Assessing invertebrate herbivory in human‐modified tropical forest canopies

1. Studies on the effects of human‐driven forest disturbance usually focus on either biodiversity or carbon dynamics but much less is known about ecosystem processes that span different trophic levels. Herbivory is a fundamental ecological process for ecosystem functioning, but it remains poorly quantified in human‐modified tropical rainforests.
2. Here, we present the results of the largest study to date on the impacts of human disturbances on herbivory. We quantified the incidence (percentage of leaves affected) and severity (the percentage of leaf area lost) of canopy insect herbivory caused by chewers, miners, and gall makers in leaves from 1,076 trees distributed across 20 undisturbed and human‐modified forest plots in the Amazon.
3. We found that chewers dominated herbivory incidence, yet were not a good predictor of the other forms of herbivory at either the stem or plot level. Chewing severity was higher in both logged and logged‐and‐burned primary forests when compared to undisturbed forests. We found no difference in herbivory severity between undisturbed primary forests and secondary forests. Despite evidence at the stem level, neither plot‐level incidence nor severity of the three forms of herbivory responded to disturbance.
4. Synthesis. Our large‐scale study of canopy herbivory confirms that chewers dominate the herbivory signal in tropical forests, but that their influence on leaf area lost cannot predict the incidence or severity of other forms. We found only limited evidence suggesting that human disturbance affects the severity of leaf herbivory, with higher values in logged and logged‐and‐burned forests than undisturbed and secondary forests. Additionally, we found no effect of human disturbance on the incidence of leaf herbivory.

     

Intermediate disturbances are a key driver of long‐term tree demography across old‐growth temperate forests

Disentangling the relative influence of background versus disturbance related mortality on forest demography is crucial for understanding long‐term dynamics and predicting the influence of global change on forests. Quantifying the rates and drivers of tree demography requires direct observations of tree populations over multiple decades, yet such studies are rare in old‐growth forest, particularly in the temperate zone of Europe. We use multi‐decade (1980–2020) monitoring of permanent plots, including observations of mode of mortality and disturbance events, to quantify rates and drivers of tree demography across a network of old‐growth remnants in temperate mountain forests of Slovenia. Annual rates of mortality and recruitment varied markedly among sites and over time; census intervals that captured intermediate severity canopy disturbances caused subtle peaks in annual mortality (e.g., >2%/year), while rates of background mortality in non‐disturbed intervals averaged about 1%/year. Roughly half of the trees died from modes of mortality associated with disturbance (i.e., uprooting or snapped‐alive). Results of a Bayesian multilevel model indicate that beech (Fagus sylvatica) had a higher likelihood of disturbance related mortality compared to fir (Abies alba), which mainly died standing, and there was a notable increase in the odds of disturbance mortality with increasing diameter for all species. Annual recruitment rates were consistently low at sites (<0.5%) that lacked evidence of disturbance, but often exceeded 3% on sites with higher levels of past canopy mortality. Recruitment was dominated by beech on sites with more diffuse background mortality, while the less shade tolerant maple (Acer pseudoplatanus) recruited following known disturbance events. Our study highlights the important role of stand‐scale, partial canopy disturbance for long‐term forest demography. These results suggest that subtle climate‐driven changes in the regime of intermediate severity disturbances could have an important influence on future forest dynamics and warrant attention.     

Agent‐based modeling of the effects of forest dynamics,selective logging,and fragment size on epiphyte communities

