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Article ; Online: Differences in regeneration niche mediate how disturbance severity and microclimate affect forest species composition

Becker, Kendall M.L. / Lutz, James A.

Forest Ecology and Management. 2023 Sept., v. 544 p.121190-

2023

Abstract: Climate change is altering forest composition through species-specific responses to fire and drought. Future forest composition will depend on how the different regeneration niches of co-occurring species align with current environmental conditions, ... ...

Abstract	Climate change is altering forest composition through species-specific responses to fire and drought. Future forest composition will depend on how the different regeneration niches of co-occurring species align with current environmental conditions, especially after fire, which can promote germination by exposing mineral soil. Few studies, however, have examined the effects of disturbance severity and microclimate on post-fire regeneration to define and compare the regeneration niches of co-occurring tree species. We used seven years of annual demography and microenvironment data from a 25.6-ha fully censused, stem-mapped forest dynamics plot in California, USA, to examine how disturbance severity, snow duration, and temperature extremes affect the survival of Abies concolor and Pinus lambertiana seedlings that germinated naturally after a low- to moderate-severity fire. We defined disturbance severity at the microsite level, based on characteristics of the substrate, and at the neighborhood level, based on tree mortality. Both disturbance severity and snow duration had species-specific effects on seedling survival, but these differed by life stage. During the germination year, later snow disappearance was associated with a 0.5 increase in survival probability for A. concolor but hardly affected P. lambertiana; in contrast, higher neighborhood disturbance severity increased survival of both species. After the germination year, higher substrate burn severity was associated with a 0.8 increase in survival probability for A. concolor but hardly affected P. lambertiana; higher neighborhood disturbance severity and later snow disappearance increased annual survival of both species, but maximum summer temperature had minimal effect. Overall, available seed, higher substrate burn severity, higher neighborhood disturbance severity, and later snow disappearance promoted natural regeneration. However, lower substrate burn severity and earlier snow disappearance in the germination year disadvantaged A. concolor seedlings, increasing the relative abundance of P. lambertiana seedlings compared to the local tree population. Our results indicate that natural post-fire compositional shifts toward drought-tolerant Pinus species–and away from less drought-tolerant Abies species–are possible in the Sierra Nevada, with potential benefits for forest persistence under climate change. Broadly, we show that species differences in regeneration niches shape how disturbance severity and microclimate affect forest species composition.
Keywords	Abies concolor ; Pinus lambertiana ; administrative management ; burn severity ; climate change ; demography ; drought ; drought tolerance ; forest dynamics ; forests ; germination ; microclimate ; mineral soils ; mountains ; natural regeneration ; probability ; seedlings ; snow ; species diversity ; summer ; temperature ; tree mortality ; trees ; California ; Low-severity fire ; Moderate-severity fire ; Old-growth forest ; Post-fire seedlings ; Regeneration niche ; Rim fire ; Smithsonian ForestGEO ; Yosemite Forest Dynamics Plot
Language	English
Dates of publication	2023-09
Publishing place	Elsevier B.V.
Document type	Article ; Online
ZDB-ID	751138-3
ISSN	0378-1127
ISSN	0378-1127
DOI	10.1016/j.foreco.2023.121190
Database	NAL-Catalogue (AGRICOLA)

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Article ; Online: Shared friends counterbalance shared enemies in old forests.

Germain, Sara J / Lutz, James A

Ecology

2021 Volume 102, Issue 11, Page(s) e03495

Abstract: Mycorrhizal mutualisms are nearly ubiquitous across plant communities. Yet, it is still unknown whether facilitation among plants arises primarily from these mycorrhizal networks or from physical and ecological attributes of plants themselves. Here, we ... ...

