterial and fungal communities showed that shifts in soil microbial community structure were related to plant traits representing the quality of resources available to soil microorganisms: the construction cost of leaf material, foliar carbon‐to‐nitrogen ratios, and leaf dry matter content. Thus, our study provides evidence that climate change could affect soil microbial communities indirectly via changes in plant inputs and highlights the importance of considering long‐term climate change effects, especially in nutrient‐poor systems with slow‐growing vegetation.     

Long‐term phosphite application maintains species assemblages,richness and structure of plant communities invaded by Phytophthora cinnamomi

The impact of the plant pathogen Phytophthora cinnamomi and the fungicide phosphite on species assemblages, richness, abundance and vegetation structure was quantified at three sites in Kwongkan communities in the Southwest Australian Floristic Region. Healthy and diseased vegetation treated with phosphite over 7–16 years was compared with non‐treated healthy and diseased vegetation. After site differences, disease had the greatest effect on species assemblages, species richness and richness within families. Disease significantly reduced cover in the upper and lower shrub layers and increased sedge and bare ground cover. Seventeen of 21 species assessed from the families Ericaceae, Fabaceae, Myrtaceae and Proteaceae were significantly less abundant in non‐treated diseased vegetation. In diseased habitats, phosphite treatment significantly reduced the loss of shrub cover and reduced bare ground and sedge cover. In multivariate analysis of species assemblages, phosphite‐treated diseased plots grouped more closely with healthy plots. Seven of 17 susceptible species were significantly more abundant in phosphite‐treated diseased plots compared with diseased non‐treated plots. The abundance of seven of 10 Phytophthora‐susceptible species was significantly higher along transects in phosphite‐treated vegetation. Comparison of the floristics of healthy non‐treated with healthy‐treated plots showed no significant differences in species assemblages. Of 21 species assessed, three increased in abundance and only one decreased significantly in phosphite‐treated healthy plots. In three Kwongkan communities of the SWAFR, P. cinnamomi had a profound impact on species assemblages, richness, abundance and vegetation structure. There was no evidence of adverse effects of phosphite treatment on phosphorus‐sensitive species, even after fire. Treatment with phosphite enhanced the survival of key susceptible species and mitigated disease‐mediated changes in vegetation structure. In the absence of alternative methods of control in native communities, phosphite will continue to play an important role in the protection of high priority species and communities at risk of extinction due to P. cinnamomi.     

Effects of plant diversity,N fertilization,and elevated carbon dioxide on grassland soil N cycling in a long‐term experiment

The effects of global environmental changes on soil nitrogen (N) pools and fluxes have consequences for ecosystem functions such as plant productivity and N retention. In a 13‐year grassland experiment, we evaluated how elevated atmospheric carbon dioxide (CO₂), N fertilization, and plant species richness alter soil N cycling. We focused on soil inorganic N pools, including ammonium and nitrate, and two N fluxes, net N mineralization and net nitrification. In contrast with existing hypotheses, such as progressive N limitation, and with observations from other, often shorter, studies, elevated CO₂ had relatively static and small, or insignificant, effects on soil inorganic N pools and fluxes. Nitrogen fertilization had inconsistent effects on soil N transformations, but increased soil nitrate and ammonium concentrations. Plant species richness had increasingly positive effects on soil N transformations over time, likely because in diverse subplots the concentrations of N in roots increased over time. Species richness also had increasingly positive effects on concentrations of ammonium in soil, perhaps because more carbon accumulated in soils of diverse subplots, providing exchange sites for ammonium. By contrast, subplots planted with 16 species had lower soil nitrate concentrations than less diverse subplots, especially when fertilized, probably due to greater N uptake capacity of subplots with 16 species. Monocultures of different plant functional types had distinct effects on N transformations and nitrate concentrations, such that not all monocultures differed from diverse subplots in the same manner. The first few years of data would not have adequately forecast the effects of N fertilization and diversity on soil N cycling in later years; therefore, the dearth of long‐term manipulations of plant species richness and N inputs is a hindrance to forecasting the state of the soil N cycle and ecosystem functions in extant plant communities.     