1. Forest canopies play a crucial role in structuring communities of vascular epiphytes by providing substrate for colonization, by locally varying microclimate, and by causing epiphyte mortality due to branch or tree fall. However, as field studies in the three‐dimensional habitat of epiphytes are generally challenging, our understanding of how forest structure and dynamics influence the structure and dynamics of epiphyte communities is scarce.
2. Mechanistic models can improve our understanding of epiphyte community dynamics. We present such a model that couples dispersal, growth, and mortality of individual epiphytes with substrate dynamics, obtained from a three‐dimensional functional–structural forest model, allowing the study of forest–epiphyte interactions. After validating the epiphyte model with independent field data, we performed several theoretical simulation experiments to assess how (a) differences in natural forest dynamics, (b) selective logging, and (c) forest fragmentation could influence the long‐term dynamics of epiphyte communities.
3. The proportion of arboreal substrate occupied by epiphytes (i.e., saturation level) was tightly linked with forest dynamics and increased with decreasing forest turnover rates. While species richness was, in general, negatively correlated with forest turnover rates, low species numbers in forests with very‐low‐turnover rates were due to competitive exclusion when epiphyte communities became saturated. Logging had a negative impact on epiphyte communities, potentially leading to a near‐complete extirpation of epiphytes when the simulated target diameters fell below a threshold. Fragment size had no effect on epiphyte abundance and saturation level but correlated positively with species numbers.
4. Synthesis: The presented model is a first step toward studying the dynamic forest–epiphyte interactions in an agent‐based modeling framework. Our study suggests forest dynamics as key factor in controlling epiphyte communities. Thus, both natural and human‐induced changes in forest dynamics, for example, increased mortality rates or the loss of large trees, pose challenges for epiphyte conservation.

     

Two decades of demography reveals that seed and seedling transitions limit population persistence in a translocated shrub

Background and Aims

Olearia flocktoniae is an endangered shrub that was passively translocated from its natural ecosystem, where it has since gone extinct. This study aimed to determine sensitivities vital to populations persisting in human-created areas.

Methods

Population colonization, longevity and extinction were investigated over 20 years using 133 populations. Seed-bank longevity was determined from germination trials of seeds exhumed from extinct and extant sites via a 10-year glasshouse trial and by in situ sowing experiments. From 27 populations, 98 cohorts were followed and matrix models of transitions from seeds to adults were used to evaluate the intrinsic rate of population growth against disturbance histories. Ten populations (38 cohorts) with different disturbance histories were used to evaluate sensitivities in vital rates.

Key Results

Most populations had few individuals (∼30) and were transient (<5 years above ground). The intrinsic population growth rate was rarely >1 and all but two populations were extinct at year 20. Seeds were short-lived in situ. Although >1000 seeds per plant were produced annually in most populations, sensitivity analysis showed that the transition to the seed bank and the transition from the seed bank to seedlings are key vulnerabilities in the life-cycle.

Conclusions

Seedling establishment is promoted by recent disturbance. Increasing the number of disturbance events in populations, even severe disturbances that almost extirpate populations, significantly increases longer-term population persistence. Only populations that were disturbed annually survived the full 20 years of the study. The results show that translocated populations of O. flocktoniae will fail to persist without active management.     

Tree recruitment is determined by stand structure and shade tolerance with uncertain role of climate and water relations

1. Tree regeneration is a key process for long‐term forest dynamics, determining changes in species composition and shaping successional trajectories. While tree regeneration is a highly stochastic process, tree regeneration studies often cover narrow environmental gradients only, focusing on specific forest types or species in distinct regions. Thus, the larger‐scale effects of temperature, water availability, and stand structure on tree regeneration are poorly understood.
2. We investigated these effects in respect of tree recruitment (in‐growth) along wide environmental gradients using forest inventory data from Flanders (Belgium), northwestern Germany, and Switzerland covering more than 40 tree species. We employed generalized linear mixed models to capture the abundance of tree recruitment in response to basal area, stem density, shade casting ability of a forest stand as well as site‐specific degree‐day sum (temperature), water balance, and plant‐available water holding capacity. We grouped tree species to facilitate comparisons between species with different levels of tolerance to shade and drought.
3. Basal area and shade casting ability of the overstory had generally a negative impact on tree recruitment, but the effects differed between levels of shade tolerance of tree recruitment in all study regions. Recruitment rates of very shade‐tolerant species were positively affected by shade casting ability. Stem density and summer warmth (degree‐day sum) had similar effects on all tree species and successional strategies. Water‐related variables revealed a high degree of uncertainty and did not allow for general conclusions. All variables had similar effects independent of the varying diameter thresholds for tree recruitment in the different data sets.
4. Synthesis: Shade tolerance and stand structure are the main drivers of tree recruitment along wide environmental gradients in temperate forests. Higher temperature generally increases tree recruitment rates, but the role of water relations and drought tolerance remains uncertain for tree recruitment on cross‐regional scales.

