The Habitat Amount Hypothesis implies negative effects of habitat fragmentation on species richness

The habitat amount hypothesis (HAH) predicts that species richness in a habitat site increases with the amount of habitat in the ‘local landscape’ defined by an appropriate distance around the site, with no distinct effects of the size of the habitat patch in which the site is located. It has been stated that a consequence of the HAH, if supported, would be that it is unnecessary to consider habitat configuration to predict or manage biodiversity patterns, and that conservation strategies should focus on habitat amount regardless of fragmentation. Here, I assume that the HAH holds and apply the HAH predictions to all habitat sites over entire landscapes that have the same amount of habitat but differ in habitat configuration. By doing so, I show that the HAH actually implies clearly negative effects of habitat fragmentation, and of other spatial configuration changes, on species richness in all or many of the habitat sites in the landscape, and that these habitat configuration effects are distinct from those of habitat amount in the landscape. I further show that, contrary to current interpretations, the HAH is compatible with a steeper slope of the species–area relationship for fragmented than for continuous habitat, and with higher species richness for a single large patch than for several small patches with the same total area (SLOSS). This suggests the need to revise the ways in which the HAH has been interpreted and can be actually tested. The misinterpretation of the HAH has arisen from confounding and overlooking the differences in the spatial scales involved: the individual habitat site at which the HAH gives predictions, the local landscape around an individual site and the landscapes or regions (with multiple habitat sites and different local landscapes) that need to be analysed and managed. The HAH has been erroneously viewed as negating or diminishing the relevance of fragmentation effects, while it actually supports the importance of habitat configuration for biodiversity. I conclude that, even in the cases where the HAH holds, habitat fragmentation and configuration are important for understanding and managing species distributions in the landscape.     

Group living enhances individual resources discrimination: the use of public information by cockroaches to assess shelter quality

In group-living organisms, consensual decision of site selection results from the interplay between individual responses to site characteristics and to group-members. Individuals independently gather personal information by exploring their environment. Through social interaction, the presence of others provides public information that could be used by individuals and modulates the individual probability of joining/leaving a site. The way that individual's information processing and the network of interactions influence the dynamics of public information (depending on population size) that in turn affect discrimination in site quality is a central question. Using binary choice between sheltering sites of different quality, we demonstrate that cockroaches in group dramatically outperform the problem-solving ability of single individual. Such use of public information allows animals to discriminate between alternatives whereas isolated individuals are ineffective (i.e. the personal discrimination efficiency is weak). Our theoretical results, obtained from a mathematical model based on behavioral rules derived from experiments, highlight that the collective discrimination emerges from competing amplification processes relying on the modulation of the individual sheltering time without shelters comparison and communication modulation. Finally, we well demonstrated here the adaptive value of such decision algorithm. Without any behavioral change, the system is able to shift to a more effective strategy when alternatives are present: the modification of the spatio-temporal distributions of individuals leading to the collective selection of the best resource. This collective discrimination implying such parsimonious and widespread mechanism must be shared by many group living-species.     

Self-amplification as a source of interindividual variability: Shelter selection in cockroaches

Although group effect and collective decisions have been described in many insect species, the behavioral mechanisms involved in the process remain poorly documented at the individual level. We examined how individual behavior depends on the environmental context and we precisely characterized the behavioral rules leading to settlement of individual cockroaches in resting site. We focused on the spatial and temporal distribution of individuals in absence of conspecifics. Using isolated adult males of the cockroach Periplaneta americana, we showed that the quality of resting sites and the duration of the settlement exerted an influence on the individual decision-making: the probability of leaving a resting site decreased with the time spent under a shelter. A numerical model derived from experimental data suggested that this simple rule of self-amplification can also account for the interindividual variability.     

