Use and partition of space and resources by two coexisting Rhyacophila species (Trichoptera) in a high mountain stream

Life history, distribution, drift and food habits of two coexisting Rhyacophila species, Rhyacophila evoluta and Rhyacophila intermedia were studied in a high mountain stream in the Pyrénées.The life histories of both species took two years downstream and three years upstream. Several generations and a large range of larval instars permanently coexisted in the river.Slight temporal segregation of the life histories and differences in size and food habits of the coexisting larvae partially reduced the interspecific competition and allowed an optimal use of space. Instars IV of R. evoluta and V of R. intermedia, which had the most similar biological characteristics had the same microdistribution when they lived alone but used different habitats when they coexisted.     

The development of radioresistance in irradiated drosophila nebulosa populations

The adaptive value of experimental populations of Drosophila nebulosa, irradiated each generation with doses of 500, 1000 and 2000 R of Co-60 gamma-rays, was measured with respect to their ability to compete with non-irradiated Drosophila willistoni. Two groups of experiments were developed: (a) to investigate the effects of repeated irradiation each generation on D. nebulosa under interspecific competition; (b) to demonstrate the radioresistance of D. nebulosa populations with different histories of irradiation. The results obtained through the interspecific competition were confirmed by tests of egg-laying capacity, hatchability and productivity of D. nebulosa from the competing populations. The populations with previous histories of gamma-irradiation were shown to be less susceptible than control populations to the detrimental effects of a new exposure to irradiation. This suggests that radioresistance developed in the D. nebulosa populations that had been irradiated over several generations.     

Influence of nymphal size, sex and morphotype on microdistribution of Deleatidium (Ephemeroptera: Leptophlebiidae) in a New Zealand river

• 1 The interaction of water depth and velocity with size, sex and morphotype of nymphs belonging to the leptophlebiid mayfly genus Deleatidium was investigated in a New Zealand river.
• 2 Velocity had a significant effect on distribution such that larger nymphs tended to be found more commonly in faster water. Depth also influenced size distribution, but in different ways at different sites.
• 3 Water velocity or depth alone did not significantly influence the distribution of sexes (1.0 mm head width), but the combined effect of these two factors was significant. Final instar males appeared to be relatively more abundant in slower water than final instar females.
• 4 Nymphs (1.0 mm head width) belonging to the informal lillii group (apically pointed gills) were present over a wide range of depth and velocity conditions, but those belonging to the myzobranchia group (round-tipped gills) were largely restricted to fast water (>0.9ms^?1).
• 5 Several mechanisms thought to cause variations in microdistribution are discussed. These include differential effects of oxygen availability, food requirements and drag forces, nymphal behaviour, and selection of oviposition sites by adults.

     

Diet and food selection in the endemic Hawaiian amphidromous goby, Sicyopterus stimpsoni (Pisces: Gobiidae)

Synopsis The ecological role of Sicyopterus stimpsoni as a nearly exclusive algal grazer in Hawaiian streams is established through gut content analysis of 192 fish from Wainiha River on the northern island of Kaua'i. Algae in three phyla (Chlorophyta, Cyanophyta, and Chrysophyta) were found to be the primary components of the fish's diet (94.62% of dry biomass). Aquatic insect immatures (primarily Chironomidae) accounted for most of the remaining food biomass (5.37%). Poorly developed gill rakers and high gut-to-length ratios provide evidence for adaptation to herbivory and morphological separation of S. stimpsoni from sympatric gobiods. Interspecific competition for the green alga, Cladophora sp., is suggested as an important feature of biotic interactions among native stream fishes but is mitigated by interspecific differences in food preference and utilization. A Category V hurricane which devastated the island in September 1992 provided a fortuitous opportunity to study disturbance influences on benthic food resources in the stream. Disturbance regimes influenced food selection of S. stimpsoni by altering the abundance and composition of stream algae. Rapid adjustment of S. stimpsoni to changes occurring in algal resource abundance illustrates its superb adaptation to life in a swift-water environment subjected to periodic flood-induced disturbance.     