     

Plant–plant interactions change during succession on nurse logs in a northern temperate rainforest

Plant–plant interactions change through succession from facilitative to competitive. At early stages of succession, early‐colonizing plants can increase the survival and reproductive output of other plants by ameliorating disturbance and stressful conditions. At later stages of succession, plant interactions are more competitive as plants put more energy toward growth and reproduction. In northern temperate rainforests, gap dynamics result in tree falls that facilitate tree regeneration (nurse logs) and bryophyte succession. How bryophyte‐tree seedling interactions vary through log succession remains unclear. We examined the relationships of tree seedlings, bryophyte community composition, bryophyte depth, and percent canopy cover in 166 1.0 m² plots on nurse logs and the forest floor in the Hoh rainforest in Washington, USA, to test the hypothesis that bryophyte‐tree seedling interactions change from facilitative to competitive as the log decays. Tree seedling density was highest on young logs with early‐colonizing bryophyte species (e.g., Rhizomnium glabrescens) and lowest on decayed logs with Hylocomium splendens, a long‐lived moss that reaches depths >20 cm. As a result, bryophyte depth increased with nurse log decay and was negatively associated with tree seedling density. Tree seedling density was 4.6× higher on nurse logs than on the forest floor, which was likely due to competitive exclusion by forest floor plants, such as H. splendens. Nurse logs had 17 species of bryophytes while the forest floor had six, indicating that nurse logs contribute to maintaining bryophyte diversity. Nurse logs enable both tree seedlings and smaller bryophyte species to avoid competition with forest floor plants, including the dominant bryophyte, H. splendens. H. splendens is likely a widespread driver of plant community structure given its dominance in northern temperate forests. Our findings indicate that plant–plant interactions shift with succession on nurse logs from facilitative to competitive and, thus, influence forest community structure and dynamics.     

Effects of forest disturbance on the fitness of an endemic rodent in a biodiversity hotspot

Praomys delectorum occurs abundantly in both disturbed and intact forests in the Ukaguru Mountains within the Eastern Arc Mountains (EAM), Morogoro, Tanzania. While previous studies have reported that anthropogenic disturbances such as grazing, wood cutting, and harvesting have a positive effect on the population density of P. delectorum, the impact of habitat disturbance on its demographic traits is still unknown. We performed a capture–mark–recapture study in both disturbed and intact forests from June 2018 to February 2020 in order to investigate the effects of habitat disturbance on abundance and two demographic traits: survival and maturation of P. delectorum in the Ukaguru Mountains. We found no variation in abundance or maturation between intact and disturbed forests, but habitat type did affect survival. However, this effect was sex‐dependent since female survival was higher in disturbed forests, while male survival remained similar across the two forest types potentially due to differences in predation pressure or food availability between the two habitats. Continuous demographic monitoring of P. delectorum in EAM is necessary given that the increasing human population surrounding the landscape is leading to higher deforestation rates and expansion of the pine plantation in the forest reserve.     