Impact of landscape fragmentation on a specialised woodland bat,Rhinolophus hipposideros

For habitat specialists, fragmentation has major consequences as it means less suitable habitat for the species to live in. In a fragmented landscape, we would expect larger, but spatially more clustered, foraging ranges. We studied the impact of landscape fragmentation on the foraging range and habitat exploitation of a specialised forest bat by radiotracking 16 female lesser horseshoe bats Rhinolophus hipposideros in a landscape with connected woodland structures and in a highly fragmented landscape in Carinthia, Austria.Contrary to our expectations, spatial foraging behaviour was not influenced by fragmentation. No differences in the behaviour of the bats between the sites were evident for the foraging ranges (minimum convex polygon, MCP), the core foraging areas (50% kernel), nor the mean or the maximum distances from the roost. However, in the highly fragmented landscape, the foraging activity of individuals was spatially more clustered and the overall MCP of all bats of a colony was greater compared to the less fragmented landscape.Woodland was the most important foraging habitat for the lesser horseshoe bats at both study sites. Habitat selection at the individual MCPs was evident only at the site with low fragmentation. However, in the core foraging areas, woodland was significantly selected over all other habitat types at both study sites.We conclude that (1) conservation measures for colonies of lesser horseshoe bats should be undertaken within 2.5 km of the nursery roost, (2) woodland is the key foraging habitat particularly in the vicinity of the roost, and (3) any loss of woodland near the colonial roosts are likely to negatively influence the colony, since these bats do not seem to be able to adapt their spatial foraging behaviour in a degraded landscape. The inflexible spatial behaviour of this specialised bat highlights the need to compensate for any habitat loss within the foraging range of a bat colony.     

Home range size scales to habitat amount and increasing fragmentation in a mobile woodland specialist

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Population structure and landscape genetics in the endangered subterranean rodent Ctenomys porteousi

In order to devise adequate conservation and management strategies for endangered species, it is important to incorporate a reliable understanding of its spatial population structure, detecting the existence of demographic partitions throughout its geographical range and characterizing the distribution of its genetic diversity. Moreover, in species that occupy fragmented habitats it is essential to know how landscape characteristics may affect the genetic connectivity among populations. In this study we use eight microsatellite markers to analyze population structure and gene flow patterns in the complete geographic range of the endangered rodent Ctenomys porteousi. Also, we use landscape genetics approaches to evaluate the effects of landscape configuration on the genetic connectivity among populations. In spite of geographical proximity of the sampling sites (8–27 km between the nearest sites) and the absence of marked barriers to individual movement, strong population structure and low values of gene flow were observed. Genetic differentiation among sampling sites was consistent with a simple model of isolation by distance, where peripheral areas showed higher population differentiation than those sites located in the central area of the species’ distribution. Landscape genetics analysis suggested that habitat fragmentation at regional level has affected the distribution of genetic variation among populations. The distance of sampling sites to areas of the landscape having higher habitat connectivity was the environmental factor most strongly related to population genetic structure. In general, our results indicate strong genetic structure in C. porteousi, even at a small spatial scale, and suggest that habitat fragmentation could increase the population differentiation.     

Resting ecology and behaviour of Egyptian mongooses (Herpestes ichneumon) in southwestern Spain

Nocturnal and siesta resting ecology and behaviour of 24 Egyptian mongooses were studied. Mongooses mainly used underground dens (rabbit warrens and badger setts) and thickets in both periods. Tree hollows were occasionally used. Twenty-one individuals used 338 different nocturnal resting sites on 672 occasions. Mean number of resting sites per individual was 16.1, though resting sites used by each individual increased when sample size did. Mean re-use rate was 1.99 nights/site. The majority of nocturnal resting sites were only used once, and very few more than five times. Nocturnal resting sites used two or more times were significantly more often inside than outside home range core areas, whereas resting sites used once were significantly more often outside than inside. Mean distance between consecutive nocturnal resting sites was different among classes of individuals, and annual variations were detected. Eighteen mongooses used 133 different resting sites on 171 occasions during siesta periods. Mean number of different sites used and re-use rate were 7.4 and 1.29, respectively. Types of resting sites used during siesta periods changed throughout the year with underground dens predominating in summer. Re-use rates remained constant for the whole year. The impacts of social structure and summer high temperatures on mongoose resting behaviour are discussed.     