Mayfly production in a New Zealand glacial stream and the potential effect of climate change

In contrast to the northern hemisphere where species of Chironomidae are usually the dominant benthic invertebrates in the coldest upper reaches of glacial streams, mayflies (Deleatidium spp.: Leptophlebiidae) predominate in equivalent conditions in New Zealand. We examined the life histories and annual production of Deleatidium spp. at two sites on the Matukituki River (South Island, New Zealand) and at three sites in its glacier-fed tributary, Rob Roy Stream. Mean annual water temperature at the five sites ranged from 2.1 to 7.0°C. Monthly sampling showed that mayfly populations were poorly synchronised at all sites but were probably univoltine. The large Deleatidium cornutum was the dominant mayfly species found at the upper sites (Sites 1 and 2) on Rob Roy Stream, whereas above the confluence with Matukituki River (Site 3) it co-existed with a complex of smaller species we refer to as D. “angustum”. Deleatidium “angustum” also dominated at the Matukituki sites. Deleatidium production calculated for the five sites, assuming an 11-month nymphal life, ranged from 0.48 g dry weight/m²/year (Site 1) to 3.07 g dry weight/m²/year (Site 3). The values for D. cornutum at Sites 2 and 3 are high for a species of Deleatidium and reflect its large size. This species appears to be strongly adapted for growth at low temperatures. Climate change scenarios for New Zealand predict the gradual and ultimate loss of small South Island glaciers and a consequent warming of streams as runoff from rainfall and snow melt becomes more dominant in spring. As a result, suitable habitats will be lost for cold-water specialists such as D. cornutum, and they are likely to suffer reductions in their distributional range and local extinction. In contrast, species such as those in the D. “angustum” complex may extend their ranges into streams formerly dominated by glacial meltwater. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users. Handling editor: S. Stendera     

The macroinvertebrate fauna of a New Zealand forest stream

Abstract

The distribution, abundance, and life histories of benthic invertebrates were investigated in a small, Nothofagus forest stream in North Canterbury, South Island, New Zealand. The fauna was dominated by Trichoptera and Plecoptera; Mollusca and fish were absent. Large particle detritivores and scrapers were the predominant functional groups found. Larval Philopotamidae (Trichoptera) were the only abundant filter feeders. Nymphs of the stonefly Spaniocerca zelandica and of mayflies, Deleatidium spp., were the most abundant animals on plant detritus; Deleatidium spp. were abundant on stones also. The distribution of invertebrates in riffles, loose stones, pools, and plunge pools was examined using mesh colonisation trays lifted in September, November, February, and May after respectively 88, 69, 98, and 94 days in situ. Most species were widely distributed, and sample densities of the more abundant insect species showed weak positive correlations with the biomass of detritus present in trays in most months. The turbellarian Neppia montana was overrepresented in trays compared with stream samples, but the relative abundance of most other species appeared to be similar to that on the stream bed. A large amount of silt accumulated in trays during a heavy flood, and was colonised mainly by the oligochaete Eiseniella tetraedra. Information on the life histories of 15 species of Plecoptera, Trichoptera, Ephemeroptera, and Coleoptera was obtained from a 1-year monthly sampling programme, in which 36 taxa were distinguished, and from collections of adult insects made over a 3–4-year period. Most species had poorly synchronised life histories, in that a wide size range was apparent among individuals in most months. Eight species emerged in at least 4 months; S. zelandica and the philopotamid Hydrobiosella stenocerca were taken in 8 and 9 months respectively.     

Seawater DMS in a perturbed coastal ecosystem

Intra- and interspecific niche overlap for two mayfly species with similar life cycle timing, Rhithrogena semicolorata and Ecdyonurus sp. gr. venosus, were investigated. The nymphs were classified into 5 classes according to size and spatial overlaps are measured along a substratum roughness gradient. Substratum roughness selection was investigated by defining utilisation curves, optimum and tolerance values of the nymphs in relation to larval growth. Differences between species and size classes within each species were observed. Ecdyonurus sp. gr. venosus dominated on rough substrates, whereas R. semicolorata was most abundant on smooth substrates. An intermediate value of total interspecific substratum roughness overlap (0.49) was found. Higher intraspecific than interspecific overlap values suggested a spatial niche segregation between the two species. The results suggested that the spatial niches measured, were closer to the `fundamental niches' than could be expected if competition was acting on the two studied populations.     

