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Ecology

Published by Wiley and Ecological Society of America

Online ISSN: 1939-9170

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Print ISSN: 0012-9658

Disciplines: Ecology

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Illustration of the colonization–competition (C–C) trade‐off model in a multispecies (n = 6 species) system subject to multiple environmental drivers, including habitat size (S), resource productivity (R), and disturbance (D). Panel (a): initial species diversity pattern, containing two potential processes: superior competitors displacing inferior competitors via propagules (red arrows), and species colonizing empty sites via propagules (green arrows). Panel (b): increasing resource productivity (R) is assumed to enhance the colonization rates of all species (i.e., increasing the amount of propagules, such as seeds). Panel (c): reducing habitat size (S) by directly removing sites from the initial habitat in panel (a) (i.e., habitat loss). Panel (d): stochastic disturbance resulting in species mortality randomly across the habitat (red cross). Panel (e): biodiversity pattern as the product of interactions between these three drivers (represented by bold black cross). To establish the C–C trade‐off, we assume a competitive hierarchy by ranking the species from the best competitor (species 1) to the poorest (species 6), while setting species colonization rates as c1<c2<c3<…<c6$$ {c}_1<{c}_2<{c}_3<\dots <{c}_6 $$ (represented by the amount of propagules). All icons were obtained from https://www.ztupic.com/ and are in the public domain.
Separate effects of habitat size (S), resource productivity (R), and disturbance extent (D) on species diversity (a–c and g–i) and their relative abundances (d–f and j–l) in a simple multispecies community (n = 6), with a strict competitive hierarchy by ranking the species from the best competitor (species 1) to the poorest (species 6). Note that the x‐axes of S and R are not evenly scaled in order to make the zig‐zag pattern at S < 0.5 and R < 0.5 more distinguishable. Species diversity is characterized by both species richness and the inverse Simpson index. Species colonization rates (ci) are evenly spaced in increasing order in both ranges: (Case 1) ci∈E0.2,2$$ {c}_i\in E\left[0.2,2\right] $$ with all species coexisting in an intact habitat (i.e., R = S = 1 and D = 0); and (Case 2) ci∈E0.5,2.5$$ {c}_i\in E\left[0.5,2.5\right] $$ with species competitive exclusion occurring in an intact habitat. Panels (a, d, g, and j): S = 1 and D = 0; panels (b, e, h, and k): R = 1 & D = 0; and panels (c, f, i, and l): R = S = 1. Others: species mortality rates mi = 0.1.
Interactive effects of habitat size (S), resource productivity (R), and disturbance extent (D) on biodiversity, characterized by species richness and the inverse Simpson index. Panels (a, d, g, and j): D = 0; panels (b, e, h, and k): R = 1; and panels (c, f, i, and l): S = 1. Other parameter settings are the same as in Figure 2.
Interactive effects of habitat size (S), resource productivity (R), and disturbance extent (D) on biodiversity in a large initial community (n = 100), with a strict competitive hierarchy by ranking the species from the best competitor (species 1) to the poorest (species 100). Species diversity is characterized using species richness and the inverse Simpson index. Species colonization rates (ci) are evenly spaced in increasing order in both ranges: panels (a–f) ci∈E0.12,4.575$$ {c}_i\in E\left[0.12,4.575\right] $$ in Case 1 where all species can coexist in an intact habitat (i.e., R = S = 1 and D = 0); and panels (g–l) ci∈E0.5,2.5$$ {c}_i\in E\left[0.5,2.5\right] $$ in Case 2, with species competitive exclusion occurring in an intact habitat. Panels (a, d, g, and j): D = 0, panels (b, e, h, and k): R = 1, and panels (e, f, i, and l): S = 1. Other parameters: mi = 0.1.
Separate effects of habitat size (S), resource productivity (R), and disturbance extent (D) on species richness in a large initial community (n = 100), by varying other drivers at different levels (S, R or D = 0.2, 0.5, and 0.8). Panels (a, b, g, and h): D = 0, panels (c, d, i, and j): S = 1, and panels (e, f, k, and l): R = 1. Other parameter settings are the same as in Figure 4.
Complex interactive responses of biodiversity to multiple environmental drivers

November 2024

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354 Reads

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We’re the journal for ecologists, by ecologists. Over Ecology’s 100+-year history we’ve seen, published, and furthered the sharpest conceptual thinking in our field. Today, we’re still breaking new ground. With rigorous peer review and rapid publication, we’re known globally for cutting-edge novel discoveries. Clear, concise papers spanning empirical and theoretical research, varied approaches, and every area of ecology.

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Environmental characteristics during the experiments and overview of the three studied habitats and their vascular plant communities. (a) Daily average air temperature and air relative humidity (left panel) and photosynthetically active radiation (PAR) and solar radiation (right panel) from June to August. Horizontal lines show the averages across the summer. Data were registered every 15 min at 2 m height by an in situ weather station. (b—left panel) Nonmetric multidimensional scaling on Bray–Curtis dissimilarity distances of the plant species composition of experimental plots (n = 80; see main text) at peak growing season in 2017 (stress = 0.12; nonmetric fit r² = 0.99; linear fit r² = 0.94). Data (from Petit Bon et al., 2021; Petit Bon, Bråthen, et al., 2023) were analyzed in R v. 4.3.0 with the package “vegan” (Oksanen et al., 2020). Only the 10 most abundant species, making up >90% of the aboveground biomass within plots, are shown (names follow the Svalbard Flora; https://www.svalbardflora.no). Ellipses are the 95% confidence intervals of habitat centroids (permutational‐ANOVA: r² = 0.57, p < 0.0001). The four experimental treatments (see text) are displayed, with dot size proportional to plot biomass. Fit of the soil parameters when regressed on the biplot is moisture: r² = 0.80, p = 0.0001; nitrogen (N) concentration: R² = 0.26, p = 0.0041; carbon (C) concentration: R² = 0.24, p = 0.0041; details in Appendix S1: Section S1. (b—right panel) Average (±SD) aboveground plant biomass in control plots of the three habitats, sorted according to growth forms (data from references above). Photo credits: Matteo Petit Bon.
Effects of spring goose grubbing and summer warming on ecosystem CO2 fluxes in 2017. Model predictions ± SE for (a–c) gross ecosystem productivity (GEP), (d–f) ecosystem respiration (ER), and (g–i) net ecosystem exchange (NEE) in early, peak, and late summer, separately for the three habitats. Gray panels show model predictions ± SE averaged over the summer; different letters indicate significant differences among treatments. Significant and marginally significant main and interactive effects are shown (ANOVA); when an interaction was significant (p < 0.05), its main effects are not shown. Significance: ⸙p < 0.1; *p < 0.05; **p < 0.01; and ***p < 0.001. Full ANOVA results are given in Appendix S1: Table S2. LMM parameter estimates are given in Appendix S1: Tables S3–S5. Positive and negative fluxes denote CO2 losses (the ecosystem acts as a C source) and CO2 gains (the ecosystem acts as a C sink), respectively.
Effects of spring goose grubbing and summer warming on normalized‐difference vegetation index (NDVI) in 2017. Model predictions ± SE for NDVI of (a) mesic, (b) moist, and (c) wet habitats in early, peak, and late summer. Gray panels show model predictions ± SE averaged over the summer; different letters indicate significant differences among treatments. Significant and marginally significant main and interactive effects are shown (ANOVA); when an interaction was significant (p < 0.05), its main effects are not shown. Significance: ⸙p < 0.1; *p < 0.05; **p < 0.01; and ***p < 0.001. Full ANOVA results are given in Appendix S1: Table S2. LMM parameter estimates are given in Appendix S1: Table S6.
Across‐habitat relationships between ecosystem CO2 fluxes and both vegetation and abiotic variables in 2017. Regression lines ±95% CI for relations between (a–c) gross ecosystem productivity (GEP), (d–f) ecosystem respiration (ER), and (g–i) net ecosystem exchange (NEE) and the predictors (as additive smooth terms): normalized‐difference vegetation index (NDVI), soil moisture, and moss‐mat temperature (−2 cm). For each of the three models (GEP, ER, and NEE), the relationship with each predictor is shown at the average value of the other two predictors. Adjusted R² for each model: GEP: 0.70; ER: 0.46; NEE: 0.44. Rugs on the y‐axis show predicted values, whereas rugs on the x‐axis show values of the predictors (all colored according to habitat). Significance of the smooth terms: *p < 0.05; **p < 0.01; and ***p < 0.001. ANOVA results are given in Appendix S1: Table S7. Positive and negative fluxes denote CO2 losses (the ecosystem acts as a C source) and CO2 gains (the ecosystem acts as a C sink), respectively. CO2‐flux relationships with air (+10 cm) and soil (−7 cm) temperatures were similar (not shown), plausibly because of the positive correlations among plot‐level temperatures (Appendix S1: Figure S14).
Goose grubbing and warming suppress summer net ecosystem CO2 uptake differentially across high‐Arctic tundra habitats
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December 2024