Traits explain invasion of alien plants into tropical rainforests

• 1. The establishment of new botanic gardens in tropical regions highlights a need for weed risk assessment tools suitable for tropical ecosystems. The relevance of plant traits for invasion into tropical rainforests has not been well studied.
• 2. Working in and around four botanic gardens in Indonesia where 590 alien species have been planted, we estimated the effect of four plant traits, plus time since species introduction, on: (a) the naturalization probability and (b) abundance (density) of naturalized species in adjacent native tropical rainforests; and (c) the distance that naturalized alien plants have spread from the botanic gardens.
• 3. We found that specific leaf area (SLA) strongly differentiated 23 naturalized from 78 non‐naturalized alien species (randomly selected from 577 non‐naturalized species) in our study. These trends may indicate that aliens with high SLA, which had a higher probability of naturalization, benefit from at least two factors when establishing in tropical forests: high growth rates and occupation of forest gaps. Naturalized aliens had high SLA and tended to be short. However, plant height was not significantly related to species'' naturalization probability when considered alongside other traits.
• 4. Alien species that were present in the gardens for over 30 years and those with small seeds also had higher probabilities of becoming naturalized, indicating that garden plants can invade the understorey of closed canopy tropical rainforests, especially when invading species are shade tolerant and have sufficient time to establish.
• 5. On average, alien species that were not animal dispersed spread 78 m further into the forests and were more likely to naturalize than animal‐dispersed species. We did not detect relationships between the measured traits and estimated density of naturalized aliens in the adjacent forests.
• 6. Synthesis: Traits were able to differentiate alien species from botanic gardens that naturalized in native forest from those that did not; this is promising for developing trait‐based risk assessment in the tropics. To limit the risk of invasion and spread into adjacent native forests, we suggest tropical botanic gardens avoid planting alien species with fast carbon capture strategies and those that are shade tolerant.

     

Developmental timing of extreme temperature events (heat waves) disrupts host–parasitoid interactions

1. When thermal tolerances differ between interacting species, extreme temperature events (heat waves) will alter the ecological outcomes. The parasitoid wasp Cotesia congregata suffers high mortality when reared throughout development at temperatures that are nonstressful for its host, Manduca sexta. However, the effects of short‐term heat stress during parasitoid development are unknown in this host–parasitoid system.
2. Here, we investigate how duration of exposure, daily maximum temperature, and the developmental timing of heat waves impact the performance of C. congregata and its host¸ M. sexta. We find that the developmental timing of short‐term heat waves strongly determines parasitoid and host outcomes.
3. Heat waves during parasitoid embryonic development resulted in complete wasp mortality and the production of giant, long‐lived hosts. Heat waves during the 1st‐instar had little effect on wasp success, whereas heat waves during the parasitoid''s nutritionally and hormonally critical 2nd instar greatly reduced wasp emergence and eclosion. The temperature and duration of heat waves experienced early in development determined what proportion of hosts had complete parasitoid mortality and abnormal phenotypes.
4. Our results suggest that the timing of extreme temperature events will be crucial to determining the ecological impacts on this host–parasitoid system. Discrepancies in thermal tolerance between interacting species and across development will have important ramifications on ecosystem responses to climate change.

     

Logging impacts on avian species richness and composition differ across latitudes and foraging and breeding habitat preferences

Understanding the causes underlying changes in species diversity is a fundamental pursuit of ecology. Animal species richness and composition often change with decreased forest structural complexity associated with logging. Yet differences in latitude and forest type may strongly influence how species diversity responds to logging. We performed a meta‐analysis of logging effects on local species richness and composition of birds across the world and assessed responses by different guilds (nesting strata, foraging strata, diet, and body size). This approach allowed identification of species attributes that might underlie responses to this anthropogenic disturbance. We only examined studies that allowed forests to regrow naturally following logging, and accounted for logging intensity, spatial extent, successional regrowth after logging, and the change in species composition expected due to random assembly from regional species pools. Selective logging in the tropics and clearcut logging in temperate latitudes caused loss of species from nearly all forest strata (ground to canopy), leading to substantial declines in species richness (up to 27% of species). Few species were lost or gained following any intensity of logging in lower‐latitude temperate forests, but the relative abundances of these species changed substantially. Selective logging at higher‐temperate latitudes generally replaced late‐successional specialists with early‐successional specialists, leading to no net changes in species richness but large changes in species composition. Removing less basal area during logging mitigated the loss of avian species from all forests and, in some cases, increased diversity in temperate forests. This meta‐analysis provides insights into the important role of habitat specialization in determining differential responses of animal communities to logging across tropical and temperate latitudes.     