Abstract	Mycorrhizal mutualisms are nearly ubiquitous across plant communities. Yet, it is still unknown whether facilitation among plants arises primarily from these mycorrhizal networks or from physical and ecological attributes of plants themselves. Here, we tested the relative contributions of mycorrhizae and plants to both positive and negative biotic interactions to determine whether plant-soil feedbacks with mycorrhizae neutralize competition and enemies within multitrophic forest community networks. We used Bayesian hierarchical generalized linear modeling to examine mycorrhizal-guild-specific and mortality-cause-specific woody plant survival compiled from a spatially and temporally explicit data set comprising 101,096 woody plants from three mixed-conifer forests across western North America. We found positive plant-soil feedbacks for large-diameter trees: species-rich woody plant communities indirectly promoted large tree survival when connected via mycorrhizal networks. Shared mycorrhizae primarily counterbalanced apparent competition mediated by tree enemies (e.g., bark beetles, soil pathogens) rather than diffuse competition between plants. We did not find the same survival benefits for small trees or shrubs. Our findings suggest that lower large-diameter tree mortality susceptibility in species-rich temperate forests resulted from greater access to shared mycorrhizal networks. The interrelated importance of aboveground and belowground biodiversity to large tree survival may be critical for counteracting increasing pathogen, bark beetle, and density threats.
MeSH term(s)	Bayes Theorem ; Forests ; Mycorrhizae ; Soil ; Soil Microbiology
Chemical Substances	Soil
Language	English
Publishing date	2021-08-20
Publishing country	United States
Document type	Journal Article ; Research Support, Non-U.S. Gov't ; Research Support, U.S. Gov't, Non-P.H.S.
ZDB-ID	2010140-5
ISSN	1939-9170 ; 0012-9658
ISSN (online)	1939-9170
ISSN	0012-9658
DOI	10.1002/ecy.3495
Database	MEDical Literature Analysis and Retrieval System OnLINE

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Article ; Online: The importance of regeneration processes on forest biodiversity in old-growth forests in the Pacific Northwest.

Luu, Hoang / Ris Lambers, Janneke Hille / Lutz, James A / Metz, Margaret / Snell, Rebecca S

Philosophical transactions of the Royal Society of London. Series B, Biological sciences

2024 Volume 379, Issue 1902, Page(s) 20230016

Abstract: Forest diversity is the outcome of multiple species-specific processes and tolerances, from regeneration, growth, competition and mortality of trees. Predicting diversity thus requires a comprehensive understanding of those processes. Regeneration ... ...

Abstract	Forest diversity is the outcome of multiple species-specific processes and tolerances, from regeneration, growth, competition and mortality of trees. Predicting diversity thus requires a comprehensive understanding of those processes. Regeneration processes have traditionally been overlooked, due to high stochasticity and assumptions that recruitment is not limiting for forests. Thus, we investigated the importance of seed production and seedling survival on forest diversity in the Pacific Northwest (PNW) using a forest gap model (ForClim). Equations for regeneration processes were fit to empirical data and added into the model, followed by simulations where regeneration processes and parameter values varied. Adding regeneration processes into ForClim improved the simulation of species composition, compared to Forest Inventory Analysis data. We also found that seed production was not as important as seedling survival, and the time it took for seedlings to grow into saplings was a critical recruitment parameter for accurately capturing tree species diversity in PNW forest stands. However, our simulations considered historical climate only. Due to the sensitivity of seed production and seedling survival to weather, future climate change may alter seed production or seedling survival and future climate change simulations should include these regeneration processes to predict future forest dynamics in the PNW. This article is part of the theme issue 'Ecological novelty and planetary stewardship: biodiversity dynamics in a transforming biosphere'.
MeSH term(s)	Forests ; Trees ; Biodiversity ; Seedlings ; Northwestern United States
Language	English
Publishing date	2024-04-08
Publishing country	England
Document type	Journal Article
ZDB-ID	208382-6
ISSN	1471-2970 ; 0080-4622 ; 0264-3839 ; 0962-8436
ISSN (online)	1471-2970
ISSN	0080-4622 ; 0264-3839 ; 0962-8436
DOI	10.1098/rstb.2023.0016
Database	MEDical Literature Analysis and Retrieval System OnLINE

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Article: Dancing with Douglas‐fir: Determinism dominates fungal community assembly processes

Birch, Joseph D. / Lutz, James A. / Karst, Justine

journal of ecology. 2022 Aug., v. 110, no. 8

2022

Abstract: Fungal communities can influence the productivity, composition and survival of trees through cycling nutrients, providing resources and altering pathogens. Thus, shifts in fungal communities could impact forests by altering interactions between trees and ...