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14 Reads

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Kari Anne Bråthen

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Ingibjörg S. Jónsdóttir

Environmental changes, such as climate warming and higher herbivory pressure, are altering the carbon balance of Arctic ecosystems; yet, how these drivers modify the carbon balance among different habitats remains uncertain. This hampers our ability to predict changes in the carbon sink strength of tundra ecosystems. We investigated how spring goose grubbing and summer warming—two key environmental‐change drivers in the Arctic—alter CO2 fluxes in three tundra habitats varying in soil moisture and plant‐community composition. In a full‐factorial experiment in high‐Arctic Svalbard, we simulated grubbing and warming over two years and determined summer net ecosystem exchange (NEE) alongside its components: gross ecosystem productivity (GEP) and ecosystem respiration (ER). After two years, we found net CO2 uptake to be suppressed by both drivers depending on habitat. CO2 uptake was reduced by warming in mesic habitats, by warming and grubbing in moist habitats, and by grubbing in wet habitats. In mesic habitats, warming stimulated ER (+75%) more than GEP (+30%), leading to a 7.5‐fold increase in their CO2 source strength. In moist habitats, grubbing decreased GEP and ER by ~55%, while warming increased them by ~35%, with no changes in summer‐long NEE. Nevertheless, grubbing offset peak summer CO2 uptake and warming led to a twofold increase in late summer CO2 source strength. In wet habitats, grubbing reduced GEP (−40%) more than ER (−30%), weakening their CO2 sink strength by 70%. One‐year CO2‐flux responses were similar to two‐year responses, and the effect of simulated grubbing was consistent with that of natural grubbing. CO2‐flux rates were positively related to aboveground net primary productivity and temperature. Net ecosystem CO2 uptake started occurring above ~70% soil moisture content, primarily due to a decline in ER. Herein, we reveal that key environmental‐change drivers—goose grubbing by decreasing GEP more than ER and warming by enhancing ER more than GEP—consistently suppress net tundra CO2 uptake, although their relative strength differs among habitats. By identifying how and where grubbing and higher temperatures alter CO2 fluxes across the heterogeneous Arctic landscape, our results have implications for predicting the tundra carbon balance under increasing numbers of geese in a warmer Arctic.


Geographic extent of the National Ecological Observatory Network (NEON)'s small mammal trapping data used in this study. Each circle represents one core NEON site. Site‐level tick attachment is indicated in the figure legend (defined as the percentage of white‐footed mouse, Peromyscus leucopus, captures with one or more ticks attached).
(a) Structural equation models showing direct and indirect relationships between individual conditions, behavior, and parasitism by ticks in the white‐footed mouse, Peromyscus leucopus. β values are the standardized effects sizes and represent the strength of relationships between the variables. As continuous variables were scaled prior to analyses, these effects sizes can be directly compared. R² values shown are the conditional R² for that response variable (conditional on both fixed effects and the random intercept of trapping grid ID). Marginal R² values for trappability, average movement distance, trap diversity, parasitism by larval ticks, and parasitism by nymphal ticks are 0.01, 0.02, 0.19, 0.01, and 0.02, respectively. (b) Direct, indirect, and total effects of average body mass on behavior (trappability, trap diversity, average movement distance), and the occurrence of parasitism by larval and nymphal ticks.
Predicted relationship (and 95% CI) between individual behavior, individual conditions, and parasitism by ticks (proportion of captures when an individual was observed with one or more ticks attached to the face or ears) in the white‐footed mouse, Peromyscus leucopus. (a–c) Male mice have greater occurrence of parasitism by larval ticks than females. (a, d) Mice that move farther on average have a greater occurrence of parasitism by larval and nymphal ticks. (b, e) Larger‐bodied individuals have a lower occurrence of parasitism by larval ticks but greater parasitism by nymphal ticks. (c) Male mice that are more often observed in a reproductive state have greater parasitism by larval ticks, but there is no relationship in females. Mean movement distance is in meters and mean body mass is in grams. Parasitism by larval and nymphal ticks has been back‐transformed from a logit‐transformation.
Individual variation underlies large‐scale patterns: Host conditions and behavior affect parasitism

December 2024

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9 Reads

Allison M. Brehm

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Vania R. Assis

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Lynn B. Martin

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John L. Orrock

Identifying the factors that affect host–parasite interactions is essential for understanding the ecology and dynamics of vector‐borne diseases and may be an important component of predicting human disease risk. Characteristics of hosts themselves (e.g., body condition, host behavior, immune defenses) may affect the likelihood of parasitism. However, despite highly variable rates of parasitism and infection in wild populations, identifying widespread links between individual characteristics and heterogeneity in parasite acquisition has proven challenging because many zoonoses exist over wide geographic extents and exhibit both spatial and temporal heterogeneity in prevalence and individual and population‐level effects. Using seven years of data collected by the National Ecological Observatory Network (NEON), we examined relationships among individual host condition, behavior, and parasitism by Ixodid ticks in a keystone host species, the white‐footed mouse, Peromyscus leucopus. We found that individual condition, specifically sex, body mass, and reproductive condition, had both direct and indirect effects on parasitism by ticks, but the nature of these effects differed for parasitism by larval versus nymphal ticks. We also found that condition differences influenced rodent behavior, and behavior directly affected the rates of parasitism, with individual mice that moved farther being more likely to carry ticks. This study illustrates how individual‐level data can be examined using large‐scale datasets to draw inference and uncover broad patterns in host–parasite encounters at unprecedented spatial scales. Our results suggest that intraspecific variation in the movement ecology of hosts may affect host–parasite encounter rates and, ultimately, alter zoonotic disease risk through anthropogenic modifications and natural environmental conditions that alter host space use.