Integrating omics to characterize eco‐physiological adaptations: How moose diet and metabolism differ across biogeographic zones

1. With accelerated land conversion and global heating at northern latitudes, it becomes crucial to understand, how life histories of animals in extreme environments adapt to these changes. Animals may either adapt by adjusting foraging behavior or through physiological responses, including adjusting their energy metabolism or both. Until now, it has been difficult to study such adaptations in free‐ranging animals due to methodological constraints that prevent extensive spatiotemporal coverage of ecological and physiological data.
2. Through a novel approach of combining DNA‐metabarcoding and nuclear magnetic resonance (NMR)‐based metabolomics, we aim to elucidate the links between diets and metabolism in Scandinavian moose Alces alces over three biogeographic zones using a unique dataset of 265 marked individuals.
3. Based on 17 diet items, we identified four different classes of diet types that match browse species availability in respective ecoregions in northern Sweden. Individuals in the boreal zone consumed predominantly pine and had the least diverse diets, while individuals with highest diet diversity occurred in the coastal areas. Males exhibited lower average diet diversity than females.
4. We identified several molecular markers indicating metabolic constraints linked to diet constraints in terms of food availability during winter. While animals consuming pine had higher lipid, phospocholine, and glycerophosphocholine concentrations in their serum than other diet types, birch‐ and willow/aspen‐rich diets exhibit elevated concentrations of several amino acids. The individuals with highest diet diversity had increased levels of ketone bodies, indicating extensive periods of starvation for these individuals.
5. Our results show how the adaptive capacity of moose at the eco‐physiological level varies over a large eco‐geographic scale and how it responds to land use pressures. In light of extensive ongoing climate and land use changes, these findings pave the way for future scenario building for animal adaptive capacity.

     

Modeling the demography of species providing extended parental care: A capture–recapture multievent model with a case study on polar bears (Ursus maritimus)

1. In species providing extended parental care, one or both parents care for altricial young over a period including more than one breeding season. We expect large parental investment and long‐term dependency within family units to cause high variability in life trajectories among individuals with complex consequences at the population level. So far, models for estimating demographic parameters in free‐ranging animal populations mostly ignore extended parental care, thereby limiting our understanding of its consequences on parents and offspring life histories.
2. We designed a capture–recapture multievent model for studying the demography of species providing extended parental care. It handles statistical multiple‐year dependency among individual demographic parameters grouped within family units, variable litter size, and uncertainty on the timing at offspring independence. It allows for the evaluation of trade‐offs among demographic parameters, the influence of past reproductive history on the caring parent''s survival status, breeding probability, and litter size probability, while accounting for imperfect detection of family units. We assess the model performance using simulated data and illustrate its use with a long‐term dataset collected on the Svalbard polar bears (Ursus maritimus).
3. Our model performed well in terms of bias and mean square error and in estimating demographic parameters in all simulated scenarios, both when offspring departure probability from the family unit occurred at a constant rate or varied during the field season depending on the date of capture. For the polar bear case study, we provide estimates of adult and dependent offspring survival rates, breeding probability, and litter size probability. Results showed that the outcome of the previous reproduction influenced breeding probability.
4. Overall, our results show the importance of accounting for i) the multiple‐year statistical dependency within family units, ii) uncertainty on the timing at offspring independence, and iii) past reproductive history of the caring parent. If ignored, estimates obtained for breeding probability, litter size, and survival can be biased. This is of interest in terms of conservation because species providing extended parental care are often long‐living mammals vulnerable or threatened with extinction.

