The evolutionary species pool hypothesis and patterns of freshwater diatom diversity along a pH gradient

Aim To interpret the unimodal relationship between diatom species richness and lake pH within the context of the evolutionary species pool hypothesis (SPH). We test the following primary prediction arising from the SPH: the size of the potential species pool (PSP) will increase along a gradient representing the historical commonness of different pH environments (pH commonness). To do this we assume that the present‐day spatial dominance of near‐neutral pH conditions compared with acidic and alkaline conditions reliably mimics the relative spatial availabilities of historical pH conditions among freshwater lakes. We also determine whether local richness represents a constant proportion of PSP size along the pH commonness gradient. Location Two hundred and thirty‐four lakes distributed over a 405,000 km² region of the north‐eastern United States of America. Methods Sediment diatom morphospecies lists and pH data were acquired from the US Environmental Protection Agency's Environmental Monitoring and Assessment Program (EMAP) website. Using 248 morphospecies that occurred in at least 10 of the 234 lakes, four different measures of PSPs were calculated along the pH gradient. Local species richness was equated with the number of species occurring within the lake. Alpha diversity was equated with the average species richness of lakes with similar pH values. A combination of statistical methods were employed, including correlations, quadratic regression and piecewise regression. Results PSP size increased significantly with pH commonness for all four measures of PSP size, thus supporting the primary prediction of the evolutionary SPH. Local richness comprised a larger proportion of the PSP within acidic lakes than within circumneutral lakes. Alpha diversity and lake species richness both increased significantly with pH commonness, but the former did so in a two‐step fashion. We test and reject several alternative contemporary time‐scale explanations for our findings. Main Conclusions Our findings are consistent with the hypothesis that diatom taxonomic richness is presently lower within acidic and highly alkaline lakes than in circumneutral lakes owing to the limited opportunity in space and/or time for the evolution of suitably adapted species. Whereas ecological processes can explain why certain species are excluded from particular habitats, e.g. acidic lakes, they cannot account for why so few species are adapted to those habitats in the first place.     

Community assembly along a species pool gradient: implications for multiple-scale patterns of species diversity

Various ecological processes influence patterns of species diversity at multiple spatial scales. One process that is potentially important but rarely considered is community assembly. I assembled model communities using species pools of differing size to examine how the history of community assembly may affect multi-scale diversity patterns. The model contained three scales at which diversity could be measured: local community, metacommunity, and species pool. Local species saturation occurred, as expected from the competition and predation built in the model. However, local communities did not become resistant to invasions except when the species pool was very small. Depending on dispersal rate and trophic level, the larger the species pool, the harder it was to predict which species invades which local community at a given time. Consequently, local-community dissimilarity maintained by assembly history increased linearly with pool size, even though local diversity was decoupled from pool size. These results have two implications for multi-scale diversity patterns. First, assembly history may provide an explanation for scale-dependent relationships between local and regional diversity: assembly causes the relationship to be curvilinear at one scale (local community), while linear at another (metacommunity). Second, assembly history influences how -diversity is partitioned into - and -diversity: assembly causes the relative contribution of to increase with pool size. Overall, this study suggests that community assembly history interacts with species pool size to regulate multi-scale patterns of species diversity.     

The distribution of vascular plant species and guilds in space and time along a desert gradient

Abstract. We studied distribution patterns of vascular plant species and environmental variables for three years along a permanent transect traversing a closed-drainage watershed in the northern Chihuahuan Desert of south-central New Mexico, USA. The transect extended for 2.7 km from a basin floor playa (1310 m elevation, fine-textured soil), across a piedmont slope, and onto the base of a granitic mountain (1410 m elevation, coarse-textured soil). The gradients in elevation and soils across our transect, along with variable seasonal rainfall, downslope redistribution of water and organic matter, and soil texture-related variation in infiltration, water holding capacity, and moisture release characteristics, interact to generate a complex spatial and temporal gradient of available soil water and nitrogen. We grouped plant species into guilds according to growth form and photo synthetic pathway type. These guilds are spatially and temporally differentiated along the transect such that particular groups utilize particular seasonal phases or spatial regions of the gradient. We identified six distinct plant communities along the transect. C₄ perennial grasses dominated the mesic/high nitrogen portion of the gradient, which occurred at the highest (upper piedmont grassland, dominated by Bouteloua eriopoda) and lowest (playa, dominated by Panicum obtusum) elevations along the transect. C₃ shrubs were dominant in the xeric/low nitrogen portion of the gradient located near the middle of the transect (bajada shrubland, dominated by Larrea tridentata). C₃ shrubs also dominated a narrow zone of vegetation located adjacent to the playa (playa fringe, dominated by Prosopisglandulosa). C₄ perennial grasses, C₃ subshrubs, and C₃ and C₄ perennial forbs and annuals were co-dominant in the intermediate locations along the gradient, which occurred below (mixed basin slopes) and above (lower piedmont grassland) the bajada shrubland. Life-form distribution patterns at the small scale of our study reflect some of the patterning that occurs at larger scales in response to climate gradients. The distributions of some species and guilds along the transect are apparently modified by competitive interactions.     

