Does size matter for dispersal distance?

Aim The aim of this study is to answer the questions: (1) do small organisms disperse farther than large, or vice versa; and (2) does the observed pattern differ for passive and active dispersers? These questions are central to several themes in biogeography (including microbial biogeography), macroecology, metacommunity ecology and conservation biology. Location The meta‐analysis was conducted using published data collected worldwide. Methods We collected and analysed 795 data values in the peer‐reviewed literature for direct observations of both maximal dispersal distance and mass of the dispersing organisms (e.g. seeds, not trees). Analysed taxa ranged in size from bacteria to whales. We applied macroecology analyses based on null models (using Monte Carlo randomizations) to test patterns relative to specific hypotheses. Results Collected dispersal distance and mass data spanned 9 and 21 orders of magnitude, respectively. Active dispersers dispersed significantly farther (P < 0.001) and were significantly greater in mass (P < 0.001) than passive dispersers. Overall, size matters: larger active dispersers attained greater maximum observed dispersal distances than smaller active dispersers. In contrast, passive‐disperser distances were random with respect to propagule mass, but not uniformly random, in part due to sparse data available for tiny propagules. Conclusions Size is important to maximal dispersal distance for active dispersers, but not for passive dispersers. Claims that microbes disperse widely cannot be tested by current data based on direct observations of dispersal: indirect approaches will need to be applied. Distance–mass relationships should contribute to a resolution of neutral and niche‐based metacommunity theories by helping scale expectations for dispersal limitation. Also, distance–mass relationships should inform analyses of latitudinal species richness and conservation biology topics such as fragmentation, umbrella species and taxonomic homogenization.     

Environmental heterogeneity,dispersal mode,and co‐occurrence in stream macroinvertebrates

Both environmental heterogeneity and mode of dispersal may affect species co‐occurrence in metacommunities. Aquatic invertebrates were sampled in 20–30 streams in each of three drainage basins, differing considerably in environmental heterogeneity. Each drainage basin was further divided into two equally sized sets of sites, again differing profoundly in environmental heterogeneity. Benthic invertebrate data were divided into three groups of taxa based on overland dispersal modes: passive dispersers with aquatic adults, passive dispersers with terrestrial winged adults, and active dispersers with terrestrial winged adults. The co‐occurrence of taxa in each dispersal mode group, drainage basin, and heterogeneity site subset was measured using the C‐score and its standardized effect size. The probability of finding high levels of species segregation tended to increase with environmental heterogeneity across the drainage basins. These patterns were, however, contingent on both dispersal mode and drainage basin. It thus appears that environmental heterogeneity and dispersal mode interact in affecting co‐occurrence in metacommunities, with passive dispersers with aquatic adults showing random patterns irrespective of environmental heterogeneity, and active dispersers with terrestrial winged adults showing increasing segregation with increasing environmental heterogeneity.     

Empty habitat in coastal sediments for populations of macrozoobenthos

Species with wide dispersal are expected to have little empty habitat. This was tested by analysing habitat use in the Wadden Sea near the island of Sylt (North Sea). Sampling covered 222 intertidal sites spread across the K?nigshafen tidal flats (4.5 km², mapping approach conducted once) and 270 subtidal sites along 12 km of a tidal channel system (stratified random sampling in spring and autumn). For any single species, 'suitable habitat' was extrapolated from the ranges of water depth and sediment composition present at the ten sites with the highest frequency or abundance of the species. On average, macrobenthic species actually used less than half of the suitable sites within the scale of local populations; this was far less than expected from a lognormal distribution. In the subtidal, abundance of most species changed between the two sampling seasons. A 50% increase in overall abundance was accompanied by a decrease in empty habitat of only about 25%. Thus, a doubling of abundances would not fill the empty space but just of half of it. The polychaete Scoloplos armiger was an exception in occupying almost all of its suitable habitat in the intertidal and subtidal sediments. A distinct patch of high species richness occurred where flood waters persistently form a large gyre which may enhance larval settlement. We suggest that limitations to larval settlement and/or juvenile survival primarily cause the extent of the observed habitat emptiness. Electronic Publication     

Effects of oil/gas field produced water on the macrobenthic community in a small gradient estuary

