Resource partitioning among the fishes of rainforest streams in Sri Lanka

The fish assemblage typical of small rainforest streams of Sri Lanka was investigated to see if the fishes were characterized by a high degree of specialization, expected of equilibrium communities, or if they were relatively unspecialized, expected of more stochastic (non-equilibrium) communities. Morphological features, habitats, microhabitats and diets of the 20 most abundant fish species were measured. The fishes included 11 Cyprinidae (including seven Barbus spp. and two Rasbara spp.), two Gobiidae, two Channidae, and one species each in the Cobitidae, Cyprinodontidae, Belontidae, Belonidae, and Mastacembelidae. Microhabitats were determined by measuring (for 6760 fish) water column depth, distance of fish from bottom, mean water column velocity, water velocity at fish, and substrate. Morphologically, the species showed a high degree of specialization, especially in structures related to feeding. Although several species were habitat generalists, most species occurred in distinct habitats. Within habitats, microhabitat overlap among co-occuring species was low, particularly in relation to position in the water column. Principal component analysis of the microhabitat measurements produced three new variables. Most species not clearly segregated by the overlap analysis showed segregation on the new variables. Fishes not segregated by habitat or microhabitat tended to show low dietary overlaps. Specialization in feeding habits was more pronounced among the Sri Lankan fishes than noted for fishes in small rainforest streams elsewhere, in part because of the greater reliance of the Sri Lanka fishes on autochthonous foods. Overall, the fish assemblage had the characteristics expected of an equilibrium (deterministic) assemblage.     

南岭国家级自然保护区常绿阔叶林主要树种的种间分离

种间分离的研究对揭示种间相互作用、群落组成与动态具有重要意义。本研究在南岭国家级自然保护区的常绿阔叶林内,设置了36个样方,以样方内株数≥10的27个物种为研究对象,在样方调查的基础上,运用ArcGIS软件、N×N最近邻体列联表、2×2列联表截表以及Pielou分离指数,研究了群落内27个物种的种间分离规律,采用Clark_Evans最近邻体法测定了27个物种的分布格局。结果显示:(1)27个物种中,有3个呈聚集分布,24个呈随机分布;(2)常绿阔叶林植物群落中正分离种对占59.83%,随机毗邻种对占40.17%,没有负分离,群落稳定;(3)正分离主要发生在群落优势种或建群种之间,优势种之间也有随机毗邻发生;(4)呈聚集分布的物种与其它物种主要表现为正分离,一般不表现为负分离;而呈随机分布的物种之间多数表现为随机毗邻,也有部分表现为正分离或负分离;(5)群落内27个物种互相交错分布,是全面不分离的。     

Ecomorphological correlates in tropical stream fishes of southern India

Synopsis I studied ecomorphological correlations in 18 stream fish species (belonging to five families) commonly occurring in diversity rich habitats of rivers in the central Western Ghats (Southern India). Univariate and multivariate analyses indicated a clear segregation of species in morphological and ecological (habitat) space. Cluster analyses using morphological features segregated species based mainly on their body shape and feeding parameters while habitat parameters segregated species according to their position along the water column and preference to runs or pool habitats. Principal component analyses showed that body size, head dimensions, mouth position and eye diameter were important characters for species segregation in morphological space while run or pool habitat type and the position of the fish in the water column best explained segregation of species in habitat space. Consistencies in some of the species clusters obtained from analyses on morphological characteristics and habitat preferences warrant closer examination of the relationships between them. These were studied employing multivariate methods like canonical correspondence analysis and the Mantel test. The tests revealed weak correlations indicating that in addition to morphology, other factors like behavioural, physiological and evolutionary constraints together are likely to play an important role in ecosystem structuring of these complex tropical assemblages.     