Schematic of the 3 × 2 factorial experimental design. There were three rescue treatments (no‐rescue treatment: two deceased, homogenized individuals added, where “X” indicates dead individuals; low‐rescue treatment: five individuals added at each rescue event; and high‐rescue treatment: 10 individuals added at each rescue event) and 2 pathogen treatments (pathogen‐exposed or unexposed). The pathogen treatments were applied during rescue events by adding two deceased, homogenized infected individuals (no‐rescue treatment) or two live infected individuals (low‐ and high‐rescue treatments; shown as gray/solid individuals). Each treatment was replicated 10 times for 60 total experimental populations.
Population abundance through time. Pathogen treatments are shown as blue (pathogen absence) or red/orange (pathogen presence). Rescue treatments are shown as light circles (no‐rescue treatment), diamonds (low‐rescue treatment), or dark triangles (high‐rescue treatment). Points are average values in each treatment (±SE). Arrows indicate days when rescue treatments were applied. Abundance was estimated by sampling ~18% of the population on each sampling day.
Demographic rescue falters when pathogens are present

December 2024

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1 Read

Catherine L. Searle

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Stephanie O. Gutierrez

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Ilinca I. Ciubotariu

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Mark R. Christie

As natural populations continue to decline globally, direct forms of intervention are increasingly necessary to prevent extinction. One type of intervention, known as demographic rescue, occurs when individuals are added directly to a population to increase abundance and ultimately prevent population extinction. However, the role of infectious disease in demographic rescue remains unknown. To examine the effects of pathogens on demographic rescue, we used a host–pathogen system with the aquatic crustacean Daphnia dentifera as the host and the fungus Metschnikowia bicuspidata as the pathogen. We constructed a randomized 3 × 2 factorial experiment with three rescue treatments (none, low, high) and two pathogen treatments (unexposed, exposed), where the pathogen was introduced via infected individuals during rescue events. We found that adding more individuals to demographically depressed populations increased abundance over the short term; highly supplemented populations initially had 62% more individuals than populations that had no introduced individuals. However, by the end of the experiment, populations that did not have any individuals introduced averaged 640% higher abundance than populations where infected individuals had been added. Thus, the introduction of infected individuals can result in worse demographic outcomes for populations than if no rescue is attempted.


Abundance‐mediated species interactions

December 2024

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169 Reads

Species interactions shape biodiversity patterns, community assemblage, and the dynamics of wildlife populations. Ecological theory posits that the strength of interspecific interactions is fundamentally underpinned by the population sizes of the involved species. Nonetheless, prevalent approaches for modeling species interactions predominantly center around occupancy states. Here, we use simulations to illuminate the inadequacies of modeling species interactions solely as a function of occupancy, as is common practice in ecology. We demonstrate erroneous inference into species interactions due to error in parameter estimates when considering species occupancy alone. To address this critical issue, we propose, develop, and demonstrate an abundance‐mediated interaction framework designed explicitly for modeling species interactions involving two or more species from detection/non‐detection data. We present Markov chain Monte Carlo (MCMC) samplers tailored for diverse ecological scenarios, including intraguild predation, disease‐ or predator‐mediated competition, and trophic cascades. Illustrating the practical implications of our approach, we compare inference from modeling the interactions in a three‐species network involving coyotes (Canis latrans), fishers (Pekania pennanti), and American marten (Martes americana) in North America as a function of occupancy states and as a function of abundance. When modeling interactions as a function of abundance rather than occupancy, we uncover previously unidentified interactions. Our study emphasizes that accounting for abundance‐mediated interactions rather than simple co‐occurrence patterns can fundamentally alter our comprehension of system dynamics. Through an empirical case study and comprehensive simulations, we demonstrate the importance of accounting for abundance when modeling species interactions, and we present a statistical framework equipped with MCMC samplers to achieve this paradigm shift in ecological research.



Conceptual model showing how environmental conditions affect forest attributes (Research Question 1), and how environmental conditions and forest attributes together affect forest biomass stocks and productivity (aboveground living biomass, aboveground dead biomass, fine root biomass, soil organic matter, aboveground biomass productivity, and litter production, Research Question 2). Clipart images from Flaticon.com.
Structural equation models (SEM) for (a) total biomass stock (the sum of aboveground living biomass, aboveground dead biomass, fine root biomass, and soil organic matter, in tons per hectare); (b) total aboveground biomass productivity (the sum of aboveground biomass productivity and annual litter production, in tons per hectare per year); and bar graphs showing beta coefficients of each factor on (c) total biomass stock and (d) total aboveground biomass productivity based on (a) and (b). Direct and indirect effects of environmental conditions (climatic wetness, stand age, and soil nutrients) and direct effects of structural attributes (i.e., tree density or stand basal area), taxonomic attributes (i.e., species richness per plot or rarefied species richness per 150 stems), and functional attributes (i.e., a community‐weighted mean [CWM] leaf nitrogen concentration [LNC] or wood density [WD]) were evaluated. In (a) and (b), for all significant relations (continuous black arrows), the beta coefficient and significance level are given (*p < 0.05, **p < 0.01, ***p < 0.001), and for all nonsignificant relations (gray, dashed arrows), no statistics are shown. R² values show the explained variance of the response variables. In (c) and (d), the filled bars show the direct effects of environmental conditions and forest attributes, and the hatched bars show the indirect effects of environmental conditions. For more statistics on the structural equation models, see Appendix S4: Table S3.
Structural equation models for (a) aboveground living biomass (AGBliving, in tons per hectare), (b) aboveground dead biomass (AGBdead, in tons per hectare), (c) fine root biomass in the top 15 cm of the soil (fine root, in tons per hectare), (d) soil organic matter in the top 15 cm of the soil (SOM, in tons per hectare), (e) aboveground biomass productivity (productivity, in tons per hectare per year), and (f) litter production (in tons per hectare per year). Direct and indirect effects of environmental conditions (climatic wetness, stand age, and soil nutrients) and direct effects of structural attributes (i.e., tree density or stand basal area), taxonomic attributes (i.e., species richness per plot or rarefied species richness per 150 stems), and functional attributes (i.e., a community‐weighted mean [CWM] leaf nitrogen concentration [LNC], leaf mass per area [LMA], or wood density [WD]) were evaluated. For all significant relations (continuous black arrows), the beta coefficient and significance level are given (*p < 0.05, **p < 0.01, ***p < 0.001), and for all nonsignificant relations (gray, dashed arrows), no statistics are shown. R² values show the explained variance of the response variables. For more statistics on the structural equation models, see Appendix S4: Table S3.
Beta coefficients of environmental conditions (climatic wetness, stand age and soil nutrients) and forest attributes (structural, taxonomic and functional attributes) on six compartments of biomass pools and productivity: (a) Aboveground living biomass (in tons per hectare), (b) aboveground dead biomass (in tons per hectare), (c) fine root biomass in the top 15 cm of the soil (in tons per hectare), (d) soil organic matter in the top 15 cm of the soil (in tons per hectare), (e) aboveground biomass productivity (in tons per hectare per year), and (f) litter production (in tons per hectare per year) based on the best models in structural equation models (Figure 3). The filled bars show the direct effects of environmental conditions and forest attributes, and the hatched bars show the indirect effects of environmental conditions.
Bivariate relationships of stand age, one soil nutrient (i.e., soil nitrogen [N] or phosphorus [P]), one structural attribute (i.e., stand basal area or tree density), one taxonomic attribute (i.e., species richness per plot or rarefied species richness per 150 stems), and one functional attribute (community‐weighted mean [CWM] leaf nitrogen concentration [LNC], leaf mass per area [LMA], or wood density [WD]) compared with aboveground living biomass (ABGliving, in tons per hectare), aboveground dead biomass (ABGdead, in tons per hectare), fine root biomass in the top 15 cm of the soil (fine root, in tons per hectare), soil organic matter in the top 15 cm of the soil (SOM, in tons per hectare), aboveground biomass productivity (productivity, in tons per hectare per year), and litter production (litter, in tons per hectare per year) in tropical dry (orange) and wet (blue) forests. The chosen environmental variables and forest attributes were the ones that were selected in the best model in the structural equation models (Figure 3). Note that these bivariate relationships are for illustration purposes only and may not necessarily provide the same results as in the structural equation models.
Drivers of biomass stocks and productivity of tropical secondary forests

