Mobility of moose—comparing the effects of wolf predation risk,reproductive status,and seasonality

In a predator–prey system, prey species may adapt to the presence of predators with behavioral changes such as increased vigilance, shifting habitats, or changes in their mobility. In North America, moose (Alces alces) have shown behavioral adaptations to presence of predators, but such antipredator behavioral responses have not yet been found in Scandinavian moose in response to the recolonization of wolves (Canis lupus). We studied travel speed and direction of movement of GPS‐collared female moose (n = 26) in relation to spatiotemporal differences in wolf predation risk, reproductive status, and time of year. Travel speed was highest during the calving (May–July) and postcalving (August–October) seasons and was lower for females with calves than females without calves. Similarly, time of year and reproductive status affected the direction of movement, as more concentrated movement was observed for females with calves at heel, during the calving season. We did not find support for that wolf predation risk was an important factor affecting moose travel speed or direction of movement. Likely causal factors for the weak effect of wolf predation risk on mobility of moose include high moose‐to‐wolf ratio and intensive hunter harvest of the moose population during the past century.     

Identifying “vital attributes” for assessing disturbance–recovery potential of seafloor communities

Despite a long history of disturbance–recovery research, we still lack a generalizable understanding of the attributes that drive community recovery potential in seafloor ecosystems. Marine soft‐sediment ecosystems encompass a range of heterogeneity from simple low‐diversity habitats with limited biogenic structure, to species‐rich systems with complex biogenic habitat structure. These differences in biological heterogeneity are a product of natural conditions and disturbance regimes. To search for unifying attributes, we explore whether a set of simple traits can characterize community disturbance–recovery potential using seafloor patch‐disturbance experiments conducted in two different soft‐sediment landscapes. The two landscapes represent two ends of a spectrum of landscape biotic heterogeneity in order to consider multi‐scale disturbance–recovery processes. We consider traits at different levels of biological organization, from the biological traits of individual species, to the traits of species at the landscape scale associated with their occurrence across the landscape and their ability to be dominant. We show that in a biotically heterogeneous landscape (Kawau Bay, New Zealand), seafloor community recovery is stochastic, there is high species turnover, and the landscape‐scale traits are good predictors of recovery. In contrast, in a biotically homogeneous landscape (Baltic Sea), the options for recovery are constrained, the recovery pathway is thus more deterministic and the scale of recovery traits important for determining recovery switches to the individual species biological traits within the disturbed patch. Our results imply that these simple, yet sophisticated, traits can be effectively used to characterize community recovery potential and highlight the role of landscapes in providing resilience to patch‐scale disturbances.     

Behavioral “bycatch” from camera trap surveys yields insights on prey responses to human‐mediated predation risk

Human disturbance directly affects animal populations and communities, but indirect effects of disturbance on species behaviors are less well understood. For instance, disturbance may alter predator activity and cause knock‐on effects to predator‐sensitive foraging in prey. Camera traps provide an emerging opportunity to investigate such disturbance‐mediated impacts to animal behaviors across multiple scales. We used camera trap data to test predictions about predator‐sensitive behavior in three ungulate species (caribou Rangifer tarandus; white‐tailed deer, Odocoileus virginianus; moose, Alces alces) across two western boreal forest landscapes varying in disturbance. We quantified behavior as the number of camera trap photos per detection event and tested its relationship to inferred human‐mediated predation risk between a landscape with greater industrial disturbance and predator activity and a “control” landscape with lower human and predator activity. We also assessed the finer‐scale influence on behavior of variation in predation risk (relative to habitat variation) across camera sites within the more disturbed landscape. We predicted that animals in areas with greater predation risk (e.g., more wolf activity, less cover) would travel faster past cameras and generate fewer photos per detection event, while animals in areas with less predation risk would linger (rest, forage, investigate), generating more photos per event. Our predictions were supported at the landscape‐level, as caribou and moose had more photos per event in the control landscape where disturbance‐mediated predation risk was lower. At a finer‐scale within the disturbed landscape, no prey species showed a significant behavioral response to wolf activity, but the number of photos per event decreased for white‐tailed deer with increasing line of sight (m) along seismic lines (i.e., decreasing visual cover), consistent with a predator‐sensitive response. The presence of juveniles was associated with shorter behavioral events for caribou and moose, suggesting greater predator sensitivity for females with calves. Only moose demonstrated a positive behavioral association (i.e., longer events) with vegetation productivity (16‐day NDVI), suggesting that for other species bottom‐up influences of forage availability were generally weaker than top‐down influences from predation risk. Behavioral insights can be gleaned from camera trap surveys and provide complementary information about animal responses to predation risk, and thus about the indirect impacts of human disturbances on predator–prey interactions.     