Non-random co-occurrence of native and exotic plant species in Mediterranean grasslands

Invasion by exotic species in Mediterranean grasslands has determined assembly patterns of native and introduced species, knowledge of which provides information on the ecological processes underlying these novel communities. We considered grasslands from Spain and Chile. For each country we considered the whole grassland community and we split species into two subsets: in Chile, species were classified as natives or colonizers (i.e. exotics); in Spain, species were classified as exclusives (present in Spain but not in Chile) or colonizers (Spanish natives and exotics into Chile). We used null models and co-occurrence indices calculated in each country for each one of 15 sites distributed along a precipitation gradient and subjected to similar silvopastoral exploitation. We compared values of species co-occurrence between countries and between species subsets (natives/colonizers in Chile; exclusives/colonizers in Spain) within each country and we characterised them according to climatic variables. We hypothesized that: a) the different coexistence time of the species in both regions should give rise to communities presenting a spatial pattern further from random in Spain than in Chile, b) the co-occurrence patterns in the grasslands are affected by mesoclimatic factors in both regions. The patterns of co-occurrence are similar in Spain and Chile, mostly showing a spatial pattern more segregated than expected by random. The colonizer species are more segregated in Spain than in Chile, possibly determined by the longer residence time of the species in the source area than in the invaded one. The segregation of species in Chile is related to water availability, being species less segregated in habitat with greater water deficit; in Spain no relationship with climatic variables was found. After an invasion process, our results suggest that the possible process of alteration of the original Chilean communities has not prevented the assembly between the native and colonizer species together.     

山西稀有濒危植物脱皮榆群落的种间分离

通过对山西太岳山脱皮榆群落1 hm²样地内出现的14种乔木进行调查,在应用ArcGIS软件分析每个个体的最近邻体植株的基础上,采用N×N最近邻体列联表及其2×2列联表截表的方法和Pielou分离指数计算分析了脱皮榆群落的种间分离规律,并根据种间分离情况以及物种对环境的适应方式,利用PCA排序将样地中14个物种进行了生态种组的划分。结果表明:（1）该群落中出现的14种乔木构成的91个种对中,随机毗邻种对占绝大多数（74.73%）,正分离种对较少（23.07%）,负分离种对极少（2.20%）,群落趋于稳定。（2）群落的种间分离在不同种之间存在着一定的差异,脱皮榆仅与其他树种随机毗邻;正分离常发生在群落中的优势种之间,而负分离常发生在群落中的伴生种之间。（3）种间分离与群落的生境异质性和物种的生态需求具有密切的关系,即两个具有不同生境要求的物种之间常常发生正分离,而具有相同或相似生境要求的物种之间常常发生负分离。（4）脱皮榆群落中14种乔木可划分为3个生态种组。研究认为,脱皮榆群落内物种的总体分离规律为全面不分离。由生境异质性、物种间不同的生物学特性和生态需求为基础的种间关系可能是维持太岳山脱皮榆群落稳定共存的机制之一。     

Static habitat partitioning and dynamic selection by sympatric young Atlantic salmon and brown trout in south-west England streams

Direct underwater observation of micro‐habitat use by 1838 young Atlantic salmon Salmo salar [mean L_T 7·9 ± 3.1(s.d.) cm, range 3·19] and 1227 brown trout Salmo trutta (L_T 10·9 ± 5·0 cm, range 3·56) showed both species were selective in habitat use, with differences between species and fish size. Atlantic salmon and brown trout selected relatively narrow ranges for the two micro‐habitat variables snout water velocity and height above bottom, but with differences between size‐classes. The smaller fishes <7 cm held positions in slower water closer to the bottom. On a larger scale, the Atlantic salmon more often used shallower stream areas, compared with brown trout. The larger parr preferred the deeper stream areas. Atlantic salmon used higher and slightly more variable mean water velocities than brown trout. Substrata used by the two species were similar. Finer substrata, although variable, were selected at the snout position, and differences were pronounced between size‐classes. On a meso‐habitat scale, brown trout were more frequently observed in slow pool‐glide habitats, while young Atlantic salmon favoured the faster high‐gradient meso‐habitats. Small juveniles <7 cm of both species were observed most frequently in riffle‐chute habitats. Atlantic salmon and brown trout segregated with respect to use of habitat, but considerable niche overlap between species indicated competitive interactions. In particular, for small fishes <7 cm of the two species, there was almost complete niche overlap for use of water depth, while they segregated with respect to water velocity. Habitat suitability indices developed for both species for mean water velocity and water depth, tended to have their optimum at lower values compared with previous studies in larger streams, with Atlantic salmon parr in the small streams occupying the same habitat as favoured by brown trout in larger streams. The data indicate both species may be flexible in their habitat selection depending on habitat availability. Species‐specific habitat overlap between streams may be complete. However, between‐species habitat partitioning remains similar.     