     

Impacts of spruce budworm defoliation on the habitat of woodland caribou,moose, and their main predators

Forest logging has contributed to the decline of several woodland caribou populations by causing the fragmentation of mature coniferous stands. Such habitat alterations could be worsened by spruce budworm (SBW) outbreaks. Using 6201 vegetation plots from provincial inventories conducted after the last SBW outbreak (1968–1992) in boreal forests of Québec (Canada), we investigated the influence of SBW‐caused tree defoliation and mortality on understory vegetation layers relevant to woodland caribou and its main predators. We found a positive association between severe outbreaks and the cover of most groups of understory plant species, especially in stands that were dominated by balsam fir before the outbreak, where a high canopy openness particularly benefited relatively fast‐growing deciduous plants. Such increases in early successional vegetation could provide high‐quality forage for moose, which is likely to promote higher wolf densities and increase predation pressure on caribou. SBW outbreaks may thus negatively affect woodland caribou by increasing predation risk, the main factor limiting caribou populations in managed forests. For the near future, we recommend updating the criteria used to define critical caribou habitat to consider the potential impacts of spruce budworm defoliation.     

Joint control of plant ecological strategy by climate,regeneration mode, and ontogeny in Northeastern Chinese forests

1. Research on how plant ecological strategies (competitive, stress‐tolerant, or ruderal) vary within species may improve our understanding of plant and community responses to climate warming and also successional changes. With increasing temperature, the importance of ruderal (R) and stress tolerance (S) components is hypothesized to decrease, while the strength of the competitive (C) component should increase. Offshoots and younger plants are predicted to have greater R and smaller S components.
2. Leaf area, leaf dry matter content, and specific leaf area were measured for 1,344 forest plants belonging to 134 species in Liangshui and Fenglin Nature Reserves in Northeastern China, and C, R, and S scores calculated for each. Linear mixed effect models were used to assess how these indicators differed among study sites (n = 2), regeneration types, ontogenetic stages, and plant life forms. The two study sites have an average annual temperature difference of 0.675°C, simulating a temperature increase of 0.630°C due to climate warming.
3. Higher temperatures reduce low‐temperature stress and frost damage, which may explain the observed decrease in R and S scores; at the same time, plant competitive ability increased, as manifested by higher C scores. This effect was most pronounced for herbaceous plants, but nearly negligible as compared to the effect of regeneration type for trees and of ontogeny for woody species. Resprouting trees and younger woody plants had higher R scores and lower S scores, a sign of adaptation to high disturbance.
4. In this study, a small increase in mean annual temperature led to shifts in CSR strategy components for herbaceous species, without altering the vegetation type or community composition. Offshoots and younger plants had higher R and lower S scores, shedding light on similar changes in the ecological strategies of tree communities during secondary succession, such as the transition of Quercus mongolica coppices to forest and age‐related changes in Populus davidiana–Betula platyphylla forests.

     

Differential spatial responses of rodents to masting on forest sites with differing disturbance history

Mast seeding, the synchronized interannual variation in seed production of trees, is a well‐known bottom‐up driver for population densities of granivorous forest rodents. Such demographic effects also affect habitat preferences of the animals: After large seed production events, reduced habitat selectivity can lead to spillover from forest patches into adjacent alpine meadows or clear‐cuts, as has been reported for human‐impacted forests. In unmanaged, primeval forests, however, gaps created by natural disturbances are typical elements, yet it is unclear whether the same spillover dynamics occur under natural conditions. To determine whether annual variation in seed production drives spillover effects in naturally formed gaps, we used 14 years of small mammal trapping data combined with seed trap data to estimate population densities of Apodemus spp. mice and bank voles (Myodes glareolus) on 5 forest sites with differing disturbance history. The study sites, located in a forest dominated by European beech (Fagus sylvatica), Norway spruce (Picea abies), and silver fir (Abies alba), consisted of two primeval forest sites with small canopy gaps, two sites with larger gaps (after an avalanche event and a windthrow event), and a managed forest stand with closed canopy as a control. Hierarchical Bayesian N‐mixture models revealed a strong influence of seed rain on small rodent abundance, which were site‐specific for M. glareolus but not for Apodemus spp. Following years of moderate or low seed crop, M. glareolus avoided open habitat patches but colonized those habitats in large numbers after full mast events, suggesting that spillover events also occur in unmanaged forests, but not in all small rodents. The species‐ and site‐specific characteristics of local density responding to food availability have potentially long‐lasting effects on forest gap regeneration dynamics and should be addressed in future studies.     