December 2024

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124 Reads

Young tropical secondary forests play an important role in the local and global carbon cycles because of their large area and rapid biomass accumulation rates. This study examines how environmental conditions and forest attributes shape biomass compartments and the productivity of young tropical secondary forests. We compared 36 young secondary forest stands that differed in the time since agricultural land abandonment (2.3–3.6 years) from dry and wet regions in Ghana. We quantified biomass stocks in living and dead stems, roots, and soil, and aboveground biomass and litter productivity. We used structural equation models to evaluate how macroclimate, soil nutrients (N, P), and forest attributes (structure, diversity, and functional composition) affect ecosystem functioning. After three years of succession, tropical wet forests stored on average 115 t biomass ha⁻¹ (the sum of aboveground living and dead biomass, belowground fine root biomass, and soil organic matter), and dry forests stored 99 t ha⁻¹. These values represent 31% (in the wet forest) and 39% (in the dry forest) of the biomass compared with neighboring old‐growth forests. The majority of forest ecosystem biomass was stored in the soil (70%) and aboveground living vegetation (25%). Macroclimate strongly shaped forest attributes, which in turn determined biomass stocks and productivity. Soil phosphorus strongly increased litter production and soil organic matter, confirming that it is a limiting element in tropical ecosystems. Tree density and species diversity increased forest biomass stocks, suggesting crown packing and complementary resource use enhance forest functioning. A more conservative trait composition (high wood density) increased biomass stocks but reduced productivity, indicating that quantity, identity, and quality of species affect ecosystem functioning.


A theoretical framework predicting the effects of the number and identity of added nutrients over contrasting spatial scales reflected in α, β, and γ diversity metrics. “A, B, C, D, E” indicate different plant species in the same plot or species pool. (ME = microelements.)
Aboveground biomass, belowground biomass, and fraction of photosynthetically active radiation (PAR) reaching the surface in plots treated without N or P (N0P0), with N but without P (N1P0), and with N&P (N1P1) with varying numbers of different subsets of other added nutrients K, Ca, S, Mg, Fe, and microelement nutrients (ME). Significant regression relationships are shown, and the specific statistics are given in Appendix S1: Table S9. The number of plots per nutrient treatment are given in Appendix S1: Table S2. Data are given for 2018, 2019, and 2020.
The α diversity (ENSpie), β diversity (Bray–Curtis dissimilarity index), and γ diversity (total number of species in pool under a specific nutrient treatment) of the plant communities in plots treated without N or P (N0P0), with N but without P (N1P0), and with N and P (N1P1) with varying numbers of different subsets of other added nutrients K, Ca, S, Mg, Fe, and microelement nutrients (ME). Significant regression relationships are shown, and the specific statistics are given in Appendix S1: Table S10. The number of plots per nutrient treatment is given in Appendix S1: Table S2. Data are given for 2018, 2019, and 2020.
Structural equation models (SEMs) showing the direct and indirect effects of the number of added nutrients and of N and N&P addition on above‐ and belowground biomass, fraction of photosynthetically active radiation (PAR) to the surface, and (a) α diversity, (b) β diversity, and (c) γ diversity. Blue and orange arrows indicate positive and negative relationships, respectively. The thickness of the line is proportional to the strength of the relationship. Dotted lines indicate insignificant links between variables. The percentages associated with the response variables represent the proportion of variance explained by the dependent variables (model R²). Numbers on the arrows are the standardized path coefficients. Significant levels of the path coefficients are indicated as follows (^p < 0.1; *p < 0.05; ***p < 0.001). SEM parameters (χ², df, and p values) are presented beneath each model.
Nutrient effects on plant diversity loss arise from nutrient identity and decreasing niche dimension

December 2024

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157 Reads

Two hypotheses have been used to explain the loss of plant diversity with nutrient addition. The nutrient identity hypothesis posits that biodiversity loss is due to a specific limiting nutrient, such as nitrogen (N) or phosphorus (P), while the niche dimension hypothesis posits that adding a larger number of limiting nutrients, regardless of their identity, results in biodiversity loss. These two hypotheses have not previously been tested together simultaneously. Here, we conduct that analysis to enable their relative effect sizes to be compared. We manipulated the supply of eight nutrients in the same experimental meadow grassland site to isolate the effects of the identity of added nutrients versus the number of added nutrients on biodiversity loss. We found support for both hypotheses, with the largest negative effects on biodiversity measures being due to N, or N and P treatment, with additional more minor effects of the number of added nutrients. Structural equation models (SEMs) suggested both identity and number of added nutrients had direct negative effects on biodiversity, likely caused by species' innate ability to competitively respond to nutrients, especially in response to disease, herbivory, and stress. SEMs also suggested indirect effects arising from nutrient‐driven increases in aboveground biomass, which resulted in intensified competition for light and the competitive exclusion of short‐statured species. These effects were exacerbated by the nutrients N and P which caused a shift in biomass accumulation from belowground to aboveground. The results highlight that a multi‐nutrient perspective will improve our ability to effectively manage, monitor, and restore ecosystems.


Invertebrate herbivores influence seagrass wasting disease dynamics

December 2024

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29 Reads

Although invertebrate herbivores commonly impact terrestrial plant diseases by facilitating transmission of plant pathogens and increasing host susceptibility to infection via wounding, less is known about the role of herbivores in marine plant disease dynamics. Importantly, transmission via herbivores may not be required in the ocean since saline ocean waters support pathogen survival and transmission. Through laboratory experiments with eelgrass (Zostera marina), we showed that isopods (Pentidotea wosnesenskii) and snails (Lacuna spp.) created grazing scars that increased disease severity and thus indirectly facilitated transmission of Labyrinthula zosterae (Lz), a protist that causes seagrass wasting disease. Experiments also quantified different feeding preferences among herbivores: Amphipods (Ampithoe lacertosa) selectively consumed diseased eelgrass, while isopods and snails selectively grazed asymptomatic leaves, suggesting different herbivore taxa may have contrasting impacts on disease dynamics. Our experiments show no sign that herbivores directly vector Lz from diseased to asymptomatic eelgrass. However, we isolated live Lz from isopod, amphipod, and snail feces and detected Lz with quantitative polymerase chain reaction in amphipods and snails, suggesting that herbivores eating diseased eelgrass could pass the live pathogen. Finally, field surveys demonstrated a close association between seagrass wasting disease and invertebrate grazing scars; disease prevalence was 29 ± 4.7% (95% CI) higher on eelgrass leaves with herbivore scars. Collectively, these findings show that some herbivores can increase eelgrass disease risk by facilitating the spread of an important pathogen via wounding, but not via direct transmission. Thus, herbivores may play different roles in plant disease dynamics in terrestrial versus marine ecosystems depending on the pathogen's ability to survive and transmit without a vector.