Scale-dependent patterns of macrofaunal distribution in soft-sediment intertidal habitats along a large-scale estuarine gradient

We investigated the pattern of distribution of intertidal soft-bottom fauna in streams and lagoons of the Uruguayan coast at three spatial scales. The Río de la Plata and the Atlantic Ocean produce on this coast a large-scale gradient in salinity, defining a freshwater (west), an estuarine (central) and a marine (east) region. Within each region, there are several streams and coastal lagoons (sites) that define a second scale of variability. A third scale is given by intertidal gradients within each site. Species richness and total abundance was low in the freshwater west region and high in the central and east regions. The community in the west region was characterized by the clam Curbicula fluminea; in the other regions, it was dominated mainly by the polychaete Heteromastus similis. The polychaete Nephtys fluviatilis was more abundant in the east region, while another polychaete, Laeonereis acuta, characterized the central region. Sediment fractions did not vary significantly at this scale. At the scale of the sites, species richness and total macrofaunal abundance were higher in coastal lagoons than in streams. Coarse sands were more common in coastal lagoons, while medium and fine sand characterized the sediment in streams. Within each site, species richness and total abundance increased towards the lower intertidal level; the macrofauna of the upper levels were a subsample of the fauna occurring at the lower levels. There was also a significantly lower proportion of fine sand at the upper level. At regional scales, the observed patterns may be indirectly or directly related to the gradient in salinity, through differential physiological tolerance to osmotic stress. At the scale of the sites, variability may be explained mainly by geomorphological and sedimentological differences between lagoons and streams. Variation among levels may be related to gradients in desiccation, colonization and predation.     

Ecological correlates of the distribution limits of two poeciliid species along a salinity gradient

Identifying the environmental factors responsible for the formation of a species' distribution limit is challenging because organisms interact in complex ways with their environments. However, the use of statistical niche models in combination with the analysis of phenotypic variation along environmental gradients can help to reduce such complexity and identify a subset of candidate factors. In the present study, we used such approaches to describe and identify factors responsible for the parapatric distribution of two closely‐related livebearer fish species along a salinity gradient in the lowlands of Trinidad, West Indies. The downstream distribution limits of Poecilia reticulata were strongly correlated with the brackish–freshwater interface. We did not observe significant phenotypic variation in life‐history traits for this species when comparing marginal with more central populations, suggesting that abrupt changes in conditions at the brackish–freshwater interface limit its distribution. By contrast, Poecilia picta was present across a wide range of salinities, although it gradually disappeared from upstream freshwater localities. In addition, P. picta populations exhibited an increase in offspring size in localities where they coexist with P. reticulata, suggesting a role for interspecific competition. The parapatric distribution of these two species, suggests that P. reticulata distributions are limited by an abiotic factor (salinity), whereas P. picta is limited by a biotic factor (interspecific competition). Similar parapatric patterns have been previously described in other systems, suggesting they might be a common pattern in nature. © 2013 The Linnean Society of London     