Individual and Group Personalities Characterise Consensus Decision‐Making in an Ant

Non‐human animals can exhibit idiosyncratic behaviour across individuals in much in the same way as humans. Animals with specific personalities may have advantages in some environments, and this idiosyncrasy may thus be of considerable ecological and evolutionary importance. In group‐living organisms, personality can occur at the level of the group as well as that of the individual. However, at present, we have very little understanding of the possible benefits of group‐level personality, and how this is linked with individual personality. In this study, I examine the influence of individual and group personality during the process of colony migration in the Japanese ant, Myrmecina nipponica. These ants use a consensus decision process to decide among alternatives when searching for a new home. Individuals contribute to this process by scouting for new nest sites, recruiting nestmates by laying pheromone trails, and carrying brood to the new site, although whether these roles are consistent among individuals and how roles are distributed within and between colonies remain unclear. Individual contributions to the nest‐site selection process were quantified over five repeated relocations in five colonies. Results demonstrate that contributions to the relocation effort were highly skewed within the colonies and that individuals were consistent in their contributions over repeated relocation events. Furthermore, the distribution of effort differed between colonies, indicating that intercolony differences in composition of behavioural types resulted in colony‐level personality. While these differences did not lead to any detectable difference in relocation performance between colonies in the simple experimental arrangement used, colony personality could influence decision outcomes in more complex environments.     

Habitat loss and fragmentation affecting mammal and bird communities—The role of interspecific competition and individual space use

Fragmentation and loss of habitat are major threats to animal communities and are therefore important to conservation. Due to the complexity of the interplay of spatial effects and community processes, our mechanistic understanding of how communities respond to such landscape changes is still poor. Modelling studies have mostly focused on elucidating the principles of community response to fragmentation and habitat loss at relatively large spatial and temporal scales relevant to metacommunity dynamics. Yet, it has been shown that also small scale processes, like foraging behaviour, space use by individuals and local resource competition are also important factors. However, most studies that consider these smaller scales are designed for single species and are characterized by high model complexity. Hence, they are not easily applicable to ecological communities of interacting individuals. To fill this gap, we apply an allometric model of individual home range formation to investigate the effects of habitat loss and fragmentation on mammal and bird communities, and, in this context, to investigate the role of interspecific competition and individual space use. Results show a similar response of both taxa to habitat loss. Community composition is shifted towards higher frequency of relatively small animals. The exponent and the 95%-quantile of the individual size distribution (ISD, described as a power law distribution) of the emerging communities show threshold behaviour with decreasing habitat area. Fragmentation per se has a similar and strong effect on mammals, but not on birds. The ISDs of bird communities were insensitive to fragmentation at the small scales considered here. These patterns can be explained by competitive release taking place in interacting animal communities, with the exception of bird's buffering response to fragmentation, presumably by adjusting the size of their home ranges. These results reflect consequences of higher mobility of birds compared to mammals of the same size and the importance of considering competitive interaction, particularly for mammal communities, in response to landscape fragmentation. Our allometric approach enables scaling up from individual physiology and foraging behaviour to terrestrial communities, and disentangling the role of individual space use and interspecific competition in controlling the response of mammal and bird communities to landscape changes.     

Interactive effects of landscape and weather on dispersal

Over the last decades, many species have been forced to track their shifting climate envelopes, and at the same time man‐induced landscape fragmentation has led to the global decrease of natural habitat availability and connectivity. The interaction between these two co‐occurring global environmental changes might have very strong effects on biodiversity that are still understudied. Species‐specific responses to these environmental changes critically depend on individual dispersal, either to track suitable climatic conditions or to cope with landscape fragmentation. Here we study how dispersal in an ectotherm is affected by interactions between landscape fragmentation and weather conditions. We show that both the emigration rates out of suitable habitats and the topology of the trajectory of dispersing individuals were affected by temperature and landscape fragmentation. The emigration rate was temperature‐dependent in fragmented landscapes, with butterflies emigrating more at high temperatures. The emigration rate was temperature insensitive in more continuous landscapes. Move length was farther at low temperatures and less at high temperatures in fragmented landscapes. Move length was less at low temperatures and farther at high temperatures in more continuous landscapes. To our knowledge only two recent studies have documented patterns of interactions between climate and fragmentation, despite the fact that they are the two main drivers of biodiversity loss worldwide. Here, we go a step further by providing mechanistic explanations to such patterns.     

Spatial and temporal variation in flight morphology in the butterfly Melitaea cinxia (Lepidoptera: Nymphalidae)

Some studies on insects have found a relationship between habitat structure and investment in dispersal-related traits. In this study we compared the morphology of the butterfly Melitaea cinxia from five sites on the large Baltic island Öland that differed markedly in degree of fragmentation and size. Both wild-caught adults and individuals reared in a split-plot design were compared. We found significant site differences in size-adjusted thorax mass and total body mass. Male thorax mass was on average larger among the sites with the highest degree of habitat fragmentation. However, due to significant sex–site interactions, males and females may have adapted differently to the habitat fragmentation. Using museum specimens, we also analysed changes in morphology, finding an increase in size (measured as head and thorax width) over time. Thorax width appears to have increased among females and decreased among males. Possible explanations include increasing fragmentation of the landscape and changes in population density.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 445–453.     