Presence of moose (<Emphasis Type="Italic">Alces alces</Emphasis>) in Southeastern Germany

Germany is situated on the western border of the distribution area of moose (Alces alces) in Europe. In the past, the number of moose sightings has been very limited and irregular in the state of Bavaria (SE Germany). Nearby populations of moose are located in neighboring countries to the east of Germany, in the Czech Republic, and Poland. A monitoring system for moose was established following a considerable increase in moose observations in Bavaria in 2007. As no reproduction has yet been recorded, moose observation has so far been attributed only to migrating individuals. Cows are more often registered than bulls (27 of 35 observations between 2004 and 2008). The reasons for the increase in migratory activities of moose have not yet been identified.     

“Micropersonality” traits and their implications for behavioral and movement ecology research

1. Many animal personality traits have implicit movement‐based definitions and can directly or indirectly influence ecological and evolutionary processes. It has therefore been proposed that animal movement studies could benefit from acknowledging and studying consistent interindividual differences (personality), and, conversely, animal personality studies could adopt a more quantitative representation of movement patterns.
2. Using high‐resolution tracking data of three‐spined stickleback fish (Gasterosteus aculeatus), we examined the repeatability of four movement parameters commonly used in the analysis of discrete time series movement data (time stationary, step length, turning angle, burst frequency) and four behavioral parameters commonly used in animal personality studies (distance travelled, space use, time in free water, and time near objects).
3. Fish showed repeatable interindividual differences in both movement and behavioral parameters when observed in a simple environment with two, three, or five shelters present. Moreover, individuals that spent less time stationary, took more direct paths, and less commonly burst travelled (movement parameters), were found to travel farther, explored more of the tank, and spent more time in open water (behavioral parameters).
4. Our case study indicates that the two approaches—quantifying movement and behavioral parameters—are broadly equivalent, and we suggest that movement parameters can be viewed as “micropersonality” traits that give rise to broad‐scale consistent interindividual differences in behavior. This finding has implications for both personality and movement ecology research areas. For example, the study of movement parameters may provide a robust way to analyze individual personalities in species that are difficult or impossible to study using standardized behavioral assays.

     

The declining occurrence of moose (Alces alces) at the southernmost edge of its range raise conservation concerns

The border region between Austria, the Czech Republic, and Germany harbors the most south‐western occurrence of moose in continental Europe. The population originated in Poland, where moose survived, immigrated from former Soviet Union or were reintroduced after the Second World War expanded west‐ and southwards. In recent years, the distribution of the nonetheless small Central European population seems to have declined, necessitating an evaluation of its current status. In this study, existing datasets of moose observations from 1958 to 2019 collected in the three countries were combined to create a database totaling 771 records (observations and deaths). The database was then used to analyze the following: (a) changes in moose distribution, (b) the most important mortality factors, and (c) the availability of suitable habitat as determined using a maximum entropy approach. The results showed a progressive increase in the number of moose observations after 1958, with peaks in the 1990s and around 2010, followed by a relatively steep drop after 2013. Mortality within the moose population was mostly due to human interactions, including 13 deadly wildlife‐vehicle collisions, particularly on minor roads, and four animals that were either legally culled or poached. Our habitat model suggested that higher altitudes (ca. 700–1,000 m a.s.l.), especially those offering wetlands, broad‐leaved forests and natural grasslands, are the preferred habitats of moose whereas steep slopes and areas of human activity are avoided. The habitat model also revealed the availability of large core areas of suitable habitat beyond the current distribution, suggesting that habitat was not the limiting factor explaining the moose distribution in the study area. Our findings call for immediate transboundary conservation measures to sustain the moose population, such as those aimed at preventing wildlife‐vehicle collisions and illegal killings. Infrastructure planning and development activities must take into account the habitat requirements of moose.     