Little is known about how climatic events (i.e., drought) inhibit or enhance movement of petroleum hydrocarbon laden sediments in estuarine systems and how this in turn effects the macrobenthic populations exposed to these sediments. Seventeen collection stations were established and monitored at New Bayou, Texas, a small gradient estuary which receives petroleum products via oil/gas field produced water discharge. Hydrographic, geologic and biotic samples were taken monthly from each station during a period of reduced rainfall. Both temperature and dissolved oxygen data taken during the study followed patterns typical for the area. Salinity values increased into the polyhaline range at near-bay stations during the reduced rain period, while upper bayou stations remained within the oligohaline range. Sediment grain sizes along the bayou were generally in the silty-clay range. Sediment hydrocarbon concentrations were highest (11.4 mg g⁻¹ dry sediment) around the produced water discharge site and lowest at near-bay stations (0.2 mg g⁻¹ dry sediment). Ninety-six macrobenthic species were collected during the study. General effects from produced water discharge on macrobenthic populations followed the classical pattern outlined by other research in shallow bay systems. A sediment hydrocarbon concentration of 2.5 mg g⁻¹ dry sediment was found to reflect the average value needed to depress population abundances. Both abundance and diversity values were lowest at central stations near the discharge site. This zone of depressed macrobenthic populations extended from the discharge site at least 107 m downstream and 46 m upstream. The zone of stimulated macrobenthic populations occurred 1486 m downstream and 381 m upstream from the discharge site.     

Effect of the Razdol’naya River on structure of the free-living nematode community of the Amursky Bay,Sea of Japan

The effect of the Razdol’naya River on the littoral and sublittoral nematode community was studied in the innermost part of the Amursky Bay. Thanks to the seasonal abundance of juveniles of macrobenthic animals, pseudomacrobenthic species prevailed in the meiobenthos littoral community. Juvenile amphipods dominated (39.5%), the percentage of nematodes was 18.4% (45 species). The nematodes dominated in the meiobenthos community (65.7%, 71 species) at the sublittoral zone. The similarity of the species structure of littoral and sublittoral nematodes reached 36.5%. Low population density (2.6±1.7 thousand ind/m²) and poor species diversity of nematodes (7 species), the dominating form being Oncholaimium paraolium, were recorded at the littoral zone, which was subjected to direct riverine influence (1‰salinity). The correlation analysis revealed no relationship between salinity and population density of nematodes at the littoral zone. At the sublittoral zone (at stations of salinity less than 2.85‰) the parameters of population density (73.9±32.3 thousand ind/m²) and species diversity (12 species) of nematodes were the lowest, Parodontophora timmica dominated. A poor expressed dependence between the density of nematode populations and salinity was revealed. The correlation between the average size of ground particles and population density of nematodes was recorded. The greatest similarity of the species structure of littoral and sublittoral nematodes (26.67%) was recorded on gravel-sand bottoms.     

Behaviour controls post-settlement dispersal by the juvenile bivalves Austrovenus stutchburyi and Macomona liliana

Post-settlement dispersal is a key process affecting the population dynamics of many soft sediment benthic invertebrates. Despite its importance, few studies have quantified those factors that influence juvenile dispersal. In a laboratory flume, we examined the effects of three flow velocities and two substrate types on the dispersal of two common intertidal bivalves: the deposit-feeding wedge shell Macomona liliana and the suspension-feeding cockle Austrovenus stutchburyi. Juveniles of three size classes (<2, 2-4 and 4-8 mm shell length) were added to cores of defaunated natural sediment or glass beads. We recorded the number of bivalves remaining in cores versus those recovered downstream either on the flume floor, in bedload traps or in a plankton net at the end of the working section of the flume after 48 h at three freestream velocities (4.8, 11.0 and 16.6 cm s⁻¹). At flow speeds of 4.8 cm s⁻¹, <5% of individuals were recovered outside the cores for both species. At higher flows, the dispersal mode (crawling, bedload or in suspension) and frequency of dispersal differed markedly between species. Austrovenus dispersed primarily by crawling in the low flow treatments. The frequency of dispersal increased substantially (2-6×) between 11.0 and 16.6 cm s⁻¹, and most Austrovenus were found in bedload traps at the highest flow. At the highest flow, twice as many Austrovenus individuals left the glass bead treatment as from the natural sediment. The number of dispersing Macomona also increased with increasing flow from the natural sediment, but numbers dispersing from glass beads were similar for the two higher flows (11.0 and 16.6 cm s⁻¹). Macomona dispersal mode was associated with size; smaller size classes were collected in the bedload traps and the plankton net in approximately equal proportions, while only a small proportion of the largest size class were collected in the plankton net. In contrast to flume dispersal experiments with live bivalves, most dead Austrovenus did not move at the highest flow speed, while most dead Macomona were transported at the highest flow speed. The live-dead comparisons, in conjunction with our primary experimental results, imply that there are active behavioural components to both water column and bedload transport. Our research emphasises that both species actively influence post-settlement transport in response to both substrate type and flow regime, and that bedload transport, often categorised as a passive transport process, is also greatly influenced by active behavioural processes.     