Coexistence of three differentDrosophila species by rescheduling their life history traits in a natural population

We present evidence for coexistence of three differentDrosophila species by rescheduling their life history traits in a natural population using the same resource, at the same time and same place.D. ananassae has faster larval development time (DT) and faster DT(egg-fly) than other two species thus utilizing the resources at maximum at both larval and adult stages respectively. Therefore,D. ananassae skips the interspecific competition at preadult stage but suffers more from intraspecific competition. However,D. melanogaster andD. biarmipes have rescheduled their various life history traits to avoid interspecific competition. Differences of ranks tests for various life history traits suggest that except for DT(egg-pupa), the difference of ranks is highest for the combination ofD. melanogaster andD. ananassae for all other life history traits. This difference is maintained by tradeoffs between larval development time and pupal period and between pupal period and DT(egg-pupa) inD. ananassae.     

Temporal resource partitioning mitigates interspecific competition and promotes coexistence among insect parasites

A key to understanding life's great diversity is discerning how competing organisms divide limiting resources to coexist in diverse communities. While temporal resource partitioning has long been hypothesized to reduce the negative effects of interspecific competition, empirical evidence suggests that time may not often be an axis along which animal species routinely subdivide resources. Here, we present evidence to the contrary in the world's most biodiverse group of animals: insect parasites (parasitoids). Specifically, we conducted a meta-analysis of 64 studies from 41 publications to determine if temporal resource partitioning via variation in the timing of a key life-history trait, egg deposition (oviposition), mitigates interspecific competition between species pairs sharing the same insect host. When competing species were manipulated to oviposit at (or near) the same time in or on a single host in the laboratory, competition was common, and one species was typically inherently superior (i.e. survived to adulthood a greater proportion of the time). In most cases, however, the inferior competitor could gain a survivorship advantage by ovipositing earlier (or in a smaller number of cases later) into shared hosts. Moreover, this positive (or in a few cases negative) priority advantage gained by the inferior competitor increased as the interval between oviposition times became greater. The results from manipulative experiments were also correlated with patterns of life-history timing and demography in nature: the more inherently competitively inferior a species was in the laboratory, the greater the interval between oviposition times of taxa in co-occurring populations. Additionally, the larger the interval between oviposition times of competing taxa, the more abundant the inferior species was in populations where competitors were known to coexist. Overall, our findings suggest that temporal resource partitioning via variation in oviposition timing may help to facilitate species coexistence and structures diverse insect communities by altering demographic measures of species success. We argue that the lack of evidence for a more prominent role of temporal resource partitioning in promoting species coexistence may reflect taxonomic differences, with a bias towards larger-sized animals. For smaller species like parasitic insects that are specialized to attack one or a group of closely related hosts, have short adult lifespans and discrete generation times, compete directly for limited resources in small, closed arenas and have life histories constrained by host phenology, temporal resource subdivision via variation in life history may play a critical role in allowing species to coexist by alleviating the negative effects of interspecific competition.     

An analysis of changes in the invertebrate community along a southern New Zealand montane stream

Changes in benthic community structure at four sites along a flood-prone montane stream continuum in the South Island of New Zealand were analysed using rank correlation (Kendall's Tau and Spearman's r_s) and a community similarity index (S_c) developed initially for stream pollution assessment. Species richness was highest in a forested headwater stream, which is attributed to its relatively greater physical stability and habitat heterogeneity. Community similarity decreased with distance apart, and pairs of open and forested sites showed the greatest similarity. Species composition and relative abundance along the continuum changed least in the numerically dominant Ephemeroptera and Plecoptera, groups in which several species had broad habitat requirements. In contrast, a number of dipteran and trichopteran species had more restricted distributions. The more specialised niche requirements of some species in these orders are attributed to their generally small size, their ability to spin silk, and their holometabolous development. Physical rather than biological factors appeared to limit the distributions of most invertebrates, and there was little evidence of ecological segregation by either food resource partitioning or differences in life history patterns amongst many closely related species. Theories that stream communities are well ordered entities maximising the efficiency of energy utilisation cannot realistically be applied to such unstable stream systems.     

Diversity and endemism in deglaciated areas: ploidy,relative genome size and niche differentiation in the Galium pusillum complex (Rubiaceae) in Northern and Central Europe

Background and Aims

Plants endemic to areas covered by ice sheets during the last glaciation represent paradigmatic examples of rapid speciation in changing environments, yet very few systems outside the harsh arctic zone have been comprehensively investigated so far. The Galium pusillum aggregate (Rubiaceae) is a challenging species complex that exhibits a marked differentiation in boreal parts of Northern Europe. As a first step towards understanding its evolutionary history in deglaciated regions, this study assesses cytological variation and ecological preferences of the northern endemics and compares the results with corresponding data for species occurring in neighbouring unglaciated parts of Central and Western Europe.