Comparative Population Genetic Structure of the Endangered Southern Brown Bandicoot,Isoodon obesulus,in Fragmented Landscapes of Southern Australia

Genetic connectivity is a key factor for maintaining the persistence of populations in fragmented landscapes. In highly modified landscapes such us peri-urban areas, organisms’ dispersal among fragmented habitat patches can be reduced due to the surrounding matrix, leading to subsequent decreased gene flow and increased potential extinction risk in isolated sub-populations. However, few studies have compared within species how dispersal/gene flow varies between regions and among different forms of matrix that might be encountered. In the current study, we investigated gene flow and dispersal in an endangered marsupial, the southern brown bandicoot (Isoodon obesulus) in a heavily modified peri-urban landscape in South Australia, Australia. We used 14 microsatellite markers to genotype 254 individuals which were sampled from 15 sites. Analyses revealed significant genetic structure. Our analyses also indicated that dispersal was mostly limited to neighbouring sites. Comparisons of these results with analyses of a different population of the same species revealed that gene flow/dispersal was more limited in this peri-urban landscape than in a pine plantation landscape approximately 400 km to the south-east. These findings increase our understanding of how the nature of fragmentation can lead to profound differences in levels of genetic connectivity among populations of the same species.     

社区水平森林景观格局动态特征与驱动因素

基于高分辨率SPOT-5影像、TM影像和农户调查数据,以及相关辅助数据,选取三峡库区典型森林大县石柱县三星乡石星村为样区,利用景观格局指数与Logistic回归分析法,对1994—2014年间的森林景观格局的动态特征及其影响因素进行分析,旨在丰富人们对社区水平森林景观格局动态的理解和认识,为森林景观恢复和可持续经营策略的制定提供科学依据。结果表明:(1)样区森林景观一直占据20年间景观基质的主导地位,空间分布上退化原始林主要集中分布在中东部低山、深丘区,次生林、退化林地和人工林呈镶嵌格局,分散于主要基质性景观中;(2)20年间样区森林景观整体呈先减少后增加的"U"型格局,表现为1994—2004年间的退化与2004—2014年间的恢复两个截然相反的过程,且这一过程主要发生在森林景观与水田、旱地、居民点、道路等接壤区;(3)2004—2014年间森林景观的演化趋势与1994—2004年间整体上呈反向趋势,空间格局上,这一阶段森林景观的恢复主要出现在1994—2004年间森林景观所发生退化的区域;(4)20年间样区森林景观在斑块水平上呈明显异质性分布,3个截面年份不同森林景观破碎化程度的总排序均为退化林地人工林次生林退化原始林,且退化原始林、次生林和退化林地的破碎度变化趋势相同,均为先增后减;(5)20年间样区森林景观在景观水平上的破碎度呈现先增强后减弱趋势,且2014年的破碎化程度恢复至低于1994年的水平,进一步说明在退化后的森林景观恢复过程中,生态完整性的恢复速度远低于破碎速度;(6)社会经济因素对20年间样区森林景观变化的影响最突出,其中,最主要的影响因素是人口密度变化,先增后减的变化使生态压力随之增减,进而导致森林景观的退化与恢复。其次是农民人均纯收入变化,特别是2004—2014年农民人均纯收入的增加,促进能源结构的转型,减少了对森林景观的依赖和干扰。空间距离对森林景观的影响具有稳定的持续作用,居民对森林景观的干扰多经由距离因素而实现。自然因素控制大地貌格局,中短时期内对森林景观变化影响不大,次生林和退化林地因处于适宜开发的边缘地带,受自然环境影响较大。     