Features of the Species and Spatial Structure of Macrobenthos in the Green Cape Lagoon (Kandalaksha Bay,White Sea)

Biology Bulletin - Abstract—Features of the species and spatial structure of sublittoral macrobenthic communities in a lagoon ecosystem of Green Cape (White Sea) have been studied. Thirty-one...     

Metacommunity structuring in stream networks: roles of dispersal mode,distance type,and regional environmental context

Within a metacommunity, both environmental and spatial processes regulate variation in local community structure. The strength of these processes may vary depending on species traits (e.g., dispersal mode) or the characteristics of the regions studied (e.g., spatial extent, environmental heterogeneity). We studied the metacommunity structuring of three groups of stream macroinvertebrates differing in their overland dispersal mode (passive dispersers with aquatic adults; passive dispersers with terrestrial adults; active dispersers with terrestrial adults). We predicted that environmental structuring should be more important for active dispersers, because of their better ability to track environmental variability, and that spatial structuring should be more important for species with aquatic adults, because of stronger dispersal limitation. We sampled a total of 70 stream riffle sites in three drainage basins. Environmental heterogeneity was unrelated to spatial extent among our study regions, allowing us to examine the effects of these two factors on metacommunity structuring. We used partial redundancy analysis and Moran's eigenvector maps based on overland and watercourse distances to study the relative importance of environmental control and spatial structuring. We found that, compared with environmental control, spatial structuring was generally negligible, and it did not vary according to our predictions. In general, active dispersers with terrestrial adults showed stronger environmental control than the two passively dispersing groups, suggesting that the species dispersing actively are better able to track environmental variability. There were no clear differences in the results based on watercourse and overland distances. The variability in metacommunity structuring among basins was not related to the differences in the environmental heterogeneity and spatial extent. Our study emphasized that (1) environmental control is prevailing in stream metacommunities, (2) dispersal mode may have an important effect on metacommunity structuring, and (3) some factors other than spatial extent or environmental heterogeneity contributed to the differences among the basins.     

Spatial and environmental effects on a rock‐pool metacommunity depend on landscape setting and dispersal mode

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Are distribution patterns linked to dispersal mechanism? An investigation using pond invertebrate assemblages

1. Dispersal ability influences the distribution and abundance of organisms, but empirical investigations of the relationship between dispersal ability and the composition of ecological assemblages are scarce. Here, we compare between-site variation in the species richness and community composition of actively and passively dispersing pond invertebrates.
2. Coleoptera (active dispersers) and microcrustacea (passive dispersers) were sampled over a season from 16 ponds within a 4-km radius in south-west England. Species richness and community composition were related to environmental variables using regression and Canonical Correspondence Analysis (CCA), respectively.
3. Coleopteran species richness was significantly and positively correlated with pond permanence and maximum area, whereas microcrustacean species richness was relatively equal across sites and did not correlate with environmental variables. The frequency of species' occurrence between sites was the same for both groups, which suggests that active and passive dispersers exhibited the same degree of dispersal.
4. Between-site variation in community composition was non-random for both groups, with pond permanence and area, together, explaining similar proportions of between-site variation for Coleoptera. Permanence was correlated most strongly with microcrustacean community composition and a high proportion (25%) of microcrustacean species were more numerous in smaller, more ephemeral ponds.
5. These data suggest that, at small spatial scales, Coleoptera which can undertake multiple dispersal events, are more likely to colonise large, more permanent ponds than passively dispersing microcrustacea. For microcrustacea, other traits (in this case those permitting existence in ephemeral habitats) may over-ride the influences of dispersal in driving between-site variation in species composition.     