Abstract	Fungal communities can influence the productivity, composition and survival of trees through cycling nutrients, providing resources and altering pathogens. Thus, shifts in fungal communities could impact forests by altering interactions between trees and their environments. Fungal community composition may be shaped by stochastic and deterministic processes such as dispersal limitation, environmental filtering and partner specificity between trees and fungi. For tree species with large geographical ranges, we expect fungal assembly processes to change with environmental variation across the range of the tree partner. Due to specificity between trees and symbiotic fungi, we expect deterministic to outweigh stochastic processes in root compared with soil fungi. As some tree species have exceptional longevity, we also expected tree age to influence fungal community assembly. We surveyed fungi in four stands of Pseudotsuga menziesii with tree ages up to 800 years along an 1,800 km transect. We sampled roots and soil around 12 P. menziesii in each stand, aged the trees, and sequenced fungal rDNA to determine composition and richness from which we calculated the relative role of deterministic and stochastic assembly processes. We used null models to evaluate the relative importance of deterministic variable and homogenizing selection, and stochastic dispersal limitation, drift and homogenizing dispersal in fungal community assembly. We detected 7,280 amplicon sequence variants with 5,270 associated with soil, 3,887 with roots and 1,877 found across both roots and soils. Deterministic processes dominated root and soil fungal communities at all sites except one where stochastic processes (i.e. dispersal limitation and drift) controlled root fungi. Despite the dominance of determinism in fungal community assembly, the proportion of processes differed by site. Assembly processes did not vary with tree age. Synthesis. Taken together, we suggest that the local environment, water limitation and partner‐preference between trees and their associated fungi, influence fungal community composition across the range of P. menziesii. We conclude that while fungal communities occurring near P. menziesii are dominated by homogenizing selection, the role of neutral processes still has a minor influence on community assembly and may be important in spatially isolated communities and those with strong gradients of fungal diversity.
Keywords	Pseudotsuga menziesii ; community structure ; environmental factors ; fungal communities ; longevity ; soil ; soil fungi ; tree age ; trees
Language	English
Dates of publication	2022-08
Size	p. 1857-1870.
Publishing place	John Wiley & Sons, Ltd
Document type	Article
Note	JOURNAL ARTICLE
ZDB-ID	3023-5
ISSN	0022-0477
ISSN	0022-0477
DOI	10.1111/1365-2745.13910
Database	NAL-Catalogue (AGRICOLA)

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Article: Climate warming may weaken stabilizing mechanisms in old forests

Germain, Sara J. / Lutz, James A.

Ecological monographs. 2022 May, v. 92, no. 2

2022

Abstract: Plant competition may intensify with climate warming, but whether this will occur equally for conspecific and heterospecific competition remains unknown. Competitive shifts have the potential to instigate community change because the relative strengths ... ...