How patrollers set foraging direction in harvester ants

Recruitment to food or nest sites is well known in ants; the recruiting ants lay a chemical trail that other ants follow to the target site, or they walk with other ants to the target site. Here we report that a different process determines foraging direction in the harvester ant Pogonomyrmex barbatus. Each day, the colony chooses from among up to eight distinct foraging trails; colonies use different trails on different days. Here we show that the patrollers regulate the direction taken by foragers each day by depositing Dufour's secretions onto a sector of the nest mound about 20 cm long and leading to the beginning of a foraging trail. The patrollers do not recruit foragers all the way to food sources, which may be up to 20 m away. Fewer foragers traveled along a trail if patrollers had no access to the sector of the nest mound leading to that trail. Adding Dufour's gland extract to patroller-free sectors of the nest mound rescued foraging in that direction, while poison gland extract did not. We also found that in the absence of patrollers, most foragers used the direction they had used on the previous day. Thus, the colony's 30-50 patrollers act as gatekeepers for thousands of foragers and choose a foraging direction, but they do not recruit and lead foragers all the way to a food source.     

The traits of “trait ecologists”: An analysis of the use of trait and functional trait terminology

Trait and functional trait approaches have revolutionized ecology improving our understanding of community assembly, species coexistence, and biodiversity loss. Focusing on traits promotes comparability across spatial and organizational scales, but terms must be used consistently. While several papers have offered definitions, it remains unclear how ecologists operationalize “trait” and “functional trait” terms. Here, we evaluate how researchers and the published literatures use these terms and explore differences among subdisciplines and study systems (taxa and biome). By conducting both a survey and a literature review, we test the hypothesis that ecologists’ working definition of “trait” is adapted or altered when confronting the realities of collecting, analyzing and presenting data. From 486 survey responses and 712 reviewed papers, we identified inconsistencies in the understanding and use of terminology among researchers, but also limited inclusion of definitions within the published literature. Discrepancies were not explained by subdiscipline, system of study, or respondent characteristics, suggesting there could be an inconsistent understanding even among those working in related topics. Consistencies among survey responses included the use of morphological, phonological, and physiological traits. Previous studies have called for unification of terminology; yet, our study shows that proposed definitions are not consistently used or accepted. Sources of disagreement include trait heritability, defining and interpreting function, and dealing with organisms in which individuals are not clearly recognizable. We discuss and offer guidelines for overcoming these disagreements. The diversity of life on Earth means traits can represent different features that can be measured and reported in different ways, and thus, narrow definitions that work for one system will fail in others. We recommend ecologists embrace the breadth of biodiversity using a simplified definition of “trait” more consistent with its common use. Trait‐based approaches will be most powerful if we accept that traits are at least as diverse as trait ecologists.     

Harvest‐induced phenotypic selection in an island population of moose,Alces alces

Empirical evidence strongly indicates that human exploitation has frequently led to rapid evolutionary changes in wild populations, yet the mechanisms involved are often poorly understood. Here, we applied a recently developed demographic framework for analyzing selection to data from a 20‐year study of a wild population of moose, Alces alces. In this population, a genetic pedigree has been established all the way back to founders. We demonstrate harvest‐induced directional selection for delayed birth dates in males and reduced body mass as calf in females. During the study period, birth date was delayed by 0.81 days per year for both sexes, whereas no significant changes occurred in calf body mass. Quantitative genetic analyses indicated that both traits harbored significant additive genetic variance. These results show that selective harvesting can induce strong selection that oppose natural selection. This may cause evolution of less favorable phenotypes that become maladaptive once harvesting ceases.     

Carbon “quantum” dots for bioapplications

Carbon “quantum” dots or carbon dots (CDots) exploit and enhance the intrinsic photoexcited state properties and processes of small carbon nanoparticles via effective nanoparticle surface passivation by chemical functionalization with organic species. The optical properties and photoinduced redox characteristics of CDots are competitive to those of established conventional semiconductor quantum dots and also fullerenes and other carbon nanomaterials. Highlighted here are major advances in the exploration of CDots for their serving as high-performance yet nontoxic fluorescence probes for one- and multi-photon bioimaging in vitro and in vivo, and for their uniquely potent antimicrobial function to inactivate effectively and efficiently some of the toughest bacterial pathogens and viruses under visible/natural or ambient light conditions. Opportunities and challenges in the further development of the CDots platform and related technologies are discussed.     

