Habitat filtering masks the detection of phylogenetic and functional density dependence on seedling survival across seasons in a subtropical forest, Southwestern China

The neighborhood effect has become an important framework with which to study the mechanisms that maintain the coexistence of tree species. Phylogenetic relatedness among neighboring plants directly affects species coexistence and the maintenance of tree diversity. And some studies have reported that seedling performance is negatively correlated with phylogenetic relatedness, which termed phylogenetic negative density dependence. Soil-borne fungal pathogens affected seedling performance of phylogenetically related host species, i.e., phylogenetic Janzen–Connell effect. Seedlings may be particularly vulnerable to habitat and neighbor characteristics. Although previous studies have demonstrated the influence of neighborhood effects, phylogenetic relatedness, and habitat filtering on seedling survival, growth, and mortality, the effect of variation in these factors on seedling abundance remains unclear. To address this question, we used a 4-ha (200 m × 200 m) and monitored four-year (2020–2023) seedling dataset from a mid-montane humid evergreen broad-leaved subtropical forest in the Gaoligong Mountains, Yunnan, Southwestern China, and which consisted of 916 seedlings belonging to 56 species. The results of generalized linear mixed models showed no significant effect of conspecific adult neighbors on seedling abundance at any of the intervals evaluated. In contrast, we found evidence of phylogenetic distance density dependence in the forests of the Gaoligong Mountains. Specifically, there was a significant positive effect of the relative average phylogenetic distance between heterospecific adult neighbors and focal seedlings on focal seedling abundance in 2020; however, the relative average phylogenetic distance between heterospecific seedling neighbors and focal seedlings had a significant negative effect on seedling abundance over the four-year period (2020–2023). Among the habitat factors, only light (canopy opening) had a negative effect on seedling abundance in all four years. Light resources may be a limiting factor for seedlings, and determine seedling dynamics in subtropical forests. Overall, our results demonstrated that phylogenetic density dependence and habitat filtering affected subtropical seedling abundance. Our findings provide new evidence of the impact of phylogenetic density dependence on seedling abundance in a subtropical mid-montane humid evergreen broad-leaved forest and highlight the need to incorporate the neighborhood effect, phylogenetic relatedness, and habitat factors in models assessing seedling abundance.

Density dependence and habitat filtering have been proposed to aid in understanding community assembly and species coexistence. Phylogenetic relatedness between neighbors was used as a proxy for assessing the degree of ecological similarity among species. There are different conclusions regarding the neighborhood effect in previous studies with different phylogenetic indices or at different spatiotemporal scales. However, the effects of density dependence, neighbor phylogenetic relatedness, and habitat filtering on seedling survival with different phylogenetic indices or at different temporal and spatial scales are poorly understood. We monitored 916 seedlings representing 56 woody plant species within a 4-ha forest dynamics plot for 4 years (from 2020 to 2023) in a subtropical mid-mountain moist evergreen broad-leaved forest in the Gaoligong Mountains, Southwestern China. Using generalized linear mixed models, we tested whether and how four phylogenetic indices: total phylogenetic distance (TOTPd), average phylogenetic distance (AVEPd), relative average phylogenetic distance (APd'), and relative nearest taxon phylogenetic distance (NTPd'), three temporals (1, 2, and 3 years), and spatial scales (1, 2, and 4 ha) affect the effect of density dependence, phylogenetic density dependence, and habitat filtering on seedling survival. We found evidence of the effect of phylogenetic density dependence in the 4-ha forest dynamics plot. The effects of density dependence, phylogenetic density dependence, and habitat filtering on seedling survival were influenced by phylogenetic indices and temporal and spatial scales. The effects of phylogenetic density dependence and habitat filtering on seedling survival were more conspicuous only at 1-year intervals, compared with those at 2- and 3-year intervals. We did not detect any effects of neighborhood or habitat factors on seedling survival at small scales (1 and 2 ha), although these effects were more evident at the largest spatial scale (4 ha). These findings highlight that the effects of local neighborhoods and habitats on seedling survival are affected by phylogenetic indices as well as temporal and spatial scales. Our study suggested that phylogenetic index APd', shortest time scale (1 year), and largest spatial scales (4 ha) were suitable for neighborhood studies in a mid-mountain moist evergreen broad-leaved forest in Gaoligong Mountains. Phylogenetic indices and spatiotemporal scales have important impacts on the results of the neighborhood studies.