Abstract	Plant competition may intensify with climate warming, but whether this will occur equally for conspecific and heterospecific competition remains unknown. Competitive shifts have the potential to instigate community change because the relative strengths of conspecific and heterospecific negative density dependence mediate the stabilizing mechanisms underpinning species coexistence. We examined a mature temperate forest to assess both direct and indirect climate effects at multiple scales: individual species, interspecies relationships, and community stability mechanisms. Our coupled approach (1) quantified tree mortality risk dependence on the interactive effects of competition, climatic water deficit, snowpack, and soil moisture for 28,913 trees over 8 years (3149 mortalities), then (2) used a climate‐projection ensemble to forecast changes in conspecific and heterospecific competition from 2020 to 2100. We predict that projected climate warming will destabilize the foundational forest community by increasing the strength of heterospecific competition at a greater rate and to a greater degree than conspecific competition for four of five abundant tree species, particularly on dry microsites. Modeling showed that these findings were most pronounced after the year 2038, at which point snowpacks were projected to be too small to ameliorate the effects of drought on competitive interactions. Our finding that heterospecific competition is more sensitive than conspecific competition to climate warming may indicate the impending loss of ecosystem functioning. We join the growing body of work showing a predominance of indirect drought effects, yet coupled climate models still fail to consider how changing community dynamics may impact forest cover and, in turn, disrupt forest–climate carbon feedbacks. Ecosystems sharing characteristics with our example forest—those with low species richness and therefore a limited biodiversity insurance effect—may be similarly vulnerable to climate‐mediated destabilization. In such communities, increased heterospecific competition among even a small number of species can more easily destabilize communities without recourse from redundant species. This study of an overlooked but vital mechanism of community change can be adapted by research in a range of ecosystems to improve the understanding of climate change consequences.
Keywords	carbon ; climate ; conspecificity ; drought ; ecosystems ; insurance ; intraspecific competition ; plant competition ; risk ; snowpack ; soil water ; species richness ; temperate forests ; tree mortality ; trees
Language	English
Dates of publication	2022-05
Publishing place	John Wiley & Sons, Inc.
Document type	Article
Note	JOURNAL ARTICLE
ZDB-ID	410216-2
ISSN	1557-7015 ; 0012-9615
ISSN (online)	1557-7015
ISSN	0012-9615
DOI	10.1002/ecm.1508
Database	NAL-Catalogue (AGRICOLA)

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Article: Soil Enzyme Activity and Soil Nutrients Jointly Influence Post-Fire Habitat Models in Mixed-Conifer Forests of Yosemite National Park, USA

Tamjidi, Jelveh / Lutz, James A

Fire. 2020 Sept. 23, v. 3, no. 4

2020

Abstract: Disentangling the relative importance of habitat filtering and dispersal limitations at local scales (<1 km²) in shaping species composition remains an important question in community ecology. Previous studies have examined the relative importance of ... ...

Abstract	Disentangling the relative importance of habitat filtering and dispersal limitations at local scales (<1 km²) in shaping species composition remains an important question in community ecology. Previous studies have examined the relative importance of these mechanisms using topography and selected soil properties. We examined both topography and edaphic properties from 160 locations in the recently burned 25.6 ha Yosemite Forest Dynamics Plot (YFDP) in Yosemite National Park, California, USA. In addition to eight soil chemical properties, we included phosphatases and urease enzymes in a definition of habitat niches, primarily because of their rapid changes with fire (compared to soil nutrients) and also their role in ecosystem function. We applied environmental variables to the distributions of 11 species. More species–habitat associations were defined by soil properties (54.5%) than topographically-defined habitat (45.4%). We also examined the relative importance of spatial and environmental factors in species assemblage. Proportions explained by spatial and environmental factors differed among species and demographic metrics (stem abundance, basal area increment, mortality, and recruitment). Spatial factors explained more variation than environmental factors in stem abundance, mortality, and recruitment. The contributions of urease and acid phosphatase to habitat definition were significant for species abundance and basal area increment. These results emphasize that a more complete understanding of niche parameters is needed beyond simple topographic factors to explain species habitat preference. The stronger contribution of spatial factors suggests that dispersal limitation and unmeasured environmental variables have high explanatory power for species assemblage in this coniferous forest.
Keywords	acid phosphatase ; coniferous forests ; ecological function ; enzyme activity ; forest dynamics ; habitat preferences ; mortality ; national parks ; soil enzymes ; species abundance ; topography ; urease ; California
Language	English
Dates of publication	2020-0923
Publishing place	Multidisciplinary Digital Publishing Institute
Document type	Article
Note	NAL-AP-2-clean
ISSN	2571-6255
DOI	10.3390/fire3040054
Database	NAL-Catalogue (AGRICOLA)

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Article: The Post-Fire Assembly Processes of Tree Communities Based on Spatial Analysis of a Sierra Nevada Mixed-Conifer Forest

Tamjidi, Jelveh / Lutz, James A

Fire. 2020 Dec. 18, v. 3, no. 4

2020

Abstract: Understanding the mechanisms underlying tree spatial arrangements may provide significant insights into the processes in the maintenance of species coexistence. We examined the potential role of habitat heterogeneity, dispersal limitation, negative ... ...

Abstract	Understanding the mechanisms underlying tree spatial arrangements may provide significant insights into the processes in the maintenance of species coexistence. We examined the potential role of habitat heterogeneity, dispersal limitation, negative density dependence, fire history, and unilateral intraspecific and interspecific interactions of adults on juveniles in shaping the spatial patterns of four dominant tree species (Abies concolor, Pinus lambertiana, Calocedrus decurrens, and Quercus kelloggii) after fire in the Yosemite Forest Dynamic Plot, California, USA. We used the univariate pair correlation function and implemented three point pattern processes (homogeneous Poisson process, inhomogeneous Poisson process, and homogeneous Thomas process) to evaluate the potential contributions of habitat filtering and dispersal limitation. We used a bivariate null model to evaluate unilateral intraspecific and interspecific interactions of adults on juveniles. We also used the pairwise correlation function to investigate the spatial patterns of density dependence. To understand the effect of fire, we used the univariate pair correlation function to investigate pattern changes during the six years following fire. We compared spatial pattern changes in both sprouting species (Quercus kelloggii) and seeding species (Abies concolor), and also examined the changes in patterns of large-diameter individuals of Abies concolor, Pinus lambertiana, and Calocedrus decurrens in 2013 (pre-fire), 2016 (two years post-fire), and 2019. Comparing the contributions of the homogeneous Thomas process and the inhomogeneous Poisson process at different spatial scales showed the importance of dispersal limitation and habitat heterogeneity at finer scales (0 m to 5 m) and coarser scales (5 m to 60 m), respectively, which suggests that the joint effects of dispersal limitation and habitat heterogeneity contribute to the spatial patterns of these three dominant tree species. Furthermore, the results showed that the young individuals of Abies concolor and Pinus lambertiana were more commonly found around the conspecific adults. Juvenile regeneration to the 1 cm diameter threshold was highly aggregated following the fire. Large-diameter trees of Abies concolor, Pinus lambertiana, and Calocedrus decurrens generally did not exhibit patterns different from complete spatial randomness (Calocedrus decurrens), or displayed only slight aggregation (Abies concolor and Pinus lambertiana). In addition, Abies concolor and Pinus lambertiana showed positive and negative conspecific density dependence in the immediate post-fire period, respectively.
Keywords	Abies concolor ; Calocedrus decurrens ; Pinus lambertiana ; Quercus kelloggii ; conspecificity ; fire history ; forests ; habitats ; juveniles ; models ; mountains ; trees ; California
Language	English
Dates of publication	2020-1218
Publishing place	Multidisciplinary Digital Publishing Institute
Document type	Article
Note	NAL-AP-2-clean
ISSN	2571-6255
DOI	10.3390/fire3040072
Database	NAL-Catalogue (AGRICOLA)

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Article ; Online: Elevated mortality rates of large trees allow for increased frequency of intermediate trees: a hypothesis supported by demographic model comparison with plot and LiDAR data

Francis, Emily J. / Lutz, James A. / Farrior, Caroline E.

Forest Ecology and Management. 2023, p.121035-

2023 , Page(s) 121035–

Abstract: Tree diameter distributions are important indicators of forest structure, are a principal element of forest carbon stock estimates, and are theoutcome of forest demography. Hump-shaped tree diameter distributions, also known as rotated-sigmoid diameter ... ...