The non-impact of hunting on moose <Emphasis Type="Italic">Alces alces</Emphasis> movement,diurnal activity,and activity range

Previous studies on moose Alces alces have suggested that interactions with humans may trigger anti-predator behaviors and generate a demographical cost. Therefore, we hypothesized that disturbances from small and big game hunting may have negative effects on moose movements, diurnal activity, and activity range. Using location data from 64 moose equipped with GPS collars from three populations (Low Alpine, Inland, Coastal) with different temporal human presence and spatial accessibility, we evaluated the impact of hunting on moose activity rhythms. On average, female moose in the low human population density (Low Alpine) area (<0.5/km²) had significantly lower movement rates during moose hunting season, but variation in movement rates among individuals were higher compared with female moose in regions with denser human populations (6–24/km²). We found no evidence that reproductive status influenced female moose sensitivity to disturbance. As expected, females used smaller activity ranges and were less active nocturnally than males. The high within-group variation suggests that current hunting disturbance levels do not alter moose population behavior in general. Our data indicate that alterations in movement were related to rutting activity, not human disturbance induced by hunting. In line with behavioral theory, our study suggests that some individuals were more sensitive to hunting disturbance than the general population. Our work suggests that individual moose may perceive human predation risk to be similar to other predation risks.     

Influence of “protective” symbionts throughout the different steps of an aphid–parasitoid interaction

Microbial associates are widespread in insects, some conferring a protection to their hosts against natural enemies like parasitoids. These protective symbionts may affect the infection success of the parasitoid by modifying behavioral defenses of their hosts, the development success of the parasitoid by conferring a resistance against it or by altering life-history traits of the emerging parasitoids. Here, we assessed the effects of different protective bacterial symbionts on the entire sequence of the host-parasitoid interaction (i.e., from parasitoid attack to offspring emergence) between the pea aphid, Acyrthosiphon pisum, and its main parasitoid, Aphidius ervi and their impacts on the life-history traits of the emerging parasitoids. To test whether symbiont-mediated phenotypes were general or specific to particular aphid–symbiont associations, we considered several aphid lineages, each harboring a different strain of either Hamiltonella defensa or Regiella insecticola, two protective symbionts commonly found in aphids. We found that symbiont species and strains had a weak effect on the ability of aphids to defend themselves against the parasitic wasps during the attack and a strong effect on aphid resistance against parasitoid development. While parasitism resistance was mainly determined by symbionts, their effects on host defensive behaviors varied largely from one aphid–symbiont association to another. Also, the symbiotic status of the aphid individuals had no impact on the attack rate of the parasitic wasps, the parasitoid emergence rate from parasitized aphids nor the life-history traits of the emerging parasitoids. Overall, no correlations between symbiont effects on the different stages of the host–parasitoid interaction was observed, suggesting no trade-offs or positive associations between symbiont-mediated phenotypes. Our study highlights the need to consider various sequences of the host-parasitoid interaction to better assess the outcomes of protective symbioses and understand the ecological and evolutionary dynamics of insect–symbiont associations.     

“Ecology of fear” in ungulates: Opportunities for improving conservation

Because ungulates are important contributors to ecosystem function, understanding the “ecology of fear” could be important to the conservation of ecosystems. Although studying ungulate ecology of fear is common, knowledge from ungulate systems is highly contested among ecologists. Here, we review the available literature on the ecology of fear in ungulates to generalize our current knowledge and how we can leverage it for conservation. Four general focus areas emerged from the 275 papers included in our literature search (and some papers were included in multiple categories): behavioral responses to predation risk (79%), physiological responses to predation risk (15%), trophic cascades resulting from ungulate responses to predation risk (20%), and manipulation of predation risk (1%). Of papers focused on behavior, 75% were about movement and habitat selection. Studies were biased toward North America (53%), tended to be focused on elk (Cervus canadensis; 29%), and were dominated by gray wolves (40%) or humans (39%) as predators of interest. Emerging literature suggests that we can utilize predation risk for conservation with top‐down (i.e., increasing predation risk) and bottom‐up (i.e., manipulating landscape characteristics to increase risk or risk perception) approaches. It is less clear whether fear‐related changes in physiology have population‐level fitness consequences or cascading effects, which could be fruitful avenues for future research. Conflicting evidence of trait‐mediated trophic cascades might be improved with better replication across systems and accounting for confounding effects of ungulate density. Improving our understanding of mechanisms modulating the nature of trophic cascades likely is most important to ensure desirable conservation outcomes. We recommend future work embrace the complexity of natural systems by attempting to link together the focal areas of study identified herein.     