Interannual variability in responses of belowground net primary productivity (NPP) and NPP partitioning to long‐term warming and clipping in a tallgrass prairie

The dynamics of belowground net primary productivity (BNPP) is of fundamental importance in understanding carbon (C) allocation and storage in grasslands. However, our knowledge of the interannual variability in response of BNPP to ongoing global warming is limited. In this study, we explored temporal responses of BNPP and net primary productivity (NPP) partitioning to warming and clipping in a tallgrass prairie in Oklahoma, USA. Infrared heaters were used to elevate soil temperature by approximately 2 °C since November 1999. Annual clipping was to mimic hay harvest. On average from 2005 to 2009, warming increased BNPP by 41.89% in the unclipped subplots and 66.93% in the clipped subplots, with significant increase observed in wet years. Clipping also had significant positive impact on BNPP, which was mostly found under warming. Overall, f_BNPP, the fraction of BNPP to NPP, increased under both warming and clipping treatments, more in dry years. Water availability (either precipitation or soil moisture) was the most limiting factor for both BNPP and f_BNPP. It strongly dominated the interannual variability in NPP, f_BNPP, and their responses to warming and clipping. Our results suggest that water availability regulates tallgrass prairie's responses to warming and land use change, which may eventually influence the global C cycle. With increasing variability in future precipitation patterns, warming effects on the vegetation in this region may become less predictable.     

Effects of long‐term cutting in a grassland system: perspectives for restoration of plant communities on nutrient‐poor soils

Abstract. The upper courses of brook valley systems harbour Nardo‐Galion saxatilis communities characteristic of oligo‐trophic soils under low‐intensity farming. Most of these communities have disappeared under intensified farming i.e. application of fertilizers. We studied the possibilities of restoration i.e. re‐establishment of the former plant community by adopting various cutting regimes after the cessation of fertilization in 1972. The various cutting regimes revealed different effects after 25 yr. Regimes with cutting every second year with or without removal of the swath, and complete abandonment deviated from the other regimes that included annual haymaking with different frequency and timing. The latter group of cutting regimes came closer to the community of an adjacent field where fertilization stopped in 1967. This field in turn harboured several Nardo‐Galion species after 25 yr of annual cutting, and showed more resemblance with a local reference community (at a distance of 500 m) that had not been fertilized since the 1940s. The local reference still does not match poorly developed Nardo‐Galion saxatilis communities found in the region of ca. 50 km around the study area, and is far from well developed Nardo‐Galion communities in the same region. The study site still harbours several species characteristic of eutrophic soil and few species characteristic of oligotrophic soil after 25 yr of annual cutting and removal of the swath. The soil seed bank harbours only few target species. Although species characteristic of oligotrophic soil are present in an adjacent field and Nardo‐Galion saxatilis species occur at 500 m, they have not (yet) established in the target area.     

Long‐term population trends in three grassland insect groups: a comparative analysis of 1951 and 2009

Development of farming practices has caused drastic changes in European agricultural landscapes during the past 50 years. As a consequence of these changes, insect diversity is widely expected to decline. We performed a comparative analysis with long‐term data of three insect groups: Auchenorrhyncha, Heteroptera and Orthoptera. In 2009, we revisited nine grassland sites in northern Germany that were originally sampled in 1951 using the same techniques and during a similar time frame. We found that the insect community exhibited no consistent trends between years. Species richness of Auchenorrhyncha and Heteroptera increased on plot level as well as on landscape level but remained unchanged for Orthoptera. Abundance of Auchenorrhyncha and Orthoptera significantly decreased, while Heteroptera increased. There is a strong trend towards homogeneity in community composition for Heteroptera and a weak one for Auchenorrhyncha. The frequency and abundance of species preferring disturbed and/or eutrophic habitats increased, whereas the number of species preferring low‐productive habitats declined. This trend is especially pronounced in Auchenorrhyncha. Generalistic species were more abundant in relative proportions as well as in absolute numbers. We hypothesize that these trends arise from alterations of Central European landscapes because of agricultural intensification over the last several decades.     

Press–pulse interactions: effects of warming,N deposition,altered winter precipitation,and fire on desert grassland community structure and dynamics