Fish and invertebrate communities show greater day–night partitioning on tropical than temperate reefs

December 2024

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56 Reads

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1 Citation

Diel partitioning of animals within ecological communities is widely acknowledged, yet rarely quantified. Investigation of most ecological patterns and processes involves convenient daylight sampling, with little consideration of the contributions of nocturnal taxa, particularly in marine environments. Here we assess diel partitioning of reef faunal assemblages at a continental scale utilizing paired day and night visual census across 54 shallow tropical and temperate reefs around Australia. Day–night differences were most pronounced in the tropics, with fishes and invertebrates displaying distinct and opposing diel occupancy on coral reefs. Tropical reefs in daytime were occupied primarily by fishes not observed at night (64% of all species sighted across day and night, and 71% of all individuals). By night, substantial emergence of invertebrates not otherwise detected during sunlit hours occurred (56% of all species, and 45% of individuals). Nocturnal emergence of tropical invertebrates corresponded with significant declines in the richness and biomass of predatory and herbivorous diurnal fishes. In contrast, relatively small diel changes in fishes active on temperate reefs corresponded to limited nocturnal emergence of temperate invertebrates. This reduced partitioning may, at least in part, be a result of strong top‐down pressures from fishes on invertebrate communities, either by predation or competitive interference. For shallow reefs, the diel cycle triggers distinct emergence and retreat of faunal assemblages and associated trophic patterns and processes, which otherwise go unnoticed during hours of regular scientific monitoring. Improved understanding of reef ecology, and management of reef ecosystems, requires greater consideration of nocturnal interactions. Without explicit sampling of nocturnal patterns and processes, we may be missing up to half of the story when assessing ecological interactions.


Significant relationships (after Benjamini–Hochberg false discovery rate correction) between leaf traits and survival over ontogeny. Light green points and lines represent relationships using seedling trait data and dark green points and lines represent relationships using adult trait data. Shaded area represents the 95% CIs. See Appendix S1: Table S2 for details on slope coefficients and Table 1 for abbreviations and units. Illustrations by María Natalia Umaña.
From seedlings to adults: Linking survival and leaf functional traits over ontogeny

December 2024

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66 Reads

As long‐lived tropical trees grow into the multi‐layered canopy and face different environmental conditions, the relationships between leaf traits and whole‐plant survival can vary over ontogeny. We tested the strength and direction of the relationships between leaf traits and long‐term survival data across life stages for woody species from a subtropical forest in Puerto Rico. Trait–survival relationships were largely consistent across ontogeny with conservative traits leading to higher survival rates. The stage‐specific relationship R² increased by up to one order of magnitude compared to studies not considering ontogenetic trait variations. Stage‐specific traits were significant predictors of their corresponding stage‐specific survival: Seedlings traits were better predictors of seedling survival than adult traits, and adult traits were better predictors of maximum adult survival than seedling traits. Our results suggest that stage‐specific leaf traits reflect different strategies over ontogeny and can substantially improve predictability of survival models in tropical forests.


Study area and effect plots of pooled community response to the two land cover variables: proportion of impervious surface and canopy cover within 300 m radius of the light trap. Map of impervious surface (A) and canopy cover (B) within study area is shown at a 1‐m resolution. Fixed‐effect plots predicting the effect of impervious surface on pooled frass (C, D), micro‐moth (G, H), and macro‐moth peak (K) phenology. Fixed‐effect plots also display the effect of canopy cover on pooled frass (E, F), micro‐moth (I, J), and macro‐moth peak (L) phenology. Our results provide no evidence that these urbanization variables affect the timing of the pooled abundance of moths, as the 95% CIs of all estimated slope coefficients included a slope of zero. DOY, day of year.
Predicted values of the interaction between the two land cover variables and the two life history traits, while keeping all other variables constant. The date of peak abundance for macro‐moth species did not change across an impervious surface gradient regardless of trait values (A, B). Species with cooler temperature niches are predicted to have later peak abundance in sites that are surrounded by more canopy cover, while species with warmer temperature niches are predicted to have consistent peak abundance dates (C). Smaller species were most phenologically responsive to increases in canopy cover, while larger species have consistent dates of peak abundance regardless of the amount of canopy cover (D). Niche and body mass traits are continuous and values in the legend for those traits denote mean and 1 SD for those values. Values in the legend are then used for displaying predicted interactions. Points are jittered to better display data density. DOY, day of year.
Temperature niche and body size condition phenological responses of moths to urbanization in a subtropical city

December 2024

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39 Reads

Urbanization in temperate climates often advances the beginning and peak of biological events due to multiple factors, especially urban heat islands. However, the effect of urbanization on insect phenology remains understudied in more tropical areas, where temperature may be a weaker phenological cue. We surveyed moths across an urban gradient in a subtropical city weekly for a year to test how impervious surface and canopy cover impact phenology at the caterpillar and adult life stages. For macro‐moths, we also examine how these effects vary with life history traits. When pooling all individuals, we found no effect of urbanization proxy variables on timing of caterpillar or adult phenology. At the species‐specific level, we found timing of peak adult macro‐moths is influenced by canopy cover, which also interacts with two traits: temperature niche and body size. Cold‐adapted species delay timing of peak abundance in more shaded sites, while warm‐adapted species were not affected. Smaller species, associated with lower dispersal ability, were more phenologically sensitive to canopy cover than larger bodied species. These results highlight the importance of canopy cover within cities and its interaction with species' traits in mediating impact of urbanization on moth phenology in subtropical systems.


This figure explains how our analyses detected cross‐scale redundancy (CSR) and response diversity (RD). Panel (A) shows three of the 16 farms in our dataset (black dots), with the gold shaded areas representing the agricultural land cover surrounding each farm. The concentric circles show four of our 10 radii of analysis. For each of nine bee species, we used generalized additive models (GAMs) to test the relationship between that species' pollination and percent agricultural land cover at each radius. Panel (B) shows, for one bee species (Bombus griseocollis), how the pollination‐by‐agriculture relationship changes as agricultural land cover is measured at the four radii shown in panel (A). (In this example, the GAMs produced linear fits.) Each line in panel (B) translates to one point in panel (C), which shows the r² of the pollination‐by‐agriculture relationship across different radii. In panel (D), we remove the data points and show one line per bee species (Ab, Andrena bradleyi; Av, Andrena vicina; Bb, Bombus bimaculatus; Bg, Bombus griseocollis; Bi, Bombus impatiens; Ci, Colletes inaequalis; Cv, Colletes validus; Hl, Habropoda laboriosa; Xv, Xylocopa virginica). In panel (E), we chose the best radius for each bee species (where “best” means the radius corresponding with the highest r² in panel D) and used this radius to determine how the pollination of each bee species changed with percent agricultural land cover.
Relationship between bee species richness and cross‐scale redundancy (CSR, panel A) and response diversity (RD, panel B). Each data point is one simulation, varying in richness from three to 14. Blue and red points are data points that included or did not include Andrena bradleyi (a common, blueberry specialist), respectively. Solid black lines are generalized additive models (GAMs) fit to all data points. Blue and red lines are GAMs fit to the blue and red data points, respectively. In panel (A), the three lines are nearly indistinguishable. In panel (B), a superficially positive effect of richness on RD was parsimoniously explained by considering the identity effects of A. bradleyi.
We quantified the effects of each of the nine bee species we studied on cross‐scale redundancy (CSR), response diversity (RD), and stability. Panel (A) shows how species effects are defined, using Andrena bradleyi as an example. Points are separated by whether A. bradleyi is present (blue) or absent (red) from the simulated bee community. Open circles show the results of the 466 simulations, and the larger filled circles are group mean values taken across those simulations. The difference between the group means gives the change in stability (∆S$$ \Delta \mathrm{S} $$) and response diversity (∆RD)$$ \Delta \mathrm{RD}\Big) $$ associated with the presence of A. bradleyi. The same logic would apply to ∆CSR$$ \Delta \mathrm{CSR} $$, if CSR had been on the x‐axis. In panels (B) and (C), each data point is one bee species, with x‐ and y‐values calculated as shown in panel A (i.e., ∆RD$$ \Delta \mathrm{RD} $$ on the x‐axis and ∆S$$ \Delta \mathrm{S} $$ on the y‐axis). Points are sized by each species' abundance summed across sites; we stress that point sizes therefore do not reflect CIs. (B) The link between CSR and stability is weak: No species generates a strong signal of CSR, and to the extent there is any signal it is not correlated with stability. (C) In contrast, the link between RD and stability is strong, almost entirely because A. bradleyi strongly increases both.
Dominant species stabilize pollination services through response diversity, but not cross‐scale redundancy