Methods

DNA flow cytometry was used together with confirmatory chromosome counts to determine ploidy levels and relative genome sizes in 1158 individuals from 181 populations. A formalized analysis of habitat preferences was applied to explore niche differentiation among species and ploidy levels.

Key Results

The G. pusillum complex evolved at diploid and tetraploid levels in Northern Europe, in contrast to the high-polyploid evolution of most other northern endemics. A high level of eco-geographic segregation was observed between different species (particularly along gradients of soil pH and competition) which is unusual for plants in deglaciated areas and most probably contributes to maintaining species integrity. Relative monoploid DNA contents of the species from previously glaciated regions were significantly lower than those of their counterparts from mostly unglaciated Central Europe, suggesting independent evolutionary histories.

Conclusions

The aggregate of G. pusillum in Northern Europe represents an exceptional case with a geographically vicariant and ecologically distinct diploid/tetraploid species endemic to formerly glaciated areas. The high level of interspecific differentiation substantially widens our perception of the evolutionary dynamics and speciation rates in the dramatically changing environments of Northern Europe.     

Segregation in space and time explains the coexistence of two sympatric sub‐Antarctic petrels

Biological communities are shaped by competition between and within species. Competition is often reduced by inter‐ and intraspecific specialization on resources, such as differencet foraging areas or time, allowing similar species to coexist and potentially contributing to reproductive isolation. Here, we examine the simultaneous role of temporal and spatial foraging segregation within and between two sympatric sister species of seabirds, Northern Macronectes halli and Southern Macronectes giganteus Giant Petrels. These species show marked sexual size dimorphism and allochrony (with earlier breeding by Northern Giant Petrels) but this is the first study to test for differences in foraging behaviours and areas across the entire breeding season both between the two species and between the sexes. We tracked males and females of both species in all breeding stages at Bird Island, South Georgia, to test how foraging distribution, behaviour and habitat use vary between and within species in biological time (incubation, brood‐guard or post‐brood stages) and in absolute time (calendar date). Within each breeding stage, both species took trips of comparable duration to similar areas, but due to breeding allochrony they segregated temporally. Northern Giant Petrels had a somewhat smaller foraging range than Southern Giant Petrels, reflecting their greater exploitation of local carrion and probably contributing to their recent higher population growth. Within species, segregation was spatial, with females generally taking longer, more pelagic trips than males, although both sexes of both species showed unexpectedly plastic foraging behaviour. There was little evidence of interspecific differences in habitat use. Thus, in giant petrels, temporal segregation reduces interspecific competition and sexual segregation reduces intraspecific competition. These results demonstrate how both specialization and dynamic changes in foraging strategies at different scales underpin resource division within a community.     

Multiple-stressor effects on stream invertebrates: DNA barcoding reveals contrasting responses of cryptic mayfly species

Most freshwater ecosystems are subject to multiple anthropogenic stressors, which commonly reduce biodiversity across all levels. Existing freshwater bioassessment programmes aim at identifying responses of aquatic biota to stressors. For practical reasons, higher-level taxonomic groups (e.g. genus or family) are often used in these programmes. This approach, however, may bias assessment results as different species can differ substantially in their biological traits, thus emphasising the need for species-level data. DNA barcoding can reliably generate species-level data for animals by sequencing a fragment of the mitochondrial cytochrome c oxidase subunit I gene (COI). This allows investigating species-specific responses to environmental stressors. In this study, we sampled 43 stream sites in southern New Zealand spanning wide gradients of agricultural stressors (fine sediment and nutrient levels). We first used conventional morphological assessment to determine stream invertebrate responses to the stressors, focusing on two important indicator taxa, the mayfly Deleatidium and the snail Potamopyrgus. We then tested for the presence of cryptic species in Deleatidium and Potamopyrgus using DNA barcoding of the COI gene for 520 and 305 specimens, respectively. While all Potamopyrgus specimens belonged to a single species, Deleatidium consisted of 12 distinct molecularly identified clades that likely represent distinct species. Finally, we compared stressor responses assessed at genus and species level. While overall Deleatidium abundance was unrelated to stressor levels, some of the individual clades differed clearly in the magnitude and direction of their responses to nutrient and sediment stress. While the most abundant cryptic Deleatidium clade (clade 1) showed no relationship to sediment or nutrient levels, clades 2 and 3 responded negatively to nutrient or sediment increases, respectively. These contrasting patterns indicate that individual freshwater invertebrate species, often merged to a higher taxonomic level for biomonitoring purposes, can differ substantially in their tolerance to stressors and respond in more complex ways than observed at genus level. Overall, our results highlight the considerable potential and importance of including DNA barcoding into freshwater ecosystem assessment and biomonitoring programmes.     