Mass–abundance scaling in avian communities is maintained after tropical selective logging

1. Selective logging dominates forested landscapes across the tropics. Despite the structural damage incurred, selectively logged forests typically retain more biodiversity than other forest disturbances. Most logging impact studies consider conventional metrics, like species richness, but these can conceal subtle biodiversity impacts. The mass–abundance relationship is an integral feature of ecological communities, describing the negative relationship between body mass and population abundance, where, in a system without anthropogenic influence, larger species are less abundant due to higher energy requirements. Changes in this relationship can indicate community structure and function changes.
2. We investigated the impacts of selective logging on the mass–abundance scaling of avian communities by conducting a meta‐analysis to examine its pantropical trend. We divide our analysis between studies using mist netting, sampling the understory avian community, and point counts, sampling the entire community.
3. Across 19 mist‐netting studies, we found no consistent effects of selective logging on mass–abundance scaling relative to primary forests, except for the omnivore guild where there were fewer larger‐bodied species after logging. In eleven point‐count studies, we found a more negative relationship in the whole community after logging, likely driven by the frugivore guild, showing a similar pattern.
4. Limited effects of logging on mass–abundance scaling may suggest high species turnover in logged communities, with like‐for‐like replacement of lost species with similar‐sized species. The increased negative mass–abundance relationship found in some logged communities could result from resource depletion, density compensation, or increased hunting; potentially indicating downstream impacts on ecosystem functions.
5. Synthesis and applications. Our results suggest that size distributions of avian communities in logged forests are relatively robust to disturbance, potentially maintaining ecosystem processes in these forests, thus underscoring the high conservation value of logged tropical forests, indicating an urgent need to focus on their protection from further degradation and deforestation.

     

Grow slowly,persist, dominate—Explaining beech dominance in a primeval forest

Being able to persist in deep shade is an important characteristic of juvenile trees, often leading to a strong dominance of shade‐tolerant species in forests with low canopy turnover and a low disturbance rate. While leaf, growth, and storage traits are known to be key components of shade tolerance, their interplay during regeneration development and their influence on juveniles'' survival time remains unclear. We assessed the ontogenetic effects of these three traits on the survival time of beech (Fagus sylvatica), and Norway and sycamore maples (Acer pseudoplatanus, Acer platanoides) in a primeval beech forest. Biomass allocation, age, and content of nonstructural carbohydrates (NSC) were measured in the stems and roots of 289 seedlings and saplings in high‐ and low‐vitality classes. Saplings experienced a trade‐off between absolute growth rate (AGR) and storage (NSC) as the leaf area ratio (LAR) decreases with biomass development. High LAR but low AGR and low NSC corresponded to beech with a marked ability to persist in deep shade while awaiting canopy release. In turn, a comparably small LAR in combination with a high AGR and higher storage (NSC), as observed in Norway maple and sycamore maple, reduced sapling survival time, thus offering an explanation for beech dominance and maple disappearance in the undergrowth of old‐growth beech forests.     

Classifying behavior from short‐interval biologging data: An example with GPS tracking of birds