November 2024

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28 Reads

Substantial evidence suggests that biodiversity can stabilize ecosystem function, but how it does this is less clear. In very general terms, the hypothesis is that biodiversity stabilizes function because having more species increases the role of compensatory dynamics, which occur when species in a community show different responses to the environment. Here, we focus on two forms of compensatory dynamics, cross‐scale redundancy (CSR) and response diversity (RD). CSR occurs when species respond to a disturbance at different scales such that scale‐specific disturbances do not negatively affect all species. RD occurs when species contributing to the same function show different responses to an environmental change. We developed a new analytical approach that can compare the strength of CSR and RD in the same dataset and used it to study native bee pollination of blueberry at 16 farms that varied in surrounding agricultural land use. We then asked whether CSR and RD among bee species are associated with the stability of blueberry pollination. Although CSR and RD were both present, only RD was associated with higher stability of pollination. Furthermore, the effects of RD on stability were due to a single widespread species, Andrena bradleyi, that is a specialist on blueberry and, unlike other bee species, was highly abundant at farms surrounded by intensive blueberry agriculture. Thus, the stabilizing effect we observed was attributable to an “identity effect” more than to species richness per se. Our results demonstrate how CSR and RD can be empirically measured and compared and highlight how the theoretical expectations of the biodiversity–ecosystem functioning field are not always upheld when confronted with real‐world data.


The daily energy intake of ruminant grazers can be estimated from the interaction between a cropping constraint that limits grazer intake on short‐grass sward and a digestion constraint that occurs because grass quality declines as grass matures (Fryxell, 1991; A). Both the cropping constraint and the digestion constraint are positively related to herbivore body size and optimal grass biomass for maximum daily energy intake of ruminant species scales with body size (B). Fire resets grass biomass to zero, and if grazers select grass patches based on optimal energy intake, then fire should cause temporal separation of species by body size (C). This occurs because smaller grazers initially utilize low‐biomass recent burns. Conversely, large grazers may benefit from feeding on high‐quality recently burned grass, but are constrained by the cropping rate. Species turnover occurs when grass regrows (gray shading) and moves between the optimum grass biomass of species of increasing size (C). Fire temporal effects should drive spatial separation as grazers select patches that burned at different times (D).
Burn preference indices (mean and 95% CI) from paired burned and unburned sites (A–D) and dynamic models (E–H) relative to grazer body mass for 0–3, 4–6, 7–9, and 10–12 months since fire, and daily image captures for focal grazer species as a function of time since fire (I–O) in Serengeti National Park, Tanzania. Burn preference indices are calculated as the number of images for individual species from burned sites divided by images from paired unburned sites within 3 km, plus images from burned sites, with a range from 0 to 1, where 0 is complete avoidance of burns and 1 is complete preference. Black lines represent relationships predicted by spatial lag (A), spatial error (B), and linear regression (E–H) models. Values for images per day (marginal means ± SE) are inferred from generalized linear mixed‐effects models, and each species' statistically significant (p < 0.05) peak utilization period (blue highlights) were identified using honestly significant difference (HSD)‐Tukey tests (Appendix S2: Table S1).
Time since fire interacts with herbivore intake rates to control herbivore habitat occupancy

November 2024

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49 Reads

Smaller grazers consistently show greater preference for recently burned patches than larger species. Energy optimization theory posits that this pattern is driven by small‐ versus large‐bodied herbivores seeking to maximize energy intake by choosing high‐quality recently burned grasses, or high‐quantity unburned grasses, respectively. We propose that if burn preference is driven by an energy‐maximization mechanism, then preference should change over time as grass regrows and progresses across the optimal feeding heights of herbivores of increasing body size. To test this, we used a camera trap array in the Serengeti National Park to quantify changes in the relative preference for burned patches of seven ruminant herbivore species. We compared observed patterns to simulation results from a grass production‐herbivore patch selection model. Burn preference and herbivore body size scaled negatively for 6 months after fire, but this relationship disappeared after 7 months when smaller species stopped selecting burns, and larger herbivores selected burns after 10 months, in a reversal of classic grazer succession. Simulations recreated the former but not the latter relationship, suggesting that an energy‐maximization mechanism can drive allometric scaling of burn preference immediately after fire, but over longer periods, grazer‐driven feedbacks are required to explain large herbivore burn preferences.



Energy transfer efficiency rather than productivity determines the strength of aquatic trophic cascades

November 2024

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203 Reads

Trophic cascades are important determinants of food web dynamics and functioning, yet mechanisms accounting for variation in trophic cascade strength remain elusive. Here, we used food chain models and a mesocosm experiment (phytoplankton–zooplankton–shrimp) to disentangle the relative importance of two energetic processes driving trophic cascades: primary productivity and energy transfer efficiency. Food chain models predicted that the strength of trophic cascades was increased as the energy transfer efficiency between herbivore and predator (predator efficiency) increased, while its relationship with primary productivity was relatively weak. These model predictions were confirmed by a mesocosm experiment, which showed that the strength of trophic cascade increased with predator efficiency but remained unaffected by nutrient supply rate or primary productivity. Combined, our results indicate that the efficiency of energy transfer along the food chain, rather than the total amount of energy fixed by primary producers, determines the strength of trophic cascades. Our study provides an integrative perspective to reconcile energetic and population dynamics in food webs, which has implications for both ecological research and ecosystem management.


The effect of priming cue intensity on growth per day prior to gall formation (A) and the interaction between priming cue intensity and galling on final height (B) and flower mass (C). Plant genotype type did not interact with priming cue intensity, so differences by genotype are omitted for clarity. Note that in (B) and (C) the effects of galling and cue strength are taken from the residuals of a model examining the effect of fly population, plant genotype, experimental week, and initial height (B only) or final height (C only) on either height or flower mass to control for these factors and more accurately reflect effect sizes (see Appendix S1: Section S1). *Indicate that galled and not galled plants differed within a priming cue treatment or that all plants in the full treatment had greater flower mass than control plants in brackets in (C). Bars (A) that do not share a letter were significantly different based on a Tukey‐Kramer comparison (α = 0.05). Error bars are one SE.
The proportion of Solidago altissima plants forming galls by (A) plant genotype, (B) ovipositing fly population, and (C) cue exposure. There were no interactions between the three manipulated variables, so they are each presented in separate panels. Treatments that do not share a letter were significantly different based on a Tukey–Kramer comparison (α = 0.05).
Latency (in seconds) to ovipuncture Solidago altissima buds, showing interactions by female fly population and plant genotype (A) or priming cue strength (B). *Indicates a significant difference between Bell0 and S110 genotypes. Bars of the same shade that do not share a letter are significantly different based on a Tukey–Kramer comparison (α = 0.05). S110 plants did not vary by fly population (A), and Bellefonte and State College females did not vary by priming cue strength (B); thus, letters for these bars are omitted. Error bars are one SE.
An insect pheromone primes tolerance of herbivory in goldenrod plants

November 2024

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47 Reads

Environmental cues that predict increased risk of herbivory can prime plant defenses; however, few studies have explored how such cues elicit broader plant responses, including potential effects on plant growth and other resource allocations that may affect tolerance to herbivore damage. We exposed goldenrod plants (Solidago altissima) to varying concentrations of the putative sex pheromone of a gall‐inducing herbivore, which has previously been implicated in defense priming. In experiments with two plant genotypes and three herbivore populations, any level of exposure to the pheromone enhanced tolerance of galling, rescuing flower production to levels observed for ungalled plants. Exposure to low doses of the pheromone elicited greater resistance to galling than exposure to high doses, with unexposed plants exhibiting intermediate resistance, suggesting a nonlinear relationship between exposure and defense priming. These findings suggest plant responses to environmental cues associated with biotic stressors are broader and more complex than previously appreciated.