ABSTRACTUnderstanding the mechanisms of species coexistence is a long‐standing goal in community ecology. The performance of woody seedlings across different ecological niches has been proposed as a key mechanism in community assembly. Factors influencing seedling survival can be revealed by analyzing neighboring plants in conjunction with their phylogenetic relatedness, functional traits, and the surrounding environment. In the present study, we conducted a seasonal analysis of 936 seedlings for 56 species from a 4‐ha subtropical forest over 3 years in the Gaoligong Mountains, Southwest China. Our aims were to examine the relative effects of neighbor density, habitat conditions, and seasonal climate variability on seedling survival during dry and rainy seasons, and to determine whether the sensitivity of seedling survival responding to neighbor densities and abiotic factors differs between seasons. The findings indicated that the relative importance of neighbor densities, habitat factors as well as seasonal rainfall on seedling survival varied with seasonality. During the rainy season, seedling survival was comparatively less affected by conspecific neighbor density and was mainly negatively influenced by rainfall, whereas habitat factors (including topography, soil properties, and canopy openness) were detrimental for survival to a lesser extent. There is evidence suggesting that phylogenetic negative density dependence (PNDD) and functional negative density dependence (FNDD) are more pronounced in the rainy season. In contrast, the positive effect of canopy openness was more important during the dry season. Our findings further revealed that the effects of PNDD, FNDD, canopy openness, and seasonal rainfall varied widely among species across seasons. Moreover, species differed in their ability to respond to the trade‐offs between canopy openness and rainfall in relation to phylogenetic relatedness and functional dissimilarity during the dry and rainy seasons. Overall, our results demonstrate that seasonality modulates the strength and importance of phylogenetic and functional density dependence, habitat preference, and climate for seedling survival in subtropical forests. The seasonal variations in these effects allow species to maintain coexistence across dry and rainy seasons. Consequently, seasonal variability should be accounted for in future studies in understanding the diversity of forest communities.

Snow damage to the canopy facilitates alien weed invasion in a subtropical montane primary forest in southwestern China

Increasing climate sensitivity of subtropical conifers along an aridity gradient

Species of the basidiomycetous genus Tomentella are widely distributed throughout temperate forests. Numerous studies on the taxonomy and phylogeny of Tomentella have been conducted from the temperate zone in the Northern hemisphere, but few have been from subtropical forests. In this study, four new species, T. casiae, T. guiyangensis, T. olivaceomarginata and T. rotundata from the subtropical mixed forests of Southwestern China, are described and illustrated based on morphological characteristics and phylogenetic analyses of the internal transcribed spacer regions (ITS) and the large subunit of the nuclear ribosomal RNA gene (LSU). Molecular analyses using Maximum Likelihood and Bayesian analysis confirmed the phylogenetic positions of these four new species. Anatomical comparisons among the closely related species in phylogenetic and morphological features are discussed. Four new species could be distinguished by the characteristics of basidiocarps, the color of the hymenophoral surface, the size of the basidia, the shape of the basidiospores and some other features.

Abstract. Chronically elevated nitrogen (N) deposition has led to severe nutrient imbalance in forest soils. Particularly in tropical and subtropical forest ecosystems, increasing N loading has aggravated phosphorus (P) limitation of biomass production, and has resulted in elevated emissions of nitrous oxide (N2O) and reduced uptake of methane (CH4), both of which are important greenhouse gases. Yet, the interactions of N and P and their effects on greenhouse gas emissions remain elusive. Here, we report N2O and CH4 emissions together with soil N and P data for a period of 18 months following a single P addition (79 kg P ha−1, as NaH2PO4 powder) to an N-saturated, Masson pine-dominated forest soil at TieShanPing (TSP), Chongqing, south-western (SW) China. We observed a significant decline in both nitrate (NO3−) concentrations in soil water (5 and 20 cm depths) and in soil N2O emissions, following P application. We hypothesise that enhanced N uptake by plants in response to P addition, resulted in less available NO3− for denitrification. By contrast to most other forest ecosystems, TSP is a net source of CH4. P addition significantly decreased CH4 emissions and turned the soil from a net source into a net sink. Based on our observation and previous studies in South America and China, we believe that P addition relieves N inhibition of CH4 oxidation. Within the 1.5 years after P addition, no significant increase of forest growth was observed and P stimulation of forest N uptake by understorey vegetation remains to be confirmed. Our study indicates that P fertilisation of N-saturated, subtropical forest soils may mitigate N2O and CH4 emissions, in addition to alleviating nutrient imbalances and reducing losses of N through NO3− leaching.