黔中喀斯特山地城市景观稳定性评价与特征分析

城市景观稳定性是对城市生态环境进行分析和评价的基础,对城市景观生态安全格局和景观可持续性具有十分重要的意义。以典型喀斯特山地城市贵阳市为研究对象,以2008年、2013年和2017年景观类型为主要数据。运用蔓延度指数、斑块密度和总边缘对比度构建景观稳定性评价模型,在分析2008-2017年景观格局变化状况的基础上评价景观稳定性时空特征。并运用探索性空间数据分析方法揭示贵阳市景观稳定性时空演变规律和冷热点特征。结果表明:(1)2008-2017年贵阳市景观格局发生剧烈变化,耕地面积大幅减少,林地和建设用地面积持续增加,景观破碎化程度加剧,但破碎化趋势有所减缓。(2)10年间,贵阳市景观稳定性水平整体较低,以不稳定和较不稳定为主,表现出"先降低、后提高,总体略有降低"的趋势。(3)贵阳市景观稳定性的空间分布与演变具有明显空间聚集效应,全局Moran''s I指数介于0.2008-0.4005之间,但聚集趋势呈"总体减弱,小幅提高"的特征;此外,景观稳定性热点区主要集中在城镇建设用地或林地斑块相对集中连片、完整的区域。研究结果可为喀斯特山地城市景观空间格局优化、生态安全网络格局构建和生态文明城市建设提供重要的科学依据。     

Patterns of resource tracking by avian frugivores at multiple spatial scales: two case studies on discordance among scales

Scaling is relevant for the analysis of plant‐frugivore interaction, since the ecological and evolutionary outcomes of seed dispersal depend on the spatial and temporal scale at which frugivory patterns emerge. We analyse the relationship between fruit abundance and frugivore activity at local and landscape spatial scales in two different systems composed, respectively, by the bird‐dispersed woody plants Juniperus communis and Bursera fagaroides, and their frugivore assemblages. We use a hierarchical approach of nested patchiness of fruit‐resource, where patches are defined by individual plants within site, at the local scale, and by sites within region, at the landscape scale. The structure of patches is also described in terms of contrast (differences in fruit availability among patches) and aggregation (spatial distribution of patches). For J. communis, frugivore activity was positively related to fruit availability at the landscape scale, this pattern seldom emerging at the local scale; conversely, B. fagaroides showed a general trend of positive local pattern that disappeared at the landscape scale. These particular trends might be partially explained by differences in contrast and aggregation. The strong contrast among plants within site together with a high aggregation among sites would promote the B. fagaroides pattern to be only local, whereas in J. communis, low aggregation among sites within region would favour a sharp landscape‐scale pattern. Both systems showed discordant patterns of fruit‐resource tracking among consecutive spatial scales, but the sense of discordance differed among systems. These results, and the available multi‐scale frugivory data, suggest that discordance among successive scales allows to link directly frugivory patterns to resource‐tracking mechanisms acting at particular scales, resulting, thus, more informative than concordance observational data, in which landscape patterns might result from accumulated effect of local mechanisms. In this context, we propose new methodological approaches for a better understanding of the hierarchical behavioural mechanisms underpinning the multi‐scale resource tracking by frugivores.     

Swarm Site Fidelity in the Sex Role-Reversed Dance Fly Empis borealis

In the dance fly species Empis borealis (L.), females (1–40) gather to swarm at landmarks (swarm markers, like trees and bushes), and males carrying an insect prey visit these swarms for mating. We noticed earlier that some swarm sites were used for several years and that they appeared to be frequented by a similar number of swarming females in each year, although the numbers of females varied greatly among swarm sites and certain sites attracted more swarming individuals than others. To explore swarm site fidelity in this mating system, in 1993 we monitored the same swarm sites that we studied in 1989, addressing the questions, Would the same swarm sites still attract the same number of females and males after 4 years? and Why do some swarm sites attract more displaying females than others? The number of females swarming at the different markers in 1993 was approximately the same as 4 years earlier. Some of these swarm sites are known to have been used for 18 years. The swarm sites with the largest number of flies had a high sun exposure during the day and were found at coniferous swarm marker trees and in a mixed forest habitat. A swarm site with few females attending and with a low amount of insolation during the day can be predicted to be abandoned as a swarming site soon. Empis borealis swarm sites thus persist over many years and are attended by a similar number of individuals each year. To our knowledge, such site fidelity has not been demonstrated for any swarming insect species earlier.     

Social flexibility and social evolution in mammals: a case study of the African striped mouse (Rhabdomys pumilio)

Environmental change poses challenges to many organisms. The resilience of a species to such change depends on its ability to respond adaptively. Social flexibility is such an adaptive response, whereby individuals of both sexes change their reproductive tactics facultatively in response to fluctuating environmental conditions, leading to changes in the social system. Social flexibility focuses on individual flexibility, and provides a unique opportunity to study both the ultimate and proximate causes of sociality by comparing between solitary and group-living individuals of the same population: why do animals form groups and how is group-living regulated by the environment and the neuro-endocrine system? These key questions have been studied for the past ten years in the striped mouse Rhabdomys pumilio. High population density favours philopatry and group-living, while reproductive competition favours dispersal and solitary-living. Studies of genetic parentage reveal that relative fitness of alternative reproductive tactics depends on the prevailing environment. Tactics have different fitness under constrained ecological conditions, when competitive ability is important. Under conditions with relaxed ecological constraints, alternative tactics can yield equal fitness. Both male and female striped mice display alternative reproductive tactics based on a single strategy, i.e. all individuals follow the same decision rules. These changes are regulated by endocrine mechanisms. Social flexibility is regarded as an adaptation to unpredictably changing environments, selecting for high phenotypic flexibility based on a broad reaction norm, not on genetic polymorphism for specific tactics.     

Where to sleep in a rural landscape? A comparative study of resting sites pattern in two syntopic Martes species

The niche‐complementarity hypothesis predicts that two sympatric species must differ in their requirements for one of the three main ecological dimensions (i.e. habitat use, diet, and activity time) to coexist. European pine marten Martes martes and stone marten M. foina are syntopic medium‐sized mustelids with very similar morphology and ecology for which resting sites are a key resource. To better understand how these species coexist, we investigated whether key features of their resting site pattern (number of resting sites, area over which they are distributed, main habitat type used for resting) differed. We used diurnal telemetry to identify resident individuals (e.g. spatially stable individuals over time) and to locate them during resting periods in a fragmented forested area in France. Stone marten used fewer resting sites distributed over a smaller surface area than pine marten. Most stone marten resting sites were located in open habitat (83%) in the proximity of human habitations, whereas pine martens rested almost exclusively in forest (98%). Sex, age, and season explained some variability in both the number of resting sites and the probability of resting within forested habitat for stone marten but not pine marten. The area covered by resting sites was larger in males than in females, but age modulated this difference in an opposite way for the two species. Such a pattern was expected given the intra‐sexual territoriality and the reproductive phenology of these species. Overall, stone marten showed higher inter‐individual variability in resting site pattern than pine marten. The particular pattern observed in subadult male stone martens during summer (increase in resting site surface area and in the probability to rest in forest) may reflect an attempt to settle in forests, and we discuss these implications in the context of interspecific competition.     

Time budgets of Tibetan eared pheasants during the non-breeding season in an alpine scrub habitat

Time allocations of the group-living Tibetan eared pheasants Crossoptilon harmani with and without supplementary food were investigated by full-day sampling from winter through spring in an alpine scrub zone,south Tibet.At a flock scale,both the different food-supply flocks displayed similar daily patterns of activity,foraging in the morning and evening,and resting around midday.In terms of individual activity,either foraging or resting was highly synchronous with the flock's.Non-provisioned birds spent mor...     

Variation in local carrying capacity and the individual fate of bacterial colonizers in the phyllosphere

Using a phyllosphere model system, we demonstrated that the term ‘carrying capacity'', as it is commonly used in microbial ecology, needs to be understood as the sum of many ‘local carrying capacities'' in order to better explain and predict the course and outcome of bacterial colonization of an environment. Using a green fluorescent protein-based bioreporter system for the quantification of reproductive success (RS) in individual Erwinia herbicola cells, we were able to reconstruct the contribution of individual immigrants to bacterial population sizes on leaves. Our analysis revealed that plant foliage represents to bacteria an environment where individual fate is determined by the local carrying capacity of the site where an immigrant cell lands. With increasing inoculation densities, the RS of most immigrants declined, suggesting that local carrying capacity under the tested conditions was linked to local nutrient availability. Fitting the observed experimental data to an adapted model of phyllosphere colonization indicated that there might exist three types of sites on leaves, which differ in their frequency of occurrence and local carrying capacity. Specifically, our data were consistent with a leaf environment that is characterized by few sites where individual immigrants can produce high numbers of offspring, whereas the remainder of the leaf offered an equal number of sites with low and medium RS. Our findings contribute to a bottom–up understanding of bacterial colonization of leaf surfaces, which includes a quantifiable role of chance in the experience at the individual level and in the outcome at the population level.