Resource partitioning by three species of newts during their aquatic phase

This paper describes the multidimensional niche of syntopic smooth newts Triturus vulgaris. alpine newts Triturus alpestris. and Italian crested newts Triturus carnifex. which constitute a complex community of eight components (the three species and their larvae, plus juvenile and neotenic alpine newts) All the potentially limiting resources (seasonal time, diel time, habitat and food) are analyzed as single niche dimensions, and then the real multidimensional niche is calculated Resource states are objectively defined, using an algorithm that groups the significant resource categories The larvae were segregated by prey, season, habitat, and water depth, in order of decreasing importance, their overlap was very low when their complete multidimensional niche was considered, and therefore they have few opportunities for present–day competition The adults were largely segregated by season and water depth, and to a lesser degree by habitat and prey, their multidimensional niches overlapped to a certain degree, thus leaving conditions for ongoing competition Considenng all the species and the life stages together, the order of importance for segregation was season, prey, habitat, water depth and diel activity, with a low overlap in the complete niche Differences m prey size were probably due to morphological constraints, differences in diel activity had very little effect on segregation except m separating larvae from the adults, and were likely promoted by need to avoid mutual predation and cannibalism It remains to be tested whether seasonal, habitat, and water depth differences are due to competitive pressures or to ecophysiological constraints     

Isotopes reveal contrasting water use strategies among coexisting plant species in a Mediterranean ecosystem

Variation in the stable carbon and oxygen isotope composition (δ(13) C, Δ(18) O) of co-occurring plant species may reflect the functional diversity of water use strategies present in natural plant communities. We investigated the patterns of water use among 10 coexisting plant species representing diverse taxonomic groups and life forms in semiarid southeast Spain by measuring their leaf δ(13) C and Δ(18) O, the oxygen isotope ratio of stem water and leaf gas exchange rates. Across species, Δ(18) O was tightly negatively correlated with stomatal conductance (g(s) ), whereas δ(13) C was positively correlated with intrinsic water use efficiency (WUE(i) ). Broad interspecific variation in Δ(18) O, δ(13) C and WUE(i) was largely determined by differences in g(s) , as indicated by a strong positive correlation between leaf δ(13) C and Δ(18) O across species The 10 co-occurring species segregated along a continuous ecophysiological gradient defined by their leaf δ(13) C and Δ(18) O, thus revealing a wide spectrum of stomatal regulation intensity and contrasting water use strategies ranging from 'profligate/opportunistic' (high g(s) , low WUE(i) ) to 'conservative' (low g(s) , high WUE(i) ). Coexisting species maintained their relative isotopic rankings in 2?yr with contrasting rainfall, suggesting the existence of species-specific 'isotopic niches' that reflect ecophysiological niche segregation in dryland plant communities.     

ENVIRONMENTAL AND PHYSIOLOGICAL FACTORS INFLUENCING THE NATURAL DISTRIBUTION OF EVERGREEN AND DECIDUOUS ERICACEOUS SHRUBS ON NORTHEAST AND SOUTHWEST SLOPES OF THE SOUTHERN APPALACHIAN MOUNTAINS. I. IRRADIANCE TOLERANCE

Deciduous and evergreen species are segregated on northeast and southwest slopes of the southern Appalachian Mountains. The segregated distributions of three ericaceous shrubs (Rhododendron maximum valley positions; Rhododendron periclymenoides on northeast slopes; Kalmia latifolia on southwest slopes) were compared to the respective irradiance environments. Growth patterns of field plants, and photosynthetic acclimation of each species to three irradiance treatments in a phytotron were studied. Rhododendron maximum, an evergreen species, was found to be most sensitive to high radiation. In phytotron experiments, quantum yield, light saturated photosynthetic capacity, photosynthesis per chlorophyll, and water use efficiency decreased at high ambient irradiance for R. maximum. These characteristics limit the growth of R. maximum on high irradiance southwestern slopes. Both K. latifolia and R. periclymenoides were able to improve their photosynthetic performance at high ambient irradiance. Rhododendron periclymenoides, a deciduous species, was found to continue increasing leaf conductance at high irradiance without an increase in photosynthesis indicating a possible limitation by water in high light environments such as southwest slopes. Kalmia latifolia, an evergreen species, had reduced photosynthetic capacity and reduced water use efficiency when grown in low irradiance conditions which coincides with the higher K. latifolia abundance on high light, southwestern slopes.     