(a) Population size shows an abrupt change starting in 2016. (b) While the per capita birth rate (dashed blue) does not change, the death rate (solid black) increases with population size until there is an abrupt increase in 2016. (c) Per capita birth and death rates (blue and black) versus total population size show rough estimates of carrying capacities before (K1) and after (K2) the abrupt change in mortality in 2016. Circles give 1983–2015 and crosses give 2016–2022, with fitted lines to the birth rate (blue) and the death rates (black). (d) Muriqui stochastic age‐ and sex‐structured population projections using either the full dataset (gray) or only data before the abrupt change, 1983–2015 (blue). Data are plotted as black circles for one randomly generated dataset in which birth years for individuals with uncertain births were selected from a uniform distribution between Bmin and Bmax. The 68% and 95% prediction intervals from 2000 stochastic simulations are given with dark and light shading, respectively.
Abrupt demographic change affects projected population size: Implications for an endangered species in a protected area

November 2024

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107 Reads

Understanding how demographic parameters change with density is essential for predicting the resilience of small populations. We use long‐term, individual‐based life history data from an isolated population of the Critically Endangered Northern Muriqui (Brachyteles hypoxanthus) inhabiting a 1000‐ha protected forest to evaluate density‐dependent demographic rates before and after an abrupt population decline. We found no effect of density on fertility or birth sex ratio, but mortality rates increased linearly with log density over the 33 years of population growth (1983–2015) and the subsequent 7 years of population decline (2016–2022). We used an age‐ and sex‐structured logistic growth model to project population sizes to 2060. Under the 1983–2015 demographic profile, the projected size was 500 individuals, but this dropped to 200 when including the abrupt change. Although the abrupt decline coincided with the end of a 2‐year drought and a yellow fever outbreak, we found no statistical effects of climate or disease on the continued population decline after 2016. However, the lower projected carrying capacity for muriquis is consistent with reduced forest productivity and increased predator pressures. These findings demonstrate the value of long‐term monitoring for identifying demographic changes that affect the sustainability of wildlife populations in small protected areas.


Possible effects of resource competition, herbivory, and allelopathy on the diversity–invasibility relationship. (a) Based on resource competition alone, a high species richness should decrease invasibility of the native community. In the presence of herbivory, (b) a decrease in herbivory damage on native plants and (c) an increase in herbivory damage on the invasive plant with species richness are likely to result in steeper diversity–invasibility relationships (solid and dashed red lines in d). Concomitantly, the intercepts of these relationships also change because damage of herbivory on native plants should increase invasibility, and damage of herbivory on the invader should decrease invasibility. In the presence of allelopathy, a decrease in allelopathic inhibition of native plants (solid line in e) or an increase in allelopathic inhibition of the invader (dashed line in f) is likely to result in steeper diversity–invasibility relationships (solid and dashed blue lines in g). These lines have different intercepts because negative allelopathic effects on native plants will increase invasibility, and negative allelopathic effects on the invader will decrease invasibility. However, when increased species richness dilutes the allelochemicals to concentrations that they are no longer effective (dashed‐dotted dark blue line in f), there should be a shallower diversity–invasibility relationship (dashed‐dotted dark blue line in g).
Relationships between transformed biomass of the invader Solidago canadensis and native plant species richness (a and b), and complementarity (c and d) and selection effects (e and f) on native biomass under different herbivory and allelopathy treatments in the years 2018 (square‐root‐transformed) and 2019 (log‐transformed). The lines are based on the simple regression analysis for each treatment separately. Solid lines indicate a statistically significant relationship (p < 0.05), and dashed lines indicate no significant relationships (p > 0.05). Red., reduced.
Structural equation models testing the effects of the allelopathy, herbivory, and species richness treatments on aboveground biomass (log10‐transformed) of the invader Solidago canadensis in the years (a) 2018 and (b) 2019. For species richness, we considered indirect effects on invader biomass via light interception by the community and the availabilities of nitrogen, phosphorus, and organic carbon in the soil, as well as a direct (or residual) effect that could not be explained by those variables. Indirect effects of species richness via herbivory or allelopathy were not included as these variables were manipulated separately. Solid lines indicate significant effects, and the thickness of each line is proportional to the magnitude of the corresponding path coefficient. The numbers in bracket are SEs. Red lines indicate significant positive effects (p < 0.05), blue lines indicate significant negative effects (p < 0.05), and gray lines indicate non‐significant effects (p > 0.05). GFI, goodness‐of‐fit test; RMSEA, root mean square error of approximation.
Herbivory and allelopathy contribute jointly to the diversity–invasibility relationship

November 2024

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91 Reads

Although herbivory and allelopathy play important roles in plant invasions, their roles in mediating the effect of plant diversity on invasion resistance remain unknown. In a 2‐year field experiment, we constructed native plant communities with four levels of species richness (one, two, four, and eight species) and used a factorial combination of insecticide and activated carbon applications to reduce herbivory and allelopathy, respectively. We then invaded the communities with the introduced plant Solidago canadensis L. One year after the start of the experiment, there was no statistically significant net effect of species richness on biomass of the invader. However, a structural equation model showed that species richness had a positive direct effect on invader biomass that was partially balanced out by a negative indirect effect of species richness via increased light interception. In the second year, the relationship between invader biomass and species richness was negative when we analyzed the treatment combination with herbivory and allelopathy separately. Therefore, we conclude that joint effects of herbivory and allelopathy may play major roles in driving the diversity–invasibility relationship and should be considered in future studies.



Eco‐phenotypic feedback loops differ in multistressor environments

November 2024

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83 Reads

Natural communities are exposed to multiple environmental stressors, which simultaneously impact the population and trait dynamics of the species embedded within these communities. Given that certain traits, such as body size, are known to rapidly respond to environmental change, and given that they can strongly influence the density of populations, this raises the question of whether the strength of the eco‐phenotypic feedback loop depends on the environment, and whether stressful environments would enhance or disrupt this feedback or causal linkage. We use two competing freshwater ciliates—Colpidium striatum and Paramecium aurelia—and expose their populations to a full‐factorial design of increasing salinity and temperature conditions as well as interspecific competition. We found that salinity, temperature, and competition significantly affected the density and cell size dynamics of both species. Cell size dynamics strongly influenced density dynamics; however, the strength of this eco‐phenotypic feedback loop weakened in stressful conditions and with interspecific competition. Our study highlights the importance of studying eco‐phenotypic dynamics in different environments comprising stressful abiotic conditions and species interactions.