Age and sex-specific variation in detectability of moose (<Emphasis Type="Italic">Alces alces</Emphasis>) during the hunting season: implications for population monitoring

The sustainability of wild ungulate harvests can be greatly enhanced if monitoring data are available to permit an adaptive management approach. Utilising data provided by hunters is potentially the most cost-effective approach. In Scandinavia, observations recorded by moose (Alces alces) hunters provide a range of indices of population density, composition and reproductive performance. These are routinely used in practical management, but there are still many questions about their accuracy and precision. In this study, we availed of the fact that virtually all individual moose on the island of Vega in central Norway were marked during the period 1992–2005. Thus, we were able to compare the observation indices provided by hunters to the known size and composition of the population. The results indicate that the hunter observation indices provided accurate estimates of variation in moose cow recruitment and twinning rates. The estimates of sex ratio closely followed the pattern of annual variation but showed a consistent pattern of over-estimating the proportion of males. Thus, males seem to visually expose themselves more often for hunters than do females. The density index, measured as number of moose seen per hunter day did not to the same extent follow the variation in population density, possibly because moose were more naive in the first years after hunting was introduced or because of reduced detection conditions due to increasing scrub and tree encroachment during the study period. The results are discussed in light of their application to management and the evolutionary pressures in moose anti-predator behaviour.     

Behavior of Moose Relative to a Road Network

Abstract Roads often negatively affect terrestrial wildlife, via habitat loss or fragmentation, noise, and direct mortality. We studied moose (Alces alces) behavior relative to a road network, in an area with a history of moose-vehicle accidents, to determine when moose were crossing roadways or using areas near roads and to investigate if environmental factors were involved in this behavior. We tracked 47 adult moose with Global Positioning System collars in a study area crossed by highways and forest roads. We hypothesized that moose would avoid crossing roads but would make occasional visits to roadsides to feed on sodium-rich vegetation and avoid biting insects. Further, we expected moose avoidance to be greater for highways than forest roads. We recorded 196,710 movement segments but only observed 328 highway and 1,172 forest-road crossings (16 and 10 times lower than expected by chance). Moose usually avoided road proximity up to ≥500 m on each side but 20% of collared moose made visits to areas within 50 m of highways, which might have resulted from moose searching for sodium in vegetation and roadside salt pools. In fact, vegetation along highways had higher sodium concentrations and was browsed in similar proportions to vegetation in adjacent forest, despite moose avoidance of these zones. Moose, however, did not use areas near roads more during periods of biting insect abundance. Our results supported the hypothesis of scale-dependent selection by moose; avoidance of highways at a coarse scale may confer long-term benefits, whereas selection of highway corridors at finer scales may be part of a strategy to overcome short-term limiting factors such as sodium deficiency. We found a positive relationship between home-range size and the proportion of road axes they contained, suggesting that moose either compensated for habitat loss or made specific movements along highways to gather sodium. The presence of sodium along highways likely increases moose-vehicle accident risks. Removal of salt pools or use of a de-icing salt other than sodium chloride should render highway surroundings less attractive to moose.     

The time-energy budget of a moose

As the first step in developing an optimal foraging model for moose (Belovsky 1978), a time-energy budget for a moose was developed. For the summer period (late May through early September) at Isle Royale National Park, Michigan, we made various behavioral observations on feeding moose and inventoried their food resources. Employing these various sets of data, we were able to determine the quantity of each food consumed by moose, the time required to acquire the foods, preferences for different food plants, and differences demonstrated by the sexes. In addition to this information, we collected data on the moose's ability to utilize each of the forage plants and the population densities of moose in different years (1972–1974) in two forest types. This enabled us to examine energy flow within each of the populations and determine in what manner energy limits the moose population.     

Reconstructing landscapes of ungulate parturition and predation using vegetation phenology

Enhanced vegetation index (EVI) data can be used to identify and define the space in which ungulates practice parturition and encounter predation. This study explores the use of EVI data to identify landscapes linked to ungulate parturition and predation events across space, time, and environmental conditions. As a case study, we used the moose population (Alces alces) of northern Minnesota in the USA. Using remotely sensed EVI data rasters and global positioning system collar data, we quantified how vegetation phenology and moose movement shaped the births and predation of 52 moose calves from 2013 to 2020 on or adjacent to the Grand Portage Indian Reservation. The known sources of predation were American black bears (Ursus americanus, n = 22) and gray wolves (Canis lupus, n = 28). Satellite-derived data summarizing seasonal landscape features at the local level revealed that landscape heterogeneity use by moose can help to quantitatively identify landscapes of parturition and predation in space and time across large areas. Vegetation phenology proved to be differentiable between adult moose ranges, sites of cow parturition, and sites of calf predation. Landscape characteristics of each moose group were consistent and tractable based on environment, suggesting that sites of parturition and predation of moose are predictable in space and time. It is possible that moose selected specific landscapes for parturition despite risk of increased predation of their calves, which could be an example of an "ecological trap." This analytical framework can be employed to identify areas for future ungulate research on the impacts of landscape on parturition and predation dynamics.     