Coexistence and Habitat Preference of Two Honeyeaters and a Sunbird on Lombok,Indonesia

On the island of Lombok, Indonesia, three nectarivorous birds partially coexist: the two closely related and very similar Lichmera honeyeaters and a sunbird (Cinnyris jugularis). We investigated how these species segregated ecologically in areas where they coexisted by evaluating foraging visits and aggressive interactions at rich and poor nectar resources in different habitats (forest and open areas) along an altitudinal gradient (800–1600 m asl). The two honeyeaters were partially segregated by altitude. In the zone of overlap, Lichmera limbata dominated the richest forest resources, while Lichmera lombokia dominated the richest resources in open land. The sunbird, C. jugularis, was only observed in open habitats and mostly at poor resource patches. In the three‐species community in open habitats a dominance hierarchy was apparent with L. lombokia as the superior species and C. jugularis as the inferior species. Studies on how segregated species interact in their transition zone can help us to understand interactions between otherwise ecologically segregated species.     

Food and habitat partitioning between young-of-year alewives and rainbow smelt in southeastern Lake Ontario

Synopsis Thermal and depth distributions, diets and time of feeding of young-of-year (YOY) alewives and YOY rainbow smelt were compared for evidence of resource partitioning in southeastern Lake Ontario. YOY alewives were largely epilimnial during August and September, but moved toward the bottom during fall turnover. Alewives were most abundant in the warmest available water. YOY rainbow smelt were concentrated at depths between 10 and 30 m in August and September, but moved into deeper water at fall turnover. Depth distribution of YOY smelt was correlated with temperatures of 8–12°C Both species fed predominantly during day on zooplankton during August and September. Cyclopoid copepods were the most common prey, but bosminids, eubosminids, and occasionally calanoid copepods were frequently eaten. As YOY rainbow smelt grew (> 60 mm), they consumed more Mysis relicta and amphipods, which became the major prey of rainbow smelt by November. YOY alewives consumed mostly zooplankton in all months. Diet overlap of the two species was greatest in warm water (> 12.0° during October (94.3% similarity) and August (80.0% similarity) and lowest in November (16.9% similarity). Positive size-selection on zooplankton was found in all months for YOY rainbow smelt, but only in late September through November for YOY alewives. Thus, during thermal stratification, the species were spatially segregated by water temperature but had a high degree of overlap in time of feeding and types of prey eaten. In contrast, after fall turnover there was a greater separation in diet but a higher overlap in habitat use.     

Seed banks of a southern Australian wetland: the influence of water regime on the final floristic composition

The sediments of a southern Australian wetland complex were investigated to determine the germinable seed bank. The number of seeds ranged from 22,000 to 78,000 m^–2. Non-metric multi dimensional scaling (NMS) ordination based upon species composition separated the four basins but each contained the representatives of the major functional groups: terrestrial, amphibious and submerged. To test the influence of water regime on the final floristic composition derived from these seed banks, sediments were positioned at three elevations (0, 30 and 80 cm) and subjected to three hydrologic regimes (static water level, and draw down rates of 2.5 and 7 cm week^–1). Species compositions were analysed after 98 days via ordination. Despite significant differences in the initial seed bank composition the final floristic compositions were correlated with the water regime and independent of the initial seed bank composition. Species groups were segregated on a basis of whether sediments were continuously exposed to the atmosphere, the rate of draw down and the water depth. Moist sediments were dominated by species belonging to all the main functional groups. Sediments subject to rapid drying were dominated by terrestrial species whereas sediments that were flooded for the majority of the time were dominated by submerged species.     

Difference in cuticular transpiration and sclerophylly in juvenile and adult pine needles relates to the species-specific rates of development

Pinus species show remarkable ontogenetic differences in needle morphology (heterophylly) between juvenile and adult vegetative phases. This developmental shift may play an adaptative role in their success under diverse habitats. As a first step to know the functional differences between each vegetative phase, we compared water loss through the cuticles of juvenile and adult needles of 21-month-old nursery-grown seedlings of nine hard pine species. Cuticular transpiration (CT), calculated after complete stomatal closure, was obtained by leaf-drying curves, and was related to leaf, ontogenetic and climatic parameters. The rate of cuticular transpiration (RCT) between juvenile and adult needles differed across pine species, and in particular segregated the Mediterranean species Pinus canariensis and P. halepensis, from the Eurasian P. uncinata and introduced species P. radiata. For these species, RCT was always higher in juvenile needles. The different leaf and ontogenic parameters studied were correlated with the variation in RCT among the nine pine species. We discuss this relationship in the light of the species ecology. Besides their possible adaptive interpretation, these results suggest an underlying need to consider the ontogenetic heterophylly when assessing functional traits in hard pine seedlings, in particular those traits that govern water relations.     

Body mass‐related changes in mammal community assembly patterns during the late Quaternary of North America

The late Quaternary of North America was marked by prominent ecological changes, including the end‐Pleistocene megafaunal extinction, the spread of human settlements and the rise of agriculture. Here we examine the mechanistic reasons for temporal changes in mammal species association and body size during this time period. Building upon the co‐occurrence results from Lyons et al. (2016) – wherein each species pair was classified as spatially aggregated, segregated or random – we examined body mass differences (BMD) between each species pair for each association type and time period (Late Pleistocene: 40 000 ¹⁴C–11 700 ¹⁴C ybp, Holocene: 11 700 ¹⁴C–50 ybp and Modern: 50–0 yr). In the Late Pleistocene and Holocene, the BMD of both aggregated and segregated species pairs was significantly smaller than the BMD of random pairs. These results are consistent with environmental filtering and competition as important drivers of community structure in both time periods. Modern assemblages showed a breakdown between BMD and co‐occurrence patterns: the average BMD of aggregated, segregated and random species pairs did not differ from each other. Collectively, these results indicate that the late Quaternary mammalian extinctions not only eliminated many large‐bodied species but were followed by a re‐organization of communities that altered patterns of species coexistence and associated differences in body size.     

Microhabitat partitioning in a diverse assemblage of darters in the Allegheny River system

Synopsis Habitat partitioning among eleven species of darters (Percidae: Etheostomatini) from the Allegheny River system was studied through underwater observation. Percina caprodes and Percina copelandi showed consistent segregation from Etheostoma by occupying deeper habitats. Substrate size, depth, and water velocity were important variables by which Etheostoma species segregated. Analysis of niche breadth values indicated that species differed widely in their degree of specialization in habitat use; based on the variables measured, Etheostoma zonale was a habitat generalist whereas Etheostoma camurum, Etheostoma tippecanoe, and Percina caprodes tended towards habitat specialization. Habitat segregation appears to be an important mechanism allowing the coexistence of these closely related and ecologically similar species. Microhabitat quantification on a fine scale was important in discovering habitat differences in this diverse system.     

Size distribution of the amphipod Paramoera walkeri (Stebbing) along a depth gradient in Antarctica

Body weight distributions of the amphipod Paramoera walkeri colonizing litter-bags at different depths (2, 5, and 10 m) in Terra Nova Bay were compared. This species dominated the vagile benthic community and was found nearly alone at the shallow sites. At the deepest site it co-occurred with two smaller benthic species (the gastropod Laevilitorina antarctica and the isopod Munna antarctica) but still accounted for most of the animal biomass (80.2%). The frequency size-class distributions of the three dominant species were sharply distinct in a condition of stable coexistence. The total biomass was inversely related to the abundance of animals and diminished with increasing depth. Both species and size-class diversities were higher in deep water which offered a more diversified and favourable habitat than shallow water. At the shallow sites only the large size classes of P. walkeri were present. Differences in the mean individual ash content between samples support the hypothesis that different size individuals of P. walkeri segregated along the depth gradient on the basis of their metabolic requirements and the related ability to exploit sub-optimal environments.     

山西五鹿山白皮松群落乔灌层的种间分离

种间分离的研究对于揭示种间相互作用、群落组成与动态具有重要意义。为了探讨五鹿山自然保护区白皮松林内物种间的关系及其共存机制,运用ArcGIS软件、N×N最近邻体列联表及其2×2列联表截表的方法、Pielou的分离指数,Hegyi单木竞争指数对山西五鹿山白皮松群落55个样方内的15种乔木和灌木的种间分离规律及9种乔木的种内种间竞争关系进行研究,并根据种间分离情况以及物种对环境的适应方式,将样地中15个物种划分为3个生态种组。结果表明:(1)该群落中随机毗邻种对占绝大多数(80%),正分离种对较少(16.19%),负分离种对极少(3.81%),群落趋于稳定。(2)种间分离在不同种之间存在着一定的差异,正分离常发生在群落中的建群种或优势种之间;而负分离常发生在群落中的优势种和一些伴生种之间。(3)物种的种间分离状况与其种内种间竞争及物种的空间分布格局具有密切的关系,正分离常常发生在种内竞争大于种间竞争呈聚集分布的优势种或建群种之间;而负分离常常发生在种间竞争大于种内竞争的优势种和伴生种之间,且同一生态种组内的物种种间竞争较不同生态种组间激烈。(4)种间分离与群落的生境异质性和物种的生态需求也具有密切的关系,即两个具有不同生境要求的物种之间常常发生正分离;而具有相同或相似生境要求的物种之间常常发生负分离。(5)白皮松群落内物种的总体分离规律为全面不分离。(6)由生境异质性和物种间不同的生态适应性引起的种内种间关系是维持五鹿山自然保护区白皮松群落稳定共存的机制。     