Feeding competition between two goby species,Pomatoschistus lozanoi (de Buen) and P. minutus (Pallas), in a macrotidal saltmarsh

Two transient goby species (Pomatoschistus minutus and P. lozanoi) occurred in saltmarshes of the macrotidal Mont-Saint-Michel bay. They colonised the tidal creeks during each spring tide throughout the study conducted in 1997. P. minutus swallowed a least 4 % of its body weight. Its diet was dominated by the most abundant resident amphipod of the saltmarshes, Orchestia gammarellus. P. lozanoi ingested less food during their stay in the creek (2.4 % of its body weight). O. gammarallus also occurred in its diet, but mysids were the main food items both during flood and ebb. In addition to this trophic segregation, a temporal segregation was observed. In fact, P. minutus stayed longer than P. lozanoi in the saltmarsh; it colonised creeks first and returned to coastal waters the last. These two predatory and sympatric species have a different trophic behaviour that limits interspecific competition both from the trophic and temporal point of view.     

Dung exploitation by the dung beetle Typhaeus typhoeus (Col., Geotrupidae)

Summary The competitive interactions between roach (Rutilus rutilus) and rudd (Scardinius erythrophthalmus) were investigated in two habitats, the open water and the waterlily zone. The growth rates of both species in enclosures were lower when confined together than when alone, demonstrating interspecific competition. Allopatric roach had the highest growth rate in both habitats although the open water was preferred. The diets of both species were dominated by zooplankton in both the open water and waterlily zones. In laboratory experiments, roach had significantly higher feeding rates than rudd when fed D. magna and Cyclops sp. The impact of roach on the mean sizes and densities of zooplankton in the enclosures, together with the results from the laboratory study, indicate that roach were competitively superior in the open water. Because of high mortality in the sympatric waterlily enclosure, no conclusions about interspecific competition in this habitat could be drawn. The observed habitat segregation between roach and rudd was at least partly interactive.     

Interspecific competition among urban cockroach species

The aim of this study was to quantify and to compare the effects of intraspecific and interspecific competition among pairs of urban cockroaches [Blatta orientalis L., Periplaneta americana (L.), and Periplaneta australasiae Fabricius (Dictyoptera: Blattodea)] in relation to the limitation of resources such as shelter or food. Our approach was to assess whether the presence of one species affected the resource exploitation of another. A reduction in access to shelters or to food for one species revealed dominant/subordination relationships among species and induced spatial segregation in shelters and temporal segregation during food exploitation. The fragmentation of available resources facilitated spatial segregation and the access of more individuals of the subordinate species. Individuals of each species were aware of the presence of non‐conspecifics. The outcome of interspecific competition under laboratory conditions should help us to understand how segregated spatial distributions occur under natural conditions in urban areas.     

Food of the New Zealand trichopterans Hydrobiosis parumbripennis McFarlane and Hydropsyche colonica McLachlan

The food of the larvae of two New Zealand Trichoptera, Hydrobiosis parumbripennis and Hydropsyche colonica, was investigated between March and November 1971 at the Wainui Valley Stream, Canterbury, New Zealand. Most samples were collected from an experimental channel constructed in the stream in 1970. The first three instars of H. parumbripennis were mainly detrital feeders, whereas the two later instars were exclusively carnivorous. In contrast, H. colonica was omnivorous. The main prey taken by H. parumbripennis were larvae of a simuliid, Austrosimulium tillyardianum, Chironomidae (subfamily Orthocladiinae), and an ephemeropteran, Deleatidium sp. The first three instars appeared to select chironomids in preference to simuliids, whereas the converse was true for the final two instars. The main prey taken by H. colonica was A. tillyardianum. The size of prey and the mean number of prey found per larva of H. parumbripennis increased with each instar. Forage ratios indicated that H. parumbripennis were eating A. tillyardianum at the same relative frequency as their abundance in the fauna, but that the chironomids and Deleatidium sp. were being preferentially selected as prey. Cased caddis larvae were avoided as food items, as was the mollusc Potamopyrgus antipodarum. Although H. parumbripennis was the most important predator of A. tillyardianum in the stream, it is considered that it has little effect on the simuliid population.     