Who let the dogs out? Exploring the spatial ecology of free‐roaming domestic dogs in western Kenya

The spatial ecology of free‐roaming dogs determines their role in the transmission of zoonoses. This study describes the geographic range of and identifies sites frequently visited by free‐roaming domestic dogs in western Kenya.Eight sites in Busia county, western Kenya, were selected. At each site, ten dog‐keeping households were recruited, a questionnaire was administered, and a GPS logger was fixed around the neck of one dog in each household. Loggers were programmed to capture the dog''s position every minute, for five consecutive days. Individual summaries of GPS recordings were produced, and the daily distance traveled was calculated. 50% and 95% utilization distribution isopleths were produced, and the area within these isopleths was extracted to estimate the size of the core and extended Home Ranges (HRs), respectively. Linear regression analyses were performed to identify factors associated with the movement parameters. The centroid points of the 10, 50, and 90% isopleths were reproduced, and the corresponding sites identified on the ground.Seventy‐three dogs were included in the final analyses. The median daily distance traveled was 13.5km, while the median core and extended HRs were 0.4 and 9.3 ha, respectively. Older dogs had a larger extended HR and traveled more daily, while the effect of sex on dog movement depended on their neutering status. Dogs spent most of their time at their household; other frequently visited sites included other household compounds, fields, and rubbish dumps. One of the centroids corresponded to a field located across the international Kenya–Uganda border, emphasizing the fluidity across the border in this ecosystem. Multiple dogs visited the same location, highlighting the heterogeneous contact networks between dogs, and between dogs and people.The field data presented are of value both in understanding domestic dog ecology and resource utilization, and in contextualizing infectious and parasitic disease transmission models.     

Lateral hop movement assesses ankle dynamics and muscle activity

Ankle function is frequently measured using static or dynamic tasks in normal and injured patients. The purpose of this study was to develop a novel task to quantify ankle dynamics and muscle activity in normal subjects. Twelve subjects with no prior ankle injuries participated. Video motion analysis cameras, force platforms, and an EMG system were used to collect data during a lateral hop movement task that consisted of multiple lateral-medial hops over an obstacle. Mean (SD) inversion ankle position at contact was 4.4° (4.0) in the medial direction and -3.5° (4.4) in the lateral direction; mean (SD) tibialis anterior normalized muscle activity was 0.11 (0.08) in the medial direction and 0.16 (0.13) in the lateral direction. The lateral hop movement was shown to be an effective task for quantifying ankle kinematics, forces, moments, and muscle activities in normal subjects. Future applications will use the lateral hop movement to assess subjects with previous ankle injuries in laboratory and clinical settings.     

The formation of “mega‐flocks” depends on vegetation structure in montane coniferous forests of Taiwan

A mixed‐species bird flock is a social assemblage where two or more bird species are moving together while foraging and might benefit from increased foraging efficiency and antipredator vigilance. A “mega‐flock,” which includes flocking species from different vegetation strata, often exhibits high species diversity. Mechanisms for the formation of mega‐flocks have not yet been explored. In this study, we evaluated the influence of vegetation structure and bird species diversity in driving the occurrence of mega‐flocks. We investigated the composition of mixed‐species flocks, local bird communities, and vegetation structure in five vegetation types of two high‐elevation sites in central Taiwan. Mega‐flocks occurred more frequently in pine woodland than later successional stages of coniferous forests. However, species richness/diversity of local bird communities increased along successional stages. Therefore, vegetation variables exhibit more influence on the occurrence of mega‐flocks than local bird communities. Besides foliage height diversity, understory coverage also showed positive effects on flock size of mixed‐species flocks. Our results indicated that pine woodlands with more evenly distributed vegetation layers could facilitate the interactions of canopy and understory flocks and increase the formation of mega‐flocks and thus the complexity of mixed‐species flocks.