A global database of butterfly species native distributions

November 2024

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52 Reads

Butterflies represent a diverse group of insects, playing key ecosystem roles such as pollination and their larval form engage in herbivory. Despite their importance, comprehensive global distribution data for butterfly species are lacking. This lack of comprehensive global data has hindered many large‐scale questions in ecology, evolutionary biology, and conservation at the regional and global scales. Here, I use an integrative workflow that combines occurrence records, alpha hull polygons, species' dispersal capacity, and natural habitat and environmental variables within a framework of species distribution models to generate species‐level native distributions for butterflies at a global scale in the contemporary period. The database releases native range maps for 10,372 extant species of butterflies at a spatial grain resolution of 5 arcmin (~10 km). This database has the potential to allow unprecedented large‐scale analyses in ecology, biogeography, and conservation of butterflies. The maps are available in the WGS84 coordinate reference system (EPSG:4326 code) and stored as vector polygons in the GEOPACKAGE format for maximum compression, allowing easy data manipulation using a standard computer. I additionally provide each species' spatial raster. All maps and R scripts are open access and available for download in Dryad and Zenodo, respectively, and are guided by FAIR (Findable, Accessible, Interoperable, and Reusable) data principles. By making these data available to the scientific community, I aim to advance the sharing of biological data to stimulate more comprehensive research in ecology, biogeography, and conservation of butterflies.


Illustration of the colonization–competition (C–C) trade‐off model in a multispecies (n = 6 species) system subject to multiple environmental drivers, including habitat size (S), resource productivity (R), and disturbance (D). Panel (a): initial species diversity pattern, containing two potential processes: superior competitors displacing inferior competitors via propagules (red arrows), and species colonizing empty sites via propagules (green arrows). Panel (b): increasing resource productivity (R) is assumed to enhance the colonization rates of all species (i.e., increasing the amount of propagules, such as seeds). Panel (c): reducing habitat size (S) by directly removing sites from the initial habitat in panel (a) (i.e., habitat loss). Panel (d): stochastic disturbance resulting in species mortality randomly across the habitat (red cross). Panel (e): biodiversity pattern as the product of interactions between these three drivers (represented by bold black cross). To establish the C–C trade‐off, we assume a competitive hierarchy by ranking the species from the best competitor (species 1) to the poorest (species 6), while setting species colonization rates as c1<c2<c3<…<c6$$ {c}_1<{c}_2<{c}_3<\dots <{c}_6 $$ (represented by the amount of propagules). All icons were obtained from https://www.ztupic.com/ and are in the public domain.
Separate effects of habitat size (S), resource productivity (R), and disturbance extent (D) on species diversity (a–c and g–i) and their relative abundances (d–f and j–l) in a simple multispecies community (n = 6), with a strict competitive hierarchy by ranking the species from the best competitor (species 1) to the poorest (species 6). Note that the x‐axes of S and R are not evenly scaled in order to make the zig‐zag pattern at S < 0.5 and R < 0.5 more distinguishable. Species diversity is characterized by both species richness and the inverse Simpson index. Species colonization rates (ci) are evenly spaced in increasing order in both ranges: (Case 1) ci∈E0.2,2$$ {c}_i\in E\left[0.2,2\right] $$ with all species coexisting in an intact habitat (i.e., R = S = 1 and D = 0); and (Case 2) ci∈E0.5,2.5$$ {c}_i\in E\left[0.5,2.5\right] $$ with species competitive exclusion occurring in an intact habitat. Panels (a, d, g, and j): S = 1 and D = 0; panels (b, e, h, and k): R = 1 & D = 0; and panels (c, f, i, and l): R = S = 1. Others: species mortality rates mi = 0.1.
Interactive effects of habitat size (S), resource productivity (R), and disturbance extent (D) on biodiversity, characterized by species richness and the inverse Simpson index. Panels (a, d, g, and j): D = 0; panels (b, e, h, and k): R = 1; and panels (c, f, i, and l): S = 1. Other parameter settings are the same as in Figure 2.
Interactive effects of habitat size (S), resource productivity (R), and disturbance extent (D) on biodiversity in a large initial community (n = 100), with a strict competitive hierarchy by ranking the species from the best competitor (species 1) to the poorest (species 100). Species diversity is characterized using species richness and the inverse Simpson index. Species colonization rates (ci) are evenly spaced in increasing order in both ranges: panels (a–f) ci∈E0.12,4.575$$ {c}_i\in E\left[0.12,4.575\right] $$ in Case 1 where all species can coexist in an intact habitat (i.e., R = S = 1 and D = 0); and panels (g–l) ci∈E0.5,2.5$$ {c}_i\in E\left[0.5,2.5\right] $$ in Case 2, with species competitive exclusion occurring in an intact habitat. Panels (a, d, g, and j): D = 0, panels (b, e, h, and k): R = 1, and panels (e, f, i, and l): S = 1. Other parameters: mi = 0.1.
Separate effects of habitat size (S), resource productivity (R), and disturbance extent (D) on species richness in a large initial community (n = 100), by varying other drivers at different levels (S, R or D = 0.2, 0.5, and 0.8). Panels (a, b, g, and h): D = 0, panels (c, d, i, and j): S = 1, and panels (e, f, k, and l): R = 1. Other parameter settings are the same as in Figure 4.
Complex interactive responses of biodiversity to multiple environmental drivers

November 2024

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354 Reads

There remains considerable doubt, debate, and confusion regarding how biodiversity responds to gradients of important environmental drivers, such as habitat size, resource productivity, and disturbance. Here we develop a simple but comprehensive theoretical framework based on competition–colonization multispecies communities to examine the separate and interactive effects of these drivers. Using both numerical simulations and analytical arguments, we demonstrate that the critical trade‐off between competitive and colonization ability can lead to complex nonlinear, zig‐zag responses in both species richness and the inverse Simpson index along gradients of these drivers. Furthermore, we find strong interactions between these drivers that can dramatically shift the response of biodiversity to these gradients. The zig‐zag patterns in biodiversity along ecological gradients, together with the strong interactions between the drivers, can explain the mixed findings of empirical studies and syntheses, thereby providing a new paradigm that can reconcile debates on the relationships between biodiversity and multiple drivers.


Tree demographic drivers across temperate rain forests, after accounting for site‐, species‐, and stem‐level attributes

November 2024

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53 Reads

Diverse drivers such as climate, soil fertility, neighborhood competition, and functional traits all contribute to variation in tree stem demographic rates. However, these demographic drivers operate at different scales, making it difficult to compare the relative importance of each driver on tree demography. Using c. 20,000 stem records from New Zealand's temperate rain forests, we analyzed the growth, recruitment, and mortality rates of 48 tree species and determined the relative importance of demographic drivers in a multilevel modeling approach. Tree species' maximum height emerged as the one most strongly associated with all demographic rates, with a positive association with growth rate and negative associations with recruitment and mortality rates. Climate, soil properties, neighborhood competition, stem size, and other functional traits also played significant roles in shaping demographic rates. Forest structure and functional composition were linked to climate and soil, with warm, dry climates and fertile soil associated with higher growth and recruitment rates. Neighborhood competition affected demographic rates depending on stem size, with smaller stems experiencing stronger negative effects, suggesting asymmetric competition where larger trees exert greater competitive effects on smaller trees. Our study emphasizes the importance of considering multiple drivers of demographic rates to better understand forest tree dynamics.


Journal metrics


4.4 (2023)

Journal Impact Factor™


21%

Acceptance rate


8.3 (2023)

CiteScore™


21 days

Submission to first decision


$3,780 / £2,520 / €3,150

Article processing charge

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