Abstract	Tree diameter distributions are important indicators of forest structure, are a principal element of forest carbon stock estimates, and are theoutcome of forest demography. Hump-shaped tree diameter distributions, also known as rotated-sigmoid diameter distributions, are characterized by an increased frequency of intermediately-sized trees over what would be extrapolated from the small tree size distribution and are common in temperate forests. One hypothesis to explain hump-shaped tree size distributions is the U-shaped mortality curve, where the mortality rates of the largest canopy trees are higher than those of intermediately-sized canopy trees. However, studies that have directly tested this hypothesis by comparing tree diameter distributions predicted from forest demographic models with empirical diameter distributions are lacking, and how the U-shaped mortality curve influences other aspects of forest structure such as canopy height is not well understood. We used model-data comparisons to test the hypothesis that the U-shaped mortality curve generates the hump-shaped tree diameter distribution. We used two versions of a forest demographic model (the PPA model) to predict tree diameter distributions, one that assumed that tree demographic rates varied only between understory and canopy trees, and a second that assumed tree demographic rates varied among understory trees, small canopy trees, and large canopy trees, consistent with the U-shaped mortality curve. We compared predictions from each form of the model with empirical tree diameter distributions from the Wind River Forest Dynamics plot, a 27.2 hectare forest dynamics plot with a hump-shaped diameter distribution. We found that the model that accounted for the higher mortality rates of larger canopy trees generated a hump-shaped distribution, whereas the model that did not account for the higher mortality rates of larger trees was not able to generate a hump-shaped diameter distribution. To understand how the assumptions of forest demographic patterns affected other aspects of forest structure that could be derived from remote sensing, we compared predictions of forest canopy height from each form of the model with measurements of forest canopy height derived from airborne LiDAR. We found that accounting for the higher mortality rates of larger trees became even more critical to accurate prediction of forest canopy height because intermediate-sized trees occupy large areas in the canopy. Our results demonstrate the link between the U-shaped mortality curve and hump-shaped diameter distributions, and suggest that accurate characterization of the demography of large trees may be particularly important for predicting forestcanopy structure and carbon stocks.
Keywords	administrative management ; canopy height ; carbon ; carbon sinks ; demography ; forest canopy ; forest dynamics ; forests ; lidar ; models ; mortality ; prediction ; rivers ; tree and stand measurements ; trees ; understory ; forest demography ; U-shaped mortality ; tree diameter distribution ; forest demographic model ; rotated-sigmoid diameter distribution
Language	English
Publishing place	Elsevier B.V.
Document type	Article ; Online
Note	Pre-press version
ZDB-ID	751138-3
ISSN	0378-1127
ISSN	0378-1127
DOI	10.1016/j.foreco.2023.121035
Database	NAL-Catalogue (AGRICOLA)

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Article: Shared friends counterbalance shared enemies in old forests

Germain, Sara J. / Lutz, James A.