Nitrogen deposition has a considerable impact on biogeochemical cycling in terrestrial ecosystems. However, how litter production and element return respond to N addition remains poorly understood in nitrogen-rich subtropical regions. In this study, a 4-year nitrogen addition experiment explored its effects on foliar litter production and carbon, nitrogen and phosphorus in a subtropical Michelia wilsonii forest. A clear seasonal pattern in foliar litterfall was observed, regardless of nitrogen treatments, with a peak in spring and a smaller one in autumn. Foliar litter increased with increasing nitrogen but did not affect litter carbon concentrations and often decreased nitrogen and phosphorous concentrations. The effect of nitrogen addition was dependent on time (month/year). Carbon, nitrogen and phosphorous return showed similar bimodal seasonal patterns. Nitrogen addition increased carbon and nitrogen return but did not affect phosphorous. Our results suggest that the addition of nitrogen stimulates carbon and nutrient return via litterfall.

AimsUnderstanding the joint effects of plant development and environment on shifts of intraspecific leaf traits will advance the understandings of the causes of intraspecific trait variation. We address this question by focusing on a widespread species Clausena dunniana in a subtropical broad‐leaved forest.MethodsWe sampled 262 individuals of C. dunniana at two major topographic habitat types, the slope and hilltop, within the karst forests in Maolan Nature Reserve in southwestern China. We measured individual plant level leaf traits (i.e., specific leaf area (SLA), leaf area, leaf dry‐matter content (LDMC), and leaf thickness) that are associated with plant resource‐use strategies. We adopted a linear mixed‐effects model in which the plant size (i.e., the first principal component of plant basal diameter and plant height) and environmental factors (i.e., topographic habitat, canopy height, and rock‐bareness) were used as independent variables, to estimate their influences on the shifts of leaf traits.Key ResultsWe found that (1) plant size and the environmental factors independently drove the intraspecific leaf trait shifts of C. dunniana, of which plant size explained less variances than environmental factors. (2) With increasing plant size, C. dunniana individuals had increasingly smaller SLA but larger sized leaves. (3) The most influential environmental factor was topographic habitat; it drove the shifts of all the four traits examined. Clausena dunniana individuals on hilltops had leaf traits representing more conservative resource‐use strategies (e.g., smaller SLA, higher LDMC) than individuals on slopes. On top of that, local‐scale environmental factors further modified leaf trait shifts.ConclusionsPlant size and environment independently shaped the variations in intraspecific leaf traits of C. dunniana in the subtropical karst forest of Maolan. Compared with plant size, the environment played a more critical role in shaping intraspecific leaf trait variations, and potentially also the underlying individual‐level plant resource‐use strategies.

Severe seasonal droughts driven by global climate change significantly alter the cycling of carbon and nutrients in forest ecosystems, while the investigation into the impacts of floor mass and plant roots on soil microbial biomass within the context of recurrent seasonal droughts is still rare. To investigate the environmental determinants governing soil microbial biomass with the escalating severity of seasonal droughts, we conducted a study in a montane subtropical moist evergreen broad-leaved forest in southwestern China from June 2019 to May 2023. The study results revealed that soil microbial biomass, as well as soil moisture, floor mass, and plant roots, showed an apparent single-hump modal within one year. In the comparative analysis of the soil microbial biomass fluctuation amplitudes across control and watered plots, a discernible disparity was observed, indicating significant differences in microbial biomass dynamics between the respective experimental conditions. The pooled data revealed a statistically significant influence of seasonal drought, floor mass, plant roots, and their reciprocal interactions on the soil microbial biomass, highlighting these factors as pivotal determinants of microbial community dynamics. This study elucidates the interactive regulatory mechanisms by which seasonal drought, floor mass, and plant roots collectively modulate soil microbial biomass within tropical and subtropical forests, offering insights into the complex ecological processes governing microbial community dynamics. This interactive regulation might influence the trajectory of plant species and soil microbial communities, facilitating their adaptive development and evolutionary responses.