Evolutionary and demographic history among Maghrebian Medicago species (Fabaceae) based on the nucleotide sequences of the chloroplast DNA barcode trnH-psbA

Data from trnH-psbA intergenic spacer (cpDNA) were analyzed to elucidate molecular evolution within and among Maghrebian species of Medicago. The spacer highlighted a high interspecific variation and a low intraspecific diversity among species. Haplotype and nucleotide diversities revealed high level of variation. Parsimony and median-joining Network methods revealed (1) the segregation into 17 haplotypes; (2) the ancestral behaviour of the annual Medicago minima and (3) the clusters are independent of the geographic origin. The neutral evolution of Wright and Fisher is rejected since the Tajima's D values deviated from 0. Besides, the statistical analyses are in agreement with an evolution into stable populations' size.     

Space use and temporal partitioning of sympatric Tasmanian devils and spotted‐tailed quolls

Sympatric species can minimise interspecific competition by spatial avoidance or by altering their temporal activity to reduce encounter rates. The Tasmanian devil (Sarcophilus harrisii), the largest carnivorous marsupial, coexists with the smaller spotted‐tailed quoll (Dasyurus maculatus) in Tasmania, Australia. Quolls may be susceptible to interspecific competition from devils, because they utilise similar habitats, consume similar prey species and are displaced by devils at food sources. Such competition might cause quolls to spatially or temporally avoid devils. To investigate whether spatial or temporal avoidance occurred, we deployed GPS collars on sympatric devils and quolls and conducted a camera survey at a site in northwest Tasmania where the devil population was not affected by devil facial tumour disease. GPS tracking coincided with the lactation period when devils and quolls had young in dens and continued until weaning occurred. We found little spatial segregation of home range and core area placement between devils and quolls and among devils. Quolls showed more spatial segregation within the sexes than between them. Devils had larger home ranges than quolls. Male devils had larger home ranges than females, but there was no difference in home range size between the sexes of quolls. Females of both species travelled significantly further per night than did males. There was moderate temporal partitioning between the two species: devil activity peaked after dusk and devils remained active until the early morning, while quoll activity showed distinct peaks around dusk and dawn. In conclusion, quolls did not spatially avoid devils but moderate temporal partitioning occurred. It is plausible that quolls are active at different times of the diel cycle to reduce encountering devils, but further studies are needed to resolve the cause of this temporal partitioning.     

Habitat stability and the larval mosquito community in treeholes and other containers on a temperate Island

The temporal fluctuation of water levels and the presence of mosquito larvae were investigated for four types of small container habitats (treeholes, bamboo stumps, cemetery stone vases, and cemetery stone vessels) on Kabeshima Island in Kyushu, southwestern Japan. The probability that containers held water was positively correlated with the quantity of the preceding rainfall and with the depth and volume of the containers. It was estimated that dehydration occurred more regularly in autumn and winter than in summer. The probability that mosquito larvae were present in each type of container was positively correlated with habitat stability in terms of the probability of the existence of standing water and the coefficient of variation of the water level. Twelve species of mosquito larvae, including two rare predators, were found. Species composition differed between the different types of container. Although 4 to 10 species used each type of container, the median number of species per container was two for treeholes and bamboo stumps, and one for the others. The dominant species wasTripteroides bambusa in treeholes and bamboo stumps,Aedes albopictus in stone vases, andA. japonicus in stone vessels. The larval mosquito community, which lacked major predators, possessed the following features that may facilitate the coexistence of many species: (1) niche segregation amongst species in terms of their selection of container types; (2) an aggregated distribution of the individual species among containers of the same type; (3) high intraspecific mean crowding (and hence probably intense intraspecific competition) in the dominant species in each type of container; (4) independent species associations within the same type of container; and (5) low interspecific mean crowding (and hence probablynot intense interspecific competition) between species in the same type of container.