Ecology. 2021 Nov., v. 102, no. 11

2021

Abstract	Mycorrhizal mutualisms are nearly ubiquitous across plant communities. Yet, it is still unknown whether facilitation among plants arises primarily from these mycorrhizal networks or from physical and ecological attributes of plants themselves. Here, we tested the relative contributions of mycorrhizae and plants to both positive and negative biotic interactions to determine whether plant–soil feedbacks with mycorrhizae neutralize competition and enemies within multitrophic forest community networks. We used Bayesian hierarchical generalized linear modeling to examine mycorrhizal‐guild‐specific and mortality‐cause‐specific woody plant survival compiled from a spatially and temporally explicit data set comprising 101,096 woody plants from three mixed‐conifer forests across western North America. We found positive plant–soil feedbacks for large‐diameter trees: species‐rich woody plant communities indirectly promoted large tree survival when connected via mycorrhizal networks. Shared mycorrhizae primarily counterbalanced apparent competition mediated by tree enemies (e.g., bark beetles, soil pathogens) rather than diffuse competition between plants. We did not find the same survival benefits for small trees or shrubs. Our findings suggest that lower large‐diameter tree mortality susceptibility in species‐rich temperate forests resulted from greater access to shared mycorrhizal networks. The interrelated importance of aboveground and belowground biodiversity to large tree survival may be critical for counteracting increasing pathogen, bark beetle, and density threats.
Keywords	Bayesian theory ; bark ; bark beetles ; biodiversity ; data collection ; mycorrhizae ; pathogens ; soil ; tree mortality ; trees ; woody plants ; North America
Language	English
Dates of publication	2021-11
Publishing place	John Wiley & Sons, Ltd
Document type	Article
Note	JOURNAL ARTICLE
ZDB-ID	1797-8
ISSN	0012-9658
ISSN	0012-9658
DOI	10.1002/ecy.3495
Database	NAL-Catalogue (AGRICOLA)

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Article: Scale‐dependent species–area relationship: Niche‐based versus stochastic processes in a typical subtropical forest

Ren, Haibao / Svenning, Jens‐Christian / Mi, Xiangcheng / Lutz, James A. / Zhou, Jinxing / Ma, Keping

journal of ecology. 2022 Aug., v. 110, no. 8

2022

Abstract: Determining the patterns and drivers of the small‐scale species–area relationship (SAR) is crucial for improving our understanding of community assembly and biodiversity patterns. Niche‐based and stochastic processes are two principal categories of ... ...

Abstract	Determining the patterns and drivers of the small‐scale species–area relationship (SAR) is crucial for improving our understanding of community assembly and biodiversity patterns. Niche‐based and stochastic processes are two principal categories of mechanisms potentially driving SARs. However, their relative importance has rarely been quantified rigorously owing to scale dependence and the simplified niche volumes often used. In a fully mapped, 24‐hm² plot of a typical subtropical forest, we built the SARs and well‐defined niche hyper‐volumes of a broad range of environmental variables at scales of 10–70 m (cell sizes). We then simulated passive sampling and partitioned the variances of the SAR slopes to disentangle these two contrasting mechanisms. We found that the small‐scale SAR best followed a power‐law relationship, consistent with large‐scale SARs. The SAR slope declined with increasing scale; it was lower than expected under passive sampling at scales below 30 m and higher at larger scales. Environmental niches explained more (39%–64%) of the slope at larger scales, exceeding 50% at scales >30 m, and these niches always captured the majority of the structured slopes. Environmental position (environmental mean values) effects were steady in absolute strength across scales and explained most (98%–68%) of the niche effect, but this proportion decreased with increasing scale. The effect of environmental heterogeneity increased with spatial scales, starting to rise at the 30 m scale after controlling for environmental position. Excluding soil properties from analyses strongly reduced these niche effects, highlighting the importance of soils for structuring the small‐scale SAR. There was also substantial stochasticity in the SAR slopes, which was only partially explained by passive sampling. Synthesis. Our results show that the small‐scale SAR in the studied subtropical forest follows a power law, exhibits a scale shift in SAR slope at 30 m, and is strongly shaped by niche effects that are dominated by environmental position relative to heterogeneity. However, soil heterogeneity controls the increase in niche effect and the shift in the SAR slope with increasing spatial scales. Hence, edaphic factors can be responsible for scale dependence in small‐scale SARs, thereby linking small‐scale and large‐scale SARs.
Keywords	biodiversity ; soil heterogeneity ; tropical forests
Language	English
Dates of publication	2022-08
Size	p. 1883-1895.
Publishing place	John Wiley & Sons, Ltd
Document type	Article
Note	JOURNAL ARTICLE
ZDB-ID	3023-5
ISSN	0022-0477
ISSN	0022-0477
DOI	10.1111/1365-2745.13924
Database	NAL-Catalogue (AGRICOLA)

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