Seed size and dispersal potential of Acer rubrum (Aceraceae) samaras produced by populations in early and late successional environments

In order to determine if red maple dispersal potential or seed size change during secondary succession, samaras were collected from five populations located in early successional environments and five populations located in late successional environments. Wing loading ratios (samara mass—mg/samara area—cm²), which are inversely proportional to dispersal ability, were computed for all samaras, and seeds were excised from each samara and weighed. Samaras from the early successional red maples showed slightly but significantly lower wing loading ratios than those from the late successional environments. This result corresponds with the conclusions reached by several theoretical investigations of seed dispersal evolution that predict that recently founded populations will show greater dispersal abilities than more established populations. The earlier successional populations had slightly heavier seeds than the later successional populations, which suggests that the changes in community composition and dynamics that occur during this successional sequence do not select for heavier seeds in older red maple populations. Coefficients of variation for wing loading and seed size showed no consistent trends with successional stage, which indicates that variation in these characters does not decrease as succession proceeds.     

Changes in the functional feeding groups of macrobenthic fauna during mangrove forest succession in Zhanjiang,China

Understanding the relationships between mangrove forest succession and the functional diversity of mangrove fauna could facilitate the restoration of mangrove ecosystems, which have been severely damaged in recent decades. The current report describes changes in macrobenthic functional diversity in a mangrove chronosequence that included a primary community (unvegetated shoal), an early community (Avicennia marina), a middle community (Aegiceras corniculatum), and a late community (Bruguiera gymnorrhiza?+?Rhizophora stylosa) in Zhanjiang, China. Phytophages were the dominant macrobenthic functional feeding group regardless of mangrove succession stage, sampling season, or macrobenthic faunal parameter (species richness, abundance, and biomass). As mangrove succession progressed, the proportions of macrobenthic species richness, abundance, or biomass represented by omnivores significantly increased (except for biomass and in the late stage; ranged from 0.065 to 0.230 and 0.033 to 0.368, respectively in wet season, and 0.000 to 0.192 and 0.000 to 0.396, respectively in dry season), while the proportions significantly decreased for detritivores during the dry season (ranged from 0.156 to 0.056, 0.107 to 0.019, and 0.066 to 0.005, respectively). Non-metric multi-dimensional scaling and PERMANOVA also indicated that the structure of macrobenthic faunal functional feeding groups was significantly affected by mangrove succession. Further analyses indicated that the changes in the relative dominance among macrobenthic faunal functional feeding groups during mangrove succession were mainly associated with changes in plant density, coverage/canopy density, and total nitrogen content of sediment, i.e., they were mostly associated with changes in food sources. The results increase our understanding of the relationship between benthic functional diversity and mangrove succession and could help guide mangrove restoration in China and around the world.     

Factors influencing macroinvertebrate assemblages in artificial subtropical ponds of Taiwan

Macroinvertebrate assemblages and its association with environmental factors at the 11 artificial subtropical ponds of Taiwan were examined using the multivariate analysis software STATICO. The aims of the study were to determine whether spatial and seasonal variation of macroinvertebrate assemblages changed seasonally, to examine which environmental factors determined the spatial and temporal structure of maroinvertebrate assemblages, and to compare between-pond variations in the taxon composition of macroinvertebrates. Macroinvertebrates were collected seasonally by a corer and a sweep net in 2007, and 13 physical and chemical factors were measured at the same time. A total of 31 macroinvertebrate taxa were collected during the sampling period, and the most dominant taxa were Chironomidae (31.7% of total animal abundance) and Tubificidae (22.4%). STATICO identified pond size, pond depth, sediment depth, and altitude as the major abiotic factors and Bufo melanostictus (Amphibia) as the major biotic factor to influence macroinvertebrate assemblages at these ponds. These factors changed with seasonality. For example, the abundance of B. melanostictus was the most important factor during the spring but became much less important in other seasons. According to the spatial distribution patterns of macroinvertebrate assemblages, macroinvertebrates could be split into two groups based on their dispersal. The active dispersers, such as insect taxa, were strongly associated with pond size and the passive dispersers, such as non-insect taxa, were strongly associated with the pond depth and/or sediment depth. The results of this study suggested that pond size might influence macroinvertebrate assemblages through their dispersal mechanisms and that the environmental factors which influenced the macroinvertebrate assemblages most changed with seasons in this study area.     