1. Recent advances in digital data collection have spurred accumulation of immense quantities of data that have potential to lead to remarkable ecological insight, but that also present analytic challenges. In the case of biologging data from birds, common analytical approaches to classifying movement behaviors are largely inappropriate for these massive data sets.
2. We apply a framework for using K‐means clustering to classify bird behavior using points from short time interval GPS tracks. K‐means clustering is a well‐known and computationally efficient statistical tool that has been used in animal movement studies primarily for clustering segments of consecutive points. To illustrate the utility of our approach, we apply K‐means clustering to six focal variables derived from GPS data collected at 1–11 s intervals from free‐flying bald eagles (Haliaeetus leucocephalus) throughout the state of Iowa, USA. We illustrate how these data can be used to identify behaviors and life‐stage‐ and age‐related variation in behavior.
3. After filtering for data quality, the K‐means algorithm identified four clusters in >2 million GPS telemetry data points. These four clusters corresponded to three movement states: ascending, flapping, and gliding flight; and one non‐moving state: perching. Mapping these states illustrated how they corresponded tightly to expectations derived from natural history observations; for example, long periods of ascending flight were often followed by long gliding descents, birds alternated between flapping and gliding flight.
4. The K‐means clustering approach we applied is both an efficient and effective mechanism to classify and interpret short‐interval biologging data to understand movement behaviors. Furthermore, because it can apply to an abundance of very short, irregular, and high‐dimensional movement data, it provides insight into small‐scale variation in behavior that would not be possible with many other analytical approaches.

     

Context‐dependent outcomes in a reproductive mutualism between two freshwater fish species

1. The development of encompassing general models of ecology is precluded by underrepresentation of certain taxa and systems. Models predicting context‐dependent outcomes of biotic interactions have been tested using plants and bacteria, but their applicability to higher taxa is largely unknown.
2. We examined context dependency in a reproductive mutualism between two stream fish species: mound nest‐building bluehead chub Nocomis leptocephalus and mountain redbelly dace Chrosomus oreas, which often uses N. leptocephalus nests for spawning. We hypothesized that increased predator density and decreased substrate availability would increase the propensity of C. oreas to associate with N. leptocephalus and decrease reproductive success of both species.
3. In a large‐scale in situ experiment, we manipulated egg predator density and presence of both symbionts (biotic context), and replicated the experiment in habitats containing high‐ and low‐quality spawning substrate (abiotic context).
4. Contradictory to our first hypothesis, we observed that C. oreas did not spawn without its host. The interaction outcome switched from commensalistic to mutualistic with changing abiotic and biotic contexts, although the net outcome was mutualistic.
5. The results of this study yielded novel insight into how context dependency operates in vertebrate mutualisms. Although the dilution effect provided by C. oreas positively influenced reproductive success of N. leptocephalus, it was not enough to overcome both egg predation and poor spawning habitat quality. Outcomes of the interaction may be ultimately determined by associate density. Studies of context dependency in vertebrate systems require detailed knowledge of species life‐history traits.

     

Disturbance and the (surprising?) role of ecosystem engineering in explaining spatial patterns of non‐native plant establishment

Different conceptions of disturbance differ in the degree to which they appeal to mechanisms that are general and equivalent, or species‐, functional group‐, or interaction‐specific. Some concepts of disturbance, for example, predict that soil disturbances and herbivory have identical impacts on species richness via identical mechanisms (reduction in biomass and in competition). An alternative hypothesis is that the specific traits of disturbance agents (small mammals) and plants differentially affect the richness or abundance of different plant groups. We tested these hypotheses on a degu (Octodon degus) colony in central Chile. We ask whether native and non‐native forbs respond differently to degu bioturbation on runways versus herbivory on grazing lawns. We ask whether this can explain the increase in non‐native plants on degu colonies. We found that biopedturbation did not explain the locations of non‐native plants. We did not find direct evidence of grazing increasing non‐native herbs either, but a grazing effect appears to be mediated by grass, which is the dominant cover. Further, we provide supplementary evidence to support our interpretation that a key mechanism of non‐native spread is the formation of dry soil conditions on grazing lawns. Thus, ecosystem engineering (alteration of soil qualities) may be an outcome of disturbances, in which each interacts with specific plant traits, to create the observed pattern of non‐native spread in the colony. Based on these results, we propose to extend Jentsch and White (Ecology, 100, 2019, e02734) concept of combined pulse/ disturbance events to the long‐term process duality of ecosystem engineering/ disturbance.