Centaurea subsect. Phalolepis in Southern Italy: ongoing speciation or species overestimation? Genetic evidence based on SSRs analyses

In this paper, we investigated a set of narrow endemics of Centaurea subsect. Phalolepis from the mountains of South Italy (mainly Calabria and Salento), segregated from the widespread species Centaurea deusta, using microsatellite (SSR) markers. The goal was to analyse the genetic makeup (levels and structure) of C. deusta and the segregated species and verify whether genetic clusters were in agreement with current classification of the species. With C. deusta, we also carried out an ecological niche modelling (ENM) analysis to check its potential distribution under present climatic conditions and to project it to the Last Glacial Maximum (LGM). As also found in former studies with subsect. Phalolepis in Greece and Turkey using the same set of SSRs, genetic diversity for the segregated Italian species was higher than expected for narrow endemics with small populations. Genetic clusters, however, were not correlated with the described species and did not support the segregation of the purported narrow endemics from a widely defined C. deusta. The results of the ENM indicate that the Adriatic Sea was a migration corridor for C. deusta at the LGM.     

The microhabitat distribution of two Dactylogyrus species parasitizing the gills of the barbel, Barbus barbus

The microhabitat distribution of two congeneric species Dactylogyrus carpathicus and D. malleus (Monogenea) parasitizing the gills of the barbel (Barbus barbus L.) was investigated. We tested whether congeneric species exhibited microhabitat preference and whether interspecific interactions could be attributed to the microhabitat segregation of congeners. The outlying mean index method was used to evaluate species microhabitats. Gill variables (different microhabitats within gills) were used as environmental factors characterizing the gills. When abundances of both species were highest, and no significant difference was found between the abundance of the two species, the gill segments and gill areas were the most important factors segregating the Dactylogyrus species on the gills. Niche overlap was low within each of the four gill arches, and parasites were segregated in the same microhabitats within each gill arch. When abundances of both species were low, each monogenean species was segregated at the level of the gill arches. When abundances of both species increased, the niche and overlap between species increased. The distribution of both congeneric species confirmed microhabitat preference within the gills. The results suggest that microhabitat preference is dependent on species abundances, species being segregated in the case of low abundance, possibly to increasing mating opportunities. Both niche and overlap between species increased with species abundance. In the case of the high abundance of both species, microhabitat preference seems to be related to interspecific interactions between monogenean species, as previously found for endoparasitic species.     

Fine‐scale belowground species associations in temperate grassland

Evaluating how belowground processes contribute to plant community dynamics is hampered by limited information on the spatial structure of root communities at the scale that plants interact belowground. In this study, roots were mapped to the nearest one mm and molecularly identified by species on vertical (0–15 cm deep) surfaces of soil blocks excavated from dry and mesic grasslands in Yellowstone National Park (YNP) to examine the spatial relationships among species at the scale that roots interact. Our results indicated that average interspecific root – root distances for the majority of species were within a distance (3 mm) that roots have been shown to compete for resources. Most species placed their roots at random, although low root numbers for many species probably led to overestimating the occurrence of random patterns. According to theory, we expected that most of the remaining species would segregate their root systems to avoid competition. Instead we found that more species aggregated than segregated from others. Based on previous investigations, we hypothesize that species aggregate to increase uptake of water, nitrogen and/or phosphorus made available by neighbouring roots, or as a consequence of a reduction in the pathogenicity of soil biota growing in multispecies mixtures. Our results indicate that YNP grassland root communities are organized as closely interdigitating networks of species that potentially can support strong interactions among many species combinations. Future root research should address the prevalence and functional consequences of species aggregation across plant communities.