Distribution,substratum preference and burrowing behaviour of Lovenia elongata (Gray) (Echinoidea: Spatangoida) in the Gulf of Elat ('Aqaba), Red Sea

Lovenia elongata (Gray), the Indo-West Pacific spatangoid, was studied in the Gulf of Elat ('Aqaba) at the northern end of the Red Sea. The occurrence and local distribution of this macrobenthic burrowing species was determined and related to the particle size distribution of the substrata in which it was found. Its habitat preference is for clean, grassless, sandy bottoms in intertidal areas (beaches and sandbars), and on sublittoral level bottoms of protected bays and lagoons sheltered from heavy wave action. The preferred sediments were found to range from medium to fine sand. The population density, size frequency distribution, growth, reproduction, recruitment, and mortality of a sublittoral population of L. elongata (at Wadi Taba, Sinai) were studied.The mode of life of this burrowing heart urchin was observed in situ and in aquaria. Its burrowing and emergence behaviour are described. Experiments were carried out on the effect of particle size on burrowing behaviour. There are significant differences in the ability of L. elongata to manipulate substrata of varying grain size. The burrowing process was found to be most rapid in natural sand of medium-fine composition, corresponding to the normal sediment of the urchin's habitat. Such sediments were also found to be suitable for successful larval settlement; juveniles did not survive in coarser sand.Differences in population density and spatial distribution between various size groups are attributed to differences in their ability to manipulate the substratum. L. elongata is found to be morphologically suited to sand, particularly medium-fine sand; however, it is restricted by its morphology to shallow burrowing, since it lacks specialized mechanisms for deeper burrowing and funnel building.     

互花米草入侵对大型底栖动物群落垂直结构的影响

2012年8月,在浙江省玉环县漩门湾国家湿地公园,利用自制分层采样器,以5 cm为单位对0—25 cm泥层进行分层取样,研究潮间带互花米草滩涂和自然滩涂两种生境中大型底栖动物的垂直分布情况,以及互花米草入侵对大型底栖动物垂直结构的影响。两种生境中共获得大型底栖动物40种,其中软体动物18种,甲壳动物13种,环节动物4种,其他类群5种。在自然滩涂生境中获得的物种数为30种,互花米草生境为31种,两种生境的共有种有21种。互花米草和自然滩涂生境平均栖息密度分别为222.6株/m2和1052.8株/m2。研究结果表明:(1)大型底栖动物主要栖息在0—10 cm泥层深度;(2)大型底栖动物群落沿泥层深度存在分层分布现象;(3)互花米草入侵缩短了底栖动物的垂直分布距离,但对其主要栖息深度影响不大;(4)互花米草入侵改变了原来生境特征,导致大型底栖动物的群落结构发生变化。     

Seed Dispersal by Birds and Bats in Lowland Philippine Forest Successional Area

In the tropical forests of SE Asia, only a few studies have dealt with the role animal dispersal plays in early forest succession and rehabilitation, and a comparison of bird and bat dispersal is even rarer. We investigated seed dispersal by birds and bats in a successional area in the lowland dipterocarp forest of the Subic Watershed Forest Reserve (SWFR) in Luzon Island, Philippines. Using pairs of day and night traps, we collected seeds during 3 mo of wet season and 3 mo of dry season in a 1.2-ha study site. Bird-dispersed seeds predominated over those dispersed by bats in terms of both seed abundance and number of seed species. The most abundant endozoochorous seed species were significantly biased toward either bird or bat dispersal. Birds and bats appeared to compete more strongly for fruit resources during the dry season than during the wet season, and bats responded more to changes in the seasons than birds did. GLM analyses showed that the factor that had the strongest influence on overall seed distribution was the number of fleshy-fruited trees surrounding the traps, and that the distribution pattern of day-dispersed seeds was affected by more physical factors (number of trees, size of trees, presence of fleshy-fruited and conspecific trees) in the study site than the pattern of night-dispersed seeds were. Given that birds are the more important dispersers in the study site, restoration efforts in SWFR might benefit by focusing on attracting these dispersers into its degraded habitats.     

The swimming abilities of recently settled post-larvae of Sillaginodes punctata

Sixty-four post-larvae of the King George whiting Sillