Coexistence between two fruit fly species is supported by the different strength of intra‐ and interspecific competition

1. Unravelling the strength and modes of interspecific interactions between resident and introduced species is necessary in order to understand the basis of their coexistence or the displacement of the former by the latter. In Argentina, the indigenous Tephritidae fly Anastrepha fraterculus overlaps its distribution and host fruit with the introduced species Ceratitis capitata. 2. This study focused on the relative strength of intra‐ and interspecific competition during the larval stage as a potential factor supporting coexistence. Classical competition experiments (addition and substitution) were conducted between larvae of the two species reared in artificial larval diet. The study evaluated whether a temporal separation between oviposition events affects the outcome of the competition. 3. When both species started to consume the resource at the same time, A. fraterculus experienced a negative effect in larval survival, pupal weight and duration of larval stage, while for C. capitata, pupal weight decreased. When A. fraterculus started feeding 1 day earlier than C. capitata, the negative effects became milder, and when the temporal separation increased, these effects were reversed. Substitution experiments showed an increase in pupal weight when larvae had to share the resource with heterospecific larvae, and showed negative effects suffered for both species when they shared the resource with conspecific individuals. 4. These results suggest that intraspecific competition is stronger than interspecific competition, and a differential oviposition preference could generate an asynchrony of these species in nature. Such mechanisms could favour coexistence between A. fraterculus and C. capitata in an environment previously occupied only by the former.     

Niche overlap in allotopic and syntopic populations of sexually interacting ground‐hopper species

Abstract There is accumulating evidence that sexual interactions among species (reproductive interference) could have dramatic effects for species’ coexistence. It has been shown that the fitness of individuals can be substantially reduced as a consequence of reproductive interference. This might subsequently lead to displacement of a species (sexual exclusion). On the other hand, some evolutionary and ecological mechanisms might enable species to coexist, such as the divergence of mate recognition systems (reproductive character displacement), habitat partitioning, clumped dispersion patterns or different colonization capabilities. We have previously shown that the two ground‐hopper species Tetrix subulata and Tetrix ceperoi interact sexually in the laboratory as well as in the field. At sites where both species co‐occur niche overlap was high, suggesting that coexistence is maintained by different niche breadths rather than by habitat partitioning. To test the hypothesis that habitat partitioning does not contribute to species’ coexistence, we examined whether allotopic and syntopic populations of these two species differ in niche overlap (competitive release). Our results show that niche overlap is higher in syntopic than in allotopic populations, suggesting that the site‐specific habitat structure (heterogeneity) has a stronger influence on microhabitat utilization than the presence of heterospecifics. Hence, our data do not support the hypothesis that habitat partitioning plays a substantial role for the coexistence of these sexually interacting species.     

How do similarities in spatial distributions and interspecific associations affect the coexistence of Quercus species in the Baotianman National Nature Reserve,Henan, China

Congeneric species often have similar ecological characteristics and use similar resources. These similarities may make it easier for them to co‐occur in a similar habitat but may also lead to strong competitions that limit their coexistence. Hence, how do similarities in congeneric species affect their coexistence exactly? This study mainly used spatial point pattern analysis in two 1 hm² plots in the Baotianman National Nature Reserve, Henan, China, to compare the similarities in spatial distributions and interspecific associations of Quercus species. Results revealed that Quercus species were all aggregated under the complete spatial randomness null model, and aggregations were weaker under the heterogeneous Poisson process null model in each plot. The interspecific associations of Quercus species to non‐Quercus species were very similar in Plot 1. However, they can be either positive or negative in different plots between the co‐occurring Quercus species. The spatial distributions of congeneric species, interspecific associations with non‐Quercus species, neighborhood richness around species, and species diversity were all different between the two plots. We found that congeneric species did have some similarities, and the closely related congeneric species can positive or negative associate with each other in different plots. The co‐occurring congeneric species may have different survival strategies in different habitats. On the one hand, competition among congenerics may lead to differentiation in resource utilization. On the other hand, their similar interspecific associations can strengthen their competitive ability and promote local exclusion to noncongeneric species to obtain more living space. Our results provide new knowledge for us to better understand the coexistence mechanisms of species.     

Assessing niche partitioning of co‐occurring sibling bat species by DNA metabarcoding

Niche partitioning through foraging is a mechanism likely involved in facilitating the coexistence of ecologically similar and co‐occurring animal species by separating their use of resources. Yet, this mechanism is not well understood in flying insectivorous animals. This is particularly true of bats, where many ecologically similar or cryptic species coexist. The detailed analysis of the foraging niche in sympatric, cryptic sibling species provides an excellent framework to disentangle the role of specific niche factors likely involved in facilitating coexistence. We used DNA metabarcoding to determine the prey species consumed by a population of sympatric sibling Rhinolophus euryale and Rhinolophus mehelyi whose use of habitat in both sympatric and allopatric ranges has been well established through radio tracking. Although some subtle dietary differences exist in prey species composition, the diet of both bats greatly overlapped (O_jk = 0.83) due to the consumption of the same common and widespread moths. Those dietary differences we did detect might be related to divergences in prey availabilities among foraging habitats, which prior radio tracking on the same population showed are differentially used and selected when both species co‐occur. This minor dietary segregation in sympatry may be the result of foraging on the same prey‐types and could contribute to reduce potential competitive interactions (e.g., for prey, acoustic space). Our results highlight the need to evaluate the spatial niche dimension in mediating the co‐occurrence of similar insectivorous bat species, a niche factor likely involved in processes of bat species coexistence.     

Co‐occurrence patterns in a diverse arboreal ant community are explained more by competition than habitat requirements

A major goal of community ecology is to identify the patterns of species associations and the processes that shape them. Arboreal ants are extremely diverse and abundant, making them an interesting and valuable group for tackling this issue. Numerous studies have used observational data of species co‐occurrence patterns to infer underlying assembly processes, but the complexity of these communities has resulted in few solid conclusions. This study takes advantage of an observational dataset that is unusually well‐structured with respect to habitat attributes (tree species, tree sizes, and vegetation structure), to disentangle different factors influencing community organization. In particular, this study assesses the potential role of interspecific competition and habitat selection on the distribution patterns of an arboreal ant community by incorporating habitat attributes into the co‐occurrence analyses. These findings are then contrasted against species traits, to explore functional explanations for the identified community patterns. We ran a suite of null models, first accounting only for the species incidence in the community and later incorporating habitat attributes in the null models. We performed analyses with all the species in the community and then with only the most common species using both a matrix‐level approach and a pairwise‐level approach. The co‐occurrence patterns did not differ from randomness in the matrix‐level approach accounting for all ant species in the community. However, a segregated pattern was detected for the most common ant species. Moreover, with the pairwise approach, we found a significant number of negative and positive pairs of species associations. Most of the segregated associations appear to be explained by competitive interactions between species, not habitat affiliations. This was supported by comparisons of species traits for significantly associated pairs. These results suggest that competition is the most important influence on the distribution patterns of arboreal ants within the focal community. Habitat attributes, in contrast, showed no significant influence on the matrix‐wide results and affected only a few associations. In addition, the segregated pairs shared more biological characteristic in common than the aggregated and random ones.     

Composition of mixed‐species flocks and shifts in foraging location of flocking species on Hainan Island,China

A total of 134 bird species were recorded at Jianfengling, Hainan Island, in China from May 2000 to September 2004, of which 44 participated in one or more of 134 mixed‐species flocks. These flocks averaged 3.8 ± 0.2 species and 20.3 ± 1.2 individuals. Flocking propensity in a given species ranged from 1.5 to 100%. For flocking species, frequency of flocking and number of individuals in flocks was positively correlated with frequency and number in point counts. Among all species pairs with flocking frequency above 5%, cluster and correlation analysis indicated there were two principal groups of flocking birds – canopy species and understorey species: associations were positive within a group, but negative between groups. Canopy birds had a higher flocking propensity than understorey birds. They also made significantly less use of inner branches and trunks and greater use of middle branches, and foraged at a significantly greater height when in mixed‐species flocks than when solitary. For understorey bird species, there were no significant differences in foraging locations between solitary and mixed‐species flocks. Higher flocking frequency occurred in the wet season for canopy birds, but in the dry season for understorey birds. Overall patterns were consistent with the explanation that flocking enables an expansion of foraging niche by reducing the risk of predation.     

Density‐dependent survival varies with species life‐history strategy in a tropical forest

Species coexistence in diverse communities likely results from multiple interacting factors. Mechanisms such as conspecific negative density dependence (CNDD) and varying life‐history strategies related to resource partitioning are known to influence plant fitness, and thereby community composition and diversity. However, we have little understanding of how these mechanisms interact and how they vary across life stages. Here, we document the interaction between CNDD and life‐history strategy, based on growth‐mortality trade‐offs, from seedling to adult tree for 47 species in a tropical forest. Species’ life‐history strategies remained consistent across stages: fast‐growing species had higher mortality than slow‐growing species at all stages. In contrast, mean CNDD was strongest at early life stages (i.e. seedling, sapling). Fast‐growing species tended to suffer greater CNDD than slow‐growing species at several, but not all life stages. Overall, our results demonstrate that coexistence mechanisms interact across multiple life stages to shape diverse tree communities.     

Parasite–grass–forb interactions and rock–paper– scissor dynamics: predicting the effects of the parasitic plant Rhinanthus minor on host plant communities

1.  Parasitic plants affect the growth, reproduction and metabolism of their hosts and may also influence the outcome of competitive interactions between host species and, consequently, the structure of entire host communities.
2.  We investigate the effect of the root hemiparasitic plant Rhinanthus minor on plant community dynamics using a spatial theoretical model. The model is parameterized with data from pairwise interaction experiments under two nutrient levels between the hemiparasite and three grass species ( Cynosurus cristatus , Festuca rubra and Phleum bertolonii ) and three forb species ( Leucanthemum vulgare , Plantago lanceolata and Ranunculus acris ).
3.  Relative interaction coefficients were intransitive, with the dynamics of the system conforming to a rock–paper–scissors game. Stable deterministic dynamics emerge from parameters obtained under low-nutrient conditions. Under high-nutrient conditions, the dynamics are unstable, but are stabilized in spatially explicit models. The outcomes are sensitive to initial spatial pattern and frequency.
4.   Synthesis . This study supports the idea that hemiparasite populations may form 'shifting clouds' in natural populations and explains seemingly unpredictable shifts in host community structure following introduction of hemiparasites. Management of plant communities using hemiparasites needs to take these complex dynamics into account.     

Testing multiple pathways for impacts of the non‐native Black‐headed Weaver Ploceus melanocephalus on native birds in Iberia in the early phase of invasion

Not all non‐native species have strong negative impacts on native species. It is desirable to assess whether a non‐native species will have a negative impact at an early stage in the invasion process, when management options such as eradication are still available. Although it may be difficult to detect early impacts of non‐native species, it is necessary to ensure that management decisions can be based on case‐specific scientific evidence. We assess the impacts of a non‐native bird, the Black‐headed Weaver Ploceus melanocephalus, at an early stage in its invasion of the Iberian Peninsula. To do this we identify potential pathways by which competition for shared resources by Black‐headed Weavers could lead to population declines in two ecologically similar native species, and generate hypotheses to test for evidence of competition along these pathways. Black‐headed Weavers could potentially impact native species by displacing them from nesting habitat, or by reducing habitat quality. We found no evidence for either potential competition pathway, suggesting that Black‐headed Weavers do not currently compete with the two native species. However, it is possible that mechanisms that currently allow coexistence may not operate once Black‐headed Weavers reach higher population densities or different habitats.     

Effect of soil nutrients,neighbor identities and root separation types on intra‐ and interspecific interaction among three clonal plant species

In natural environments, plants frequently interact with both heterospecific and conspecific neighbors. The intensity of belowground plant interaction with neighboring species commonly varies with the availability of soil nutrients in the habitats. According to classical ecological theory, competition between conspecific neighbors may be more severe than competition between unrelated species due to the similar nutrient requirements of close relatives, especially when nutrients are scarce in the habitat. However, many recent studies have shown the opposite pattern, and suggested an alternative mechanism based on species recognition. Taking Zoysia sinica as the focal species, we conducted a controlled experiment to test the results of intraspecific and interspecific interactions among three clonal species Zoysia sinica, Zoysia japonica and Alternanthera philoxeroides, which represent a conspecific, a close relative and a distant relative of the focal species, respectively, and at different root treatments (no separation NS, clone separation CS and ramet separation RS) and two nutrient levels. The results showed that Z. sinica recognized conspecific plants in the NS and CS treatments, and did not show above or belowground competition with these. The performance of the focal plant (Z. sinica) was better when it was grown with a conspecific neighbor as compared to all other types of neighbors. In all root separation treatments, the competition was more intense when Z. sinica grew with a close relative (Z. japonica) than when growing with a distant relative (A. philoxeroides). Generally, competition between plants was more intense at the high nutrient level than at the low nutrient level, suggesting that both soil nutrients and a species recognition mechanism play a significant role for the intra‐ and interspecific interaction and fitness of these three neighboring clonal species.     

Rainfall variability and fine‐scale life history tradeoffs help drive niche partitioning in a desert annual plant community

Tradeoffs have long been an essential part of the canon explaining the maintenance of species diversity. Despite the intuitive appeal of the idea that no species can be a master of all trades, there has been a scarcity of linked demographic and physiological evidence to support the role of resource use tradeoffs in natural systems. Using five species of Chihuahuan desert summer annual plants, I show that demographic tradeoffs driven by short‐term soil moisture variation act as a mechanism to allow multiple species to partition a limiting resource. Specifically, by achieving highest fitness in either rainfall pulse or interpulse periods, variability reduces fitness differences through time that could promote coexistence on a limiting resource. Differences in fitness are explained in part by the response of photosynthesis to changing soil moisture. My results suggest that increasing weather variability, as predicted under climate change, could increase the opportunity for coexistence in this community.     

Colonization and coexistence of non-native ants on a model Atlantic island

Aims

Colonization by non-native ants represents one of the gravest potential threats to island ecosystems. It is necessary to identify general mechanisms by which non-native species are able to colonize and persist in order to inform future prevention and management. We studied a model-island assemblage of 17 non-native ant species with aim of identifying the spatial source of introductions and assessing how such a diversity of species are able to coexist.

Location

Data were collected on Ascension Island: an ideal study system for its intermediate area, compact shape, spatial heterogeneity, lack of native ant species, and availability of non-native ant records dating back to the 1800s.

Methods

We collected over 47,000 individual ants from 73 sites using a range of baited traps and survey techniques. We combined this novel data with past occurrence records in order to determine whether human settlements have historically been the source of ant introductions and to quantify the mean rate at which species have dispersed across the island. Analysis of standardized field data revealed the extent to which ants were partitioning ecological niche space via (1) habitat separation, (2) fine-scale resource partitioning and (3) climatic heterogeneity.

Results

Ants were radiating at a linear rate of approximately 0.5 km² per year from human settlements on this island, with the most widespread species having been introduced earliest. After accounting for incomplete colonization, we found no evidence to suggest habitat separation between species. Instead, we found significant niche separation through resource partitioning and weather-dependent activity patterns.

Main Conclusions

Our results indicate that non-native ants can coexist in very close proximity and are therefore capable of existing at great diversity on even small islands. It is inevitable that ant colonization will continue without increased biosecurity measures, habitat restoration around settlements and conservation of native species populations.     

Niche shift in four non‐native estrildid finches and implications for species distribution models

Non‐native species can have severe impacts on ecosystems. Therefore, predictions of potentially suitable areas that are at risk of the establishment of non‐native populations are desirable. In recent years, species distribution models (SDMs) have been widely applied for this purpose. However, the appropriate selection of species records, whether from the native area alone or also from the introduced range, is still a matter of debate. We combined analyses of native and non‐native realized climate niches to understand differences between models based on all locations, as well as on locations from the native range only. Our approach was applied to four estrildid finch species that have been introduced to many regions around the world. Our results showed that SDMs based on location data from native areas alone may underestimate the potential distribution of a given species. The climatic niches of species in their native ranges differed from those of their non‐native ranges. Niche comparisons resulted in low overlap values, indicating considerable niche shifts, at least in the realized niches of these species. All four species have high potential to spread over many tropical and subtropical areas. However, transferring these results to temperate areas has a high degree of uncertainty, and we urge caution when assessing the potential spread of tropical species that have been introduced to higher latitudes.     

Limited niche differentiation within remarkable co‐occurrences of congeneric species: Monomorium ants in the Australian seasonal tropics

Niche theory predicts that few closely related species can co‐occur because such species tend to be ecologically similar and niche differentiation is required to avoid competitive exclusion. We analyse the co‐occurrence of a remarkable 10–15 species of the ant genus Monomorium occurring within single 10 × 10 m plots in a tropical savanna of northern Australia. Most of the species are undescribed, so we use genetic analysis to validate our species demarcations. We document nest dispersion patterns, and investigate differentiation in the three primary niche dimensions: space, time and food. We also examine species differences in competitive abilities, by describing rates of foraging activity, foraging ranges, worker aggression, and levels of behavioural dominance. Analyses of nest and forager distributions showed very limited evidence of spatial segregation within plots. The great majority of species foraged either exclusively or primarily during daylight hours. Body size and isotopic analyses indicated very limited dietary differentiation. Such limited niche partitioning occurred despite the species differing markedly in their competitive abilities as measured by rates of resource discovery, recruitment and monopolization. Our findings defy the traditional assumption that multiple closely related and ecologically similar species of highly interactive taxa cannot co‐occur. It seems very likely that species coexistence in our study system is determined to a very large degree by stochastic processes relating to dispersal and establishment, as predicted by neutral theory. However, neutral theory assumes competitive equivalence, whereas we found very marked differences in the competitive abilities of our co‐occurring species. We suggest that competitive exclusion is prevented by the modular nature of ant colonies, with competition limiting colony performance but not preventing occurrence. We conclude that other factors that allow species persistence, and not just competitive equivalence, can allow dispersal and establishment processes to drive species coexistence.     

Thermal adaptation affects interactions between a range‐expanding and a native odonate species

1. Increasing temperature and invading species may interact in their effects on communities. In this study, we investigated how rising temperatures alter larval interactions between a naturally range‐expanding dragonfly, Crocothemis erythraea, and a native northern European species, Leucorrhinia dubia. Initial studies revealed that C. erythraea grow up to 3.5 times faster than L. dubia at temperatures above 16 °C. As a result, we hypothesised that divergent temperature responses would lead to rapid size differences between coexisting larvae and, consequently, to asymmetric intraguild predation at higher ambient temperatures. 2. Mortality and growth rates were measured in interaction treatments (with both species present) and non‐interaction controls (one species present) at four different temperature regimes: at an ambient temperature representative of central Germany, where both species overlap in distribution, and at temperatures increased by 2, 4 and 6 °C. 3. The mortality of C. erythraea did not differ between treatment and control. In contrast, mortality of L. dubia remained similar over all temperatures in the controls, but increased with temperature in the presence of the other species and was significantly higher there than in the controls. We concluded that L. dubia suffered asymmetric intraguild predation, particularly at increased temperature. Reduced growth rate of L. dubia in the interaction treatment at higher temperatures also suggested asymmetric competition for prey in the first phase of the experiment. 4. The results imply that the range expansion of C. erythraea may cause reduction in population size of syntopic L. dubia when temperature rises by more than 2 °C. The consequences for future range patterns, as well as other factors that may influence the interaction in nature, are discussed.     

A review of the occurrence of inter‐colony segregation of seabird foraging areas and the implications for marine environmental impact assessment

Understanding the determinants of species’ distributions is a fundamental aim in ecology and a prerequisite for conservation but is particularly challenging in the marine environment. Advances in bio‐logging technology have resulted in a rapid increase in studies of seabird movement and distribution in recent years. Multi‐colony studies examining the effects of intra‐ and inter‐colony competition on distribution have found that several species exhibit inter‐colony segregation of foraging areas, rather than overlapping distributions. These findings are timely given the increasing rate of human exploitation of marine resources and the need to make robust assessments of likely impacts of proposed marine developments on biodiversity. Here we review the occurrence of foraging area segregation reported by published tracking studies in relation to the density‐dependent hinterland (DDH) model, which predicts that segregation occurs in response to inter‐colony competition, itself a function of colony size, distance from the colony and prey distribution. We found that inter‐colony foraging area segregation occurred in 79% of 39 studies. The frequency of occurrence was similar across the four seabird orders for which data were available, and included species with both smaller (10–100 km) and larger (100–1000 km) foraging ranges. Many predictions of the DDH model were confirmed, with examples of segregation in response to high levels of inter‐colony competition related to colony size and proximity, and enclosed landform restricting the extent of available habitat. Moreover, as predicted by the DDH model, inter‐colony overlap tended to occur where birds aggregated in highly productive areas, often remote from all colonies. The apparent prevalence of inter‐colony foraging segregation has important implications for assessment of impacts of marine development on protected seabird colonies. If a development area is accessible from multiple colonies, it may impact those colonies much more asymmetrically than previously supposed. Current impact assessment approaches that do not consider spatial inter‐colony segregation will therefore be subject to error. We recommend the collection of tracking data from multiple colonies and modelling of inter‐colony interactions to predict colony‐specific distributions.     

Rethinking the nature of intraspecific variability and its consequences on species coexistence

Intraspecific variability (IV) has been proposed to explain species coexistence in diverse communities. Assuming, sometimes implicitly, that conspecific individuals can perform differently in the same environment and that IV increases niche overlap, previous studies have found contrasting results regarding the effect of IV on species coexistence. We aim at showing that the large IV observed in data does not mean that conspecific individuals are necessarily different in their response to the environment and that the role of high‐dimensional environmental variation in determining IV has largely remained unexplored in forest plant communities. We first used a simulation experiment where an individual attribute is derived from a high‐dimensional model, representing “perfect knowledge” of individual response to the environment, to illustrate how large observed IV can result from “imperfect knowledge” of the environment. Second, using growth data from clonal Eucalyptus plantations in Brazil, we estimated a major contribution of the environment in determining individual growth. Third, using tree growth data from long‐term tropical forest inventories in French Guiana, Panama and India, we showed that tree growth in tropical forests is structured spatially and that despite a large observed IV at the population level, conspecific individuals perform more similarly locally than compared with heterospecific individuals. As the number of environmental dimensions that are well quantified at fine scale is generally lower than the actual number of dimensions influencing individual attributes, a great part of observed IV might be represented as random variation across individuals when in fact it is environmentally driven. This mis‐representation has important consequences for inference about community dynamics. We emphasize that observed IV does not necessarily impact species coexistence per se but can reveal species response to high‐dimensional environment, which is consistent with niche theory and the observation of the many differences between species in nature.     

SESAM – a new framework integrating macroecological and species distribution models for predicting spatio‐temporal patterns of species assemblages

Two different approaches currently prevail for predicting spatial patterns of species assemblages. The first approach (macroecological modelling, MEM) focuses directly on realized properties of species assemblages, whereas the second approach (stacked species distribution modelling, S‐SDM) starts with constituent species to approximate the properties of assemblages. Here, we propose to unify the two approaches in a single ‘spatially explicit species assemblage modelling’ (SESAM) framework. This framework uses relevant designations of initial species source pools for modelling, macroecological variables, and ecological assembly rules to constrain predictions of the richness and composition of species assemblages obtained by stacking predictions of individual species distributions. We believe that such a framework could prove useful in many theoretical and applied disciplines of ecology and evolution, both for improving our basic understanding of species assembly across spatio‐temporal scales and for anticipating expected consequences of local, regional or global environmental changes. In this paper, we propose such a framework and call for further developments and testing across a broad range of community types in a variety of environments.     

The influence of light conditions and interspecific competition on the root foraging traits and seedling growth of two tree species

Abstract

Enriched nutrient patches within natural soil represent an important source of nutrients for tree growth. In the present study, pot experiments in a heterogeneous nutrient environment were conducted to investigate the influence of light conditions and interspecific competition on the root foraging traits and seedling growth of Pinus massoniana and Schima superba. The root foraging scale and the whole-seedling biomass of both species were decreased by shading. The result of this treatment was a lower sensitivity to nutrient heterogeneity in plants that underwent the shading treatment than in plants that were exposed to full-light conditions. The above-ground biomass and whole-seedling biomass of S. superba were not affected by competition with P. massoniana. In contrast, the above-ground biomass and whole-seedling biomass of P. massoniana were negatively affected by competition with S. superba. The more rapid rate of root extension and the more efficient resource uptake of S.superba appear to explain this effect. The species-specific patterns of the influence of environmental factors on foraging ability and seedling growth should be given thorough consideration and should be applied to afforestation and to the management of tree plantations.     

Effects of local biotic neighbors and habitat heterogeneity on seedling survival in a spruce‐fir valley forest,northeastern China

Seedlings are vulnerable to many biotic and abiotic agents, and studying seedling dynamics helps understand mechanisms of species coexistence. In this study, the relative importance of biotic neighbors and habitat heterogeneity to seedling survival was examined by generalized linear mixed models for 33 species in a spruce‐fir valley forest in northeastern China. The results showed that the relative importance of these factors varied with species and functional groups. Conspecific negative density dependence (CNDD) was important to the survival of Abies nephrolepis and Picea koraiensis seedling, whereas phylogenetic negative density dependence (PNDD) was critical to Pinus koraiensis and Betula platyphylla, as well as functional groups of tree, deciduous, and shade‐intolerant seedlings. For shrubs and Acer ukurunduense, habitat heterogeneity was significant. Despite of the significance of CNDD, PNDD, and habitat heterogeneity on seedling survival, large proportions of the total variance were not accounted for by the studied variables, suggesting the needs to examine the influences of other factors such as pests, diseases, herbivores, forest structure, species functional traits, and microclimatic conditions on seedling survival in the future.