As heterotrophic respiration (RH) has great potential to increase atmospheric CO2 concentrations, it is important to understand warming effects on RH for a better prediction of carbon–climate feedbacks. However, it remains unclear how RH responds to warming in subtropical forests. Here, we carried out trenching alone and trenching with warming treatments to test the climate warming effect on RH in a subtropical forest in southwestern China. During the measurement period, warming increased annual soil temperature by 2.1 °C, and increased annual mean RH by 22.9%. Warming effect on soil temperature (WET) showed very similar pattern with warming effect on RH (WERH), decreasing yearly. Regression analyses suggest that WERH was controlled by WET and also regulated by the soil water content. These results showed that the decrease of WERH was not caused by acclimation to the warmer temperature, but was instead due to decrease of WET. We therefore suggest that global warming will accelerate soil carbon efflux to the atmosphere, regulated by the change in soil water content in subtropical forests.

Negative plant–soil feedback (PSF) has been widely considered to be a primary mechanism maintaining plant diversity. Previous studies have shown that rare species suffer stronger negative conspecific PSF than common species, but it remains unclear how rare species persist if they are more strongly self‐limited. Here, we used shade‐house and field experiments to test soil feedback effects of phylogenetically related species on seedling growth, with seven species of contrasting local abundance, in a subtropical forest, China. We quantified the PSF of conspecifics and heterospecifics and assessed the phylogenetic dependence of the feedback. Both experiments showed that although rare species suffered strong negative PSF in soils of conspecifics or phylogenetically close heterospecifics, no such feedback was found in the soils of phylogenetically distant heterospecifics. In contrast, common species had no or weak negative conspecific PSF but strong heterospecific soil PSF. Synthesis. The variation in the phylogenetically dependent PSF among rare and common species evidenced in this study ensures that rare species would grow well in the neighbourhood of phylogenetically distant heterospecifics but do poorly under their own or close relatives, while common species perform relatively well in their own neighbourhood but poorly in other's neighbourhood. This phylogenetically dependent PSF facilitates the rare–common species coexistence in communities.

Mercury distribution in the foliage and soil profiles of a subtropical forest: Process for mercury retention in soils

Rising El Niño–Southern Oscillation (ENSO) variability is expected to influence Earth's forest ecosystems, through changes in how coordinated annual tree growth is across large spatiotemporal scales. However, the mechanisms by which changes in ENSO variability affect tree growth in subtropical forests remains poorly understood. We use a newly built tree ring network collected from 4,028 trees at 144 forest locations across East Asian subtropical forests (EASF), to assess long-term influences of ENSO on the spatiotemporal variability in tree radial growth across China at subcontinental scales (∼2,000 km). Our results demonstrate a west-east dipole pattern of synchronized tree growth in EASF tree populations, with positive growth responses to El Nino in southeastern China, and negative growth responses in southwestern China, likely due to different patterns in moisture limitation. Specifically, the likely contrasting effects of ENSO on drought limitation along a longitudinal gradient resulted in trees that grew more in El Niño years in eastern populations, but less in western populations. Our results also show that increasingly severe El Niño/La Niña years have caused a sharp rise in tree growth coherence over the past 150 years in these moisture-limited populations. Increasing ENSO variability, as is expected with climate change, may potentially further destabilize subtropical forest ecosystems by synchronizing tree growth to an unprecedented level, with unknown consequences for forest stability and resilience.

Rising El Niño–Southern Oscillation (ENSO) variability is expected to influence Earth’s forest ecosystems, through changes in how coordinated annual tree growth is across large spatiotemporal scales. However, the mechanisms by which changes in ENSO variability affect tree growth remains poorly understood, especially in understudied subtropical forests. We use a newly built tree ring network collected from 4,028 trees at 144 forest locations across East Asian subtropical forests (EASF) at subcontinental scales (∼2,000 km), to assess long-term influences of ENSO on the spatiotemporal variability in tree radial growth across China. Our results demonstrate a west-east dipole pattern of synchronized tree growth in EASF moisture-limited tree populations, with positive growth responses to El Nino in southeastern China, and negative growth responses in the southwestern China. Specifically, trees grew more in El Niño years in eastern populations, but less in western populations. This pattern likely results from the contrasting effects of ENSO on drought limitation along a longitudinal gradient. Our results also show that increasingly severe El Niño/La Niña years have caused a sharp rise in tree growth coherence over past 150 years in these moisture-limited populations. A further increase in climate variability, as is expected with climate change, could destabilize subtropical forest ecosystems by synchronizing tree growth to an unprecedented level. In all, our results highlight the need for further research on the ecological implications of rising synchrony, given its increasing relevance to global forest ecosystems in a time of rising climate variability.