Global environmental change is altering temperature, precipitation patterns, resource availability, and disturbance regimes. Theory predicts that ecological presses will interact with pulse events to alter ecosystem structure and function. In 2006, we established a long‐term, multifactor global change experiment to determine the interactive effects of nighttime warming, increased atmospheric nitrogen (N) deposition, and increased winter precipitation on plant community structure and aboveground net primary production (ANPP) in a northern Chihuahuan Desert grassland. In 2009, a lightning‐caused wildfire burned through the experiment. Here, we report on the interactive effects of these global change drivers on pre‐ and postfire grassland community structure and ANPP. Our nighttime warming treatment increased winter nighttime air temperatures by an average of 1.1 °C and summer nighttime air temperature by 1.5 °C. Soil N availability was 2.5 times higher in fertilized compared with control plots. Average soil volumetric water content (VWC) in winter was slightly but significantly higher (13.0% vs. 11.0%) in plots receiving added winter rain relative to controls, and VWC was slightly higher in warmed (14.5%) compared with control (13.5%) plots during the growing season even though surface soil temperatures were significantly higher in warmed plots. Despite these significant treatment effects, ANPP and plant community structure were highly resistant to these global change drivers prior to the fire. Burning reduced the cover of the dominant grasses by more than 75%. Following the fire, forb species richness and biomass increased significantly, particularly in warmed, fertilized plots that received additional winter precipitation. Thus, although unburned grassland showed little initial response to multiple ecological presses, our results demonstrate how a single pulse disturbance can interact with chronic alterations in resource availability to increase ecosystem sensitivity to multiple drivers of global environmental change.     

Post‐fire carbon dynamics in the tropical peat swamp forests of Brunei reveal long‐term elevated CH4 flux

Tropical peatlands hold about 15%–19% of the global peat carbon (C) pool of which 77% is stored in the peat swamp forests (PSFs) of Southeast Asia. Nonetheless, these PSFs have been drained, exploited for timber and land for agriculture, leading to frequent fires in the region. The physico‐chemical characteristics of peat, as well as the hydrology of PSFs are affected after a fire, during which the ecosystem can act as a C source for decades, as C emissions to the atmosphere exceed photosynthesis. In this work, we studied the longer‐term impact of fires on C cycling in tropical PSFs, hence we quantified the magnitude and patterns of C loss (CO₂, CH₄ and dissolved organic carbon) and soil‐water quality characteristics in an intact and a degraded burnt PSF in Brunei Darussalam affected by seven fires over the last 40 years. We used natural tracers such as ¹⁴C to investigate the age and sources of C contributing to ecosystem respiration (R_eco) and CH₄, while we continuously monitored soil temperature and water table (WT) level from June 2017 to January 2019. Our results showed a major difference in the physico‐chemical parameters, which in turn affected C dynamics, especially CH₄. Methane effluxes were higher in fire‐affected areas (7.8 ± 2.2 mg CH₄ m^?2 hr^?1) compared to the intact PSF (4.0 ± 2.0 mg CH₄ m^?2 hr^?1) due to prolonged higher WT and more optimal methanogenesis conditions. On the other hand, we did not find significant differences in R_eco between burnt (432 ± 83 mg CO₂ m^?2 hr^?1) and intact PSF (359 ± 76 mg CO₂ m^?2 hr^?1). Radiocarbon analysis showed overall no significant difference between intact and burnt PSF with a modern signature for both CO₂ and CH₄ fluxes implying a microbial preference for the more labile C fraction in the peat matrix.     

Experimental fire increases soil carbon dioxide efflux in a grassland long‐term multifactor global change experiment

Numerous studies have demonstrated that soil respiration rates increase under experimental warming, although the long‐term, multiyear dynamics of this feedback are not well constrained. Less is known about the effects of single, punctuated events in combination with other longer‐duration anthropogenic influences on the dynamics of soil carbon (C) loss. In 2012 and 2013, we assessed the effects of decadal‐scale anthropogenic global change – warming, increased nitrogen (N) deposition, elevated carbon dioxide (CO₂), and increased precipitation – on soil respiration rates in an annual‐dominated Mediterranean grassland. We also investigated how controlled fire and an artificial wet‐up event, in combination with exposure to the longer‐duration anthropogenic global change factors, influenced the dynamics of C cycling in this system. Decade‐duration surface soil warming (1–2 °C) had no effect on soil respiration rates, while +N addition and elevated CO₂ concentrations increased growing‐season soil CO₂ efflux rates by increasing annual aboveground net primary production (NPP) and belowground fine root production, respectively. Low‐intensity experimental fire significantly elevated soil CO₂ efflux rates in the next growing season. Based on mixed‐effects modeling and structural equation modeling, low‐intensity fire increased growing‐season soil respiration rates through a combination of three mechanisms: large increases in soil temperature (3–5 °C), significant increases in fine root production, and elevated aboveground NPP. Our study shows that in ecosystems where soil respiration has acclimated to moderate warming, further increases in soil temperature can stimulate greater soil CO₂ efflux. We also demonstrate that punctuated short‐duration events such as fire can influence soil C dynamics with implications for both the parameterization of earth system models (ESMs) and the implementation of climate change mitigation policies that involve land‐sector C accounting.