神农架自然保护区非飞行哺乳动物的物种丰富度:沿海拔梯度的分布格局

于1999～2001年调查了神农架自然保护区6个地点不同栖息地的非飞行哺乳动物的物种丰富度。栖息地分为8类：原始林、择伐林（采伐枯立木）、次生林、灌木林、草地、常年性河流水溪、农田和人居住区。小型非飞行哺乳动物调查用捕鼠夹;大型非飞行哺乳动物调查主要根据皮张收购资料以及样线法和痕迹法;用10 m×10 m的样方调查林地树种丰富度。调查发现,神农架自然保护区有非飞行哺乳动物59种［不包括引进种梅花鹿（Cervus nippon）］。在同一海拔高度,原始林通常比择伐林和次生林的物种丰富度高,说明采伐严重降低了物种丰富度。对比同一栖息地不同海拔高度的物种丰富度,我们发现,在中海拔地段（800～1700 m）物种丰富度最高：如在原始林和次生林,海拔1700 m的东溪物种丰富度最高;在择伐林,海拔800 m的九冲物种丰富度最高。聚类分析显示,6个地点的哺乳动物物种组成可以分为两组：高海拔组（2100 m以上）和中低海拔组（1700 m以下）。各地点的哺乳动物物种组成与植被的垂直分布是一致的。各地点的物种丰富度与单位面积（100 m²）树种平均丰富度、栖息地类型数和海拔高度相关。3个环境变量间也是相关的：海拔高度对单位面积树种平均丰富度和栖息地类型数有重要影响。根据研究结果提出两点保护建议：第一,保护区的移民迁出和退耕还林工程应首先在物种丰富度最高的九冲进行,而后是东溪和下谷;第二,为了增加个体流和基因流,保护区东西两片相间的非保护区地带应划入保护区,建立栖息地廊道。     

Multitaxonomic diversity patterns along a desert riparian-upland gradient

Riparian areas are noted for their high biodiversity, but this has rarely been tested across a wide range of taxonomic groups. We set out to describe species richness, species abundance, and community similarity patterns for 11 taxonomic groups (forbs & grasses, shrubs, trees, solpugids, spiders, scarab beetles, butterflies, lizards, birds, rodents, and mammalian carnivores) individually and for all groups combined along a riparian-upland gradient in semiarid southeastern Arizona, USA. Additionally, we assessed whether biological characteristics could explain variation in diversity along the gradient using five traits (trophic level, body size, life span, thermoregulatory mechanism, and taxonomic affiliation). At the level of individual groups diversity patterns varied along the gradient, with some having greater richness and/or abundance in riparian zones whereas others were more diverse and/or abundant in upland zones. Across all taxa combined, riparian zones contained significantly more species than the uplands. Community similarity between riparian and upland zones was low, and beta diversity was significantly greater than expected for most taxonomic groups, though biological traits explained little variance in diversity along the gradient. These results indicate heterogeneity amongst taxa in how they respond to the factors that structure ecological communities in riparian landscapes. Nevertheless, across taxonomic groups the overall pattern is one of greater species richness and abundance in riparian zones, coupled with a distinct suite of species.     

Occurrence patterns of facilitation by shade along a water gradient are mediated by species traits

In disturbed habitats, shade often has facilitative effects on plants by ameliorating water and thermal stresses. Facilitation by shade tends to increase as water availability decreases. At the same time, several studies have suggested that facilitation by shade is not affected by water status or collapses under extremely dry conditions. We hypothesized that traits of beneficiary plants, specifically, the flexibility in the allocation of biomass between shoots and roots, would mediate variation in the relationship between facilitation by shade and water status. To test this hypothesis, we examined the responses of two bog species to shade under various water conditions in a post-mined peatland. The seeds of Rhynchospora alba and Moliniopsis japonica were sown under three water levels (dry: 53% peat water content, wet: 77%, and control: 71%) × two shading levels (50% shaded and unshaded). The survival, biomass, and biomass allocation between the shoots and roots of the two species were monitored for two years. Shade increased the survival and biomass of both species. However, the facilitation of R. alba by shade was independent of water level, whereas the strength of the facilitative effects on M. japonica increased as water content decreased. R. alba preferentially allocated biomass to roots under dry conditions and was highly drought tolerant. M. japonica did not alter the allocation of its biomass in response to either shade or water level and was drought intolerant. Our results suggest that flexibility in biomass allocation of beneficiary plants mediates occurrence patterns of facilitation by shade along a water gradient. The facilitation of species with inflexible biomass allocation by shade through the amelioration of water stress increases as water availability decreases, whereas the facilitation of species with flexible biomass allocation is independent of water status. Such species-specific facilitation would promote the coexistence of diverse species in a community.     

Bacterial diversity patterns along a gradient of primary productivity

Primary productivity is a key determinant of biodiversity patterns in plants and animals but has not previously been shown to affect bacterial diversity. We examined the relationship between productivity and bacterial richness in aquatic mesocosms designed to mimic small ponds. We observed that productivity could influence the composition and richness of bacterial communities. We showed that, even within the same system, different bacterial taxonomic groups could exhibit different responses to changes in productivity. The richness of members of the Cytophaga‐Flavobacteria‐Bacteroides group exhibited a significant hump‐shaped relationship with productivity, as is often observed for plant and animal richness in aquatic systems. In contrast, we observed a significant U‐shaped relationship between richness and productivity for α‐proteobacteria and no discernable relationship for β‐proteobacteria. We show, for the first time, that bacterial diversity varies along a gradient of primary productivity and thus make an important step towards understanding processes responsible for the maintenance of bacterial biodiversity.     

Non-native species distribution along the elevation gradient in the western Italian Alps

Abstract

In this work the occurrence of non-native species was recorded along the elevation gradient in the Alps, in order to establish their distribution pattern, their current altitudinal limits and to elucidate which species are presumably adapted to higher elevations. Plots were located along the course of rivers in five valleys from 100 to 2100 m a.s.l. Sixty-eight non-natives were recorded in the study area. The proportion of invasives was found to be much higher in the study area then in the whole administrative region. The number of non-natives per plot decreased strongly with increasing elevation. The great majority (94%) of the non-native species grows at the lowest elevations, while only 6% survive up to 1500 m and none was found above this limit. Results were interpreted considering the factors driving the invasion process (disturbance, the native communities' resistance to invasion, propagule pressure, climatic conditions) and plant traits with particular respect to pre-adaptation to the harsh climate, which increases progressively with elevation. Results confirm that the Alps are not immune from invasion, at least up to medium elevation.     

Seed predation of non-native species along a precipitation gradient

Generalist seed predators are an important factor shaping non-native species invasion. Their effect is highly influenced by abiotic conditions, yet how the importance of this biotic filter changes in a gradient of abiotic conditions is still poorly understood. In this study, we assessed seed predation of non-native conifer species along a precipitation gradient in north-western Argentinean Patagonia. We performed a seed removal experiment over a 15-day period during the fall in sites where annual precipitation ranges from 600 to 1600 mm and vegetation ranges from steppe to forest. We used the three most common forestry species in the study area, including two species known to invade areas adjacent to plantations and one native species. Seed predation was higher in sites with higher precipitation, but no significant differences in seed predation were found among species. Small mammals were the dominant group predating seeds on this gradient. These findings highlight the importance of abiotic conditions mediating plant-granivore interactions, and the influence that abiotic conditions may have on biotic resistance to invasion.     

Numerically dominant species drive patterns in resource use along a vertical gradient in tropical ant assemblages

Resource availability can influence the foraging strategy adopted by different ant species as they endeavor to meet nutrient demands of the colony. In tropical rain forests, environmental conditions including resource availability vary over a vertical gradient. Consequently, nitrogen is predicted to become more limiting than carbohydrates toward the canopy as food webs shift to become more reliant on plant-based resources. We used a “bait-choice” experiment in a tropical rain forest to examine differences in protein and carbohydrate use with height and determined whether there were differences in response between common (numerically dominant) and rare species. Additionally, we investigated the nutrient use at the species level. Using species co-occurrence analysis, we examined interspecific competition by testing the co-occurrence of ant species at the tree level. Over the 12 trees investigated, 124 morphospecies were identified with eight species comprising 90% of total ant abundance. Species richness and protein use increased with height of bait for all species pooled and for common species but not rare species. Correspondingly, relative carbohydrate use decreased with height. We found greater species richness of rare species on carbohydrate baits compared with protein baits. Ant species were randomly distributed among trees when all species were included in co-occurrence analysis. However, when only common species were considered, segregation between species was evident among trees providing evidence for the presence of ant mosaics. Our results suggest that nitrogen limitation in the canopy may not be true for the whole ant assemblage but rather for the few common species.     

Spatial patterns of weeds along a gradient of landscape complexity

Processes that drive spatial patterning among plant species are of ongoing interest, mostly because these patterns have implications for the structure and function of plant communities. We investigated the spatial strategies of weeds focusing on how spatial patterns of weeds are mediated by agricultural landscape complexity and species life-history attributes. We quantified the spatial distribution of 110 weed species using data collected in ten landscapes in central western France along a gradient of landscape complexity, from structurally complex (numerous small fields) to structurally simple (few large fields). We then related differences observed in species’ distribution patterns to ecological attributes of species for resource exploitation and dispersion. Our study reveals that weeds were spatially aggregated at the landscape scale. Their spatial patterns are related to the frequency of occurrence of weeds but surprisingly not directly to the seed dispersal type, nor to the degree of habitat specialization. We show that landscape complexity had no direct effect on the spatial patterning of weeds but through interactions with species attributes. Our results point to the importance of interactions between landscape complexity and species attributes in the spatial patterning of weed species even in intensively managed fields. These patterns appear to be a consequence of the spatial arrangement of landscape elements as well as the result of landscape filtering on species attributes.     

Diverging drought-tolerance strategies explain tree species distribution along a fog-dependent moisture gradient in a temperate rain forest

The study of functional traits and physiological mechanisms determining species’ drought tolerance is important for the prediction of their responses to climatic change. Fog-dependent forest patches in semiarid regions are a good study system with which to gain an understanding of species’ responses to increasing aridity and patch fragmentation. Here we measured leaf and hydraulic traits for three dominant species with contrasting distributions within patches in relict, fog-dependent forests in semiarid Chile. In addition, we assessed pressure–volume curve parameters in trees growing at a dry leeward edge and wet patch core. We predicted species would display contrasting suites of traits according to local water availability: from one end favoring water conservation and reducing cavitation risk, and from the opposite end favoring photosynthetic and hydraulic efficiency. Consistent with our hypothesis, we identified a continuum of water use strategies explaining species distribution along a small-scale moisture gradient. Drimys winteri, a tree restricted to the humid core, showed traits allowing efficient water transport and high carbon gain; in contrast, Myrceugenia correifolia, a tree that occurs in the drier patch edges, exhibited traits promoting water conservation and lower gas exchange rates, as well low water potential at turgor loss point. The most widespread species, Aextoxicon punctatum, showed intermediate trait values. Osmotic compensatory mechanism was detected in M. correifolia, but not in A. punctatum. We show that partitioning of the pronounced soil moisture gradients from patch cores to leeward edges among tree species is driven by differential drought tolerance. Such differences indicate that trees have contrasting abilities to cope with future reductions in soil moisture.     

Mollusc community patterns and species response curves along a mineral richness gradient: a case study in fens

Aim The goals of this study were to: (1) compare water conductivity and pH as proxy measures of mineral richness in relation to mollusc assemblages in fens, (2) examine the patterns of mollusc species richness along the gradient of mineral richness based on these factors, (3) model species–response curves and analyse calcicole–calcifuge behaviour of molluscs, and (4) compare the results with those from other studies concerning non‐marine mollusc ecology. Location Altogether, 135 treeless spring fen sites were sampled within the area of the Western Carpathians (east Czech Republic, north‐west Slovakia and south Poland; overall extent of study area was 12,000 km²). Methods Mollusc communities were recorded quantitatively from a homogeneous area of 16 m². Water conductivity and pH were measured in the field. The patterns of local species diversity along selected gradients, and species–response curves, were modelled using generalized linear models (GLM) and generalized additive models (GAM), both using the Poisson distribution. Results When the most acid sites (practically free of molluscs) were excluded, conductivity expressed the sites’ mineral richness and base saturation within the entire gradient, in contrast to pH. In the base‐rich sites, pH did not correlate with mineral richness. A unimodal response of local species diversity to mineral richness (expressed as conductivity) was found. In the extremely mineral‐rich, tufa‐forming sites (conductivity > 600 μS cm^?1) a decrease in species diversity was encountered. Response curves of the most common species showed clear differentiation of their niches. Significant models of either unimodal or monotonic form were fitted for 18 of the 30 species analysed. Species showed five types of calcicole–calcifuge behaviour: (1) a decreasing monotonic response curve and a preference for the really acid sites; (2) a skewed unimodal response curve with the optimum shifted towards the slightly acid sites; (3) a symmetrical unimodal model response curve with the optimum in the base‐rich sites, with no or slight tufa precipitation; (4) a skewed unimodal response curve but with the optimum shifted to the more mineral‐rich sites; and (5) an increasingly monotonic response curve, the optimum in the extremely base‐rich sites with strong tufa precipitation. Main conclusions Conductivity is the only reliable proxy measure of mineral richness across the entire gradient, within the confines of this study. This information is of great ecological significance in studies of fen mollusc communities. Species richness does not increase with increasing mineral richness along the entire gradient: only a few species are able to dwell in the extremely base‐rich sites. The five types of calcicole–calcifuge behaviour seen in species living in fens have a wider application: data published so far suggest they are also applicable to mollusc communities in other habitats.     

Ecological species replacement of Liolaemus lizards along a habitat gradient

Summary Ecological species replacement of Liolaemus platei by Liolaemus lemniscatus appears to occur along a north-south oriented herb density gradient. Both congeners are the only ground-dwelling lizards in the area; this suggests that L. lemniscatus is an ecological counterpart of L. platei in herbaceous habitats. Relatively longer legs appear to be associated with the utilization of herb-free habitats by L. platei; the shorter legs possessed by L. lemniscatus are associated with the utilization of habitats of higher herb density. Morphological evidence indicates the presence of intermediate populations in the zone of intermediate herb density. Alternatives of a single-species cline versus interspecific hybridization between the two taxa are discussed.     

Gas exchange and resource-use patterns along a Mediterranean successional gradient

Abstract. Gas exchange, leaf-nitrogen concentration and water potential were measured in early and late spring in early successional herbaceous plants occurring after cutting and after fire, and in mature woody species from the Mediterranean climax community Quercetum ilicis in central Italy. Net photosynthesis peaked in early spring in all species studied when values for temperature and light were lower but leaf-nitrogen content was higher as compared to late spring, suggesting that nitrogen more than energy input controlled photosynt-hetic rates. Herbaceous pioneer species occurring after cutting showed higher field photo synthetic capacity than evergreen climax trees and shrubs. By contrast, net photosynthesis of herbaceous species occurring in a persistent stage after fire, was in the same range as that of climax trees. This evidence suggests that carbon-gaining appears to be partly related to the dynamic stage of succession and not solely to the growth form.     

Shifts in patterns of microhabitat occupation by six closely related species of mosses along a complex altitudinal gradient

Summary Changes in patterns of microhabitat occupation were examined for six closely related moss species (family Polytrichaceae) found growing together along a complex altitudinal gradient on the northeast face of Mount Washington, New Hampshire. Little evidence could be found to support the hypothesis that the relative distributions of these six moss species were determined by competitive interactions occurring among them. Instead, the data support the hypothesis that changing patterns in the relative distributions of these six moss species result from differences in microhabitat availability among sites. The moss species appear to behave in an opportunistic manner, occupying a wide array of microhabitats as these microhabitats become available to them.     

The distribution of some heathland plant species along a microtopographic gradient at Dinnet,Scotland, and their dehydration resistances

Along a microtopographic gradient in a heathland, five types of plant distribution were recognized: (i) Pyrola type, restricted to the crest of the rise; (ii) Vaccinium type, mainly on the steeper slopes and thin soil; (iii) Arctostaphylos type, ranging widely from the top to the bottom of the slope; (iv) Erica tetralix type, in the waterlogged wet places; and (v) Calluna type, showing the most extensive occurrence across the ranges of all the foregoing types. A clear separation of habitat was found between Erica cinerea and Erica tetralix. The stomatal transpiration of Calluna was the most active among five species studied. However, the cuticular transpiration rate is rather low. These facts all have a bearing on the widely dominant occurrence of Calluna in heathland. Vaccinium vitis-idaea and Arctostaphylos uva-ursi showed the lowest transpiration rates in the present study. Their cuticular transpiration rates are also low. These facts may be in accord with their inability to overcome Calluna under conditions of adequate water supply. However, they may become dominant in some sites, such as the terrace of thin soil. The buds of Erica tetralix are very susceptible to drought, because their lethal water deficit is very small. It may be a main cause of the restriction of this species to very wet places.     

Bird communities in habitats along a successional gradient: Divergent patterns of species richness,specialization and threat

The long history of human influence on northern temperate landscapes has created a mosaic of successional stages, from closed forest to open grassland. Various species thus adapted to different habitats and it is interesting to explore how these differences in species composition among particular successional stages translate into differences at the community level. For this purpose, we surveyed breeding birds in 233 patches of five different habitats covering a gradient from bare ground to forest in 29 abandoned military training sites scattered throughout the Czech Republic. Linear mixed effects modelling revealed that late-successional habitats (dense scrubland and forest) were the most species-rich, whereas early-successional stages hosted bird communities with the highest habitat specialization and threat level. These results suggest that the habitats of late-successional stages are important for the maintenance of high bird species richness, but that early-successional habitats are essential for highly specialized and threatened bird species. Given the highly adverse impacts of agricultural intensification and land abandonment on open habitats, it is necessary to promote factors creating initial successional stages suitable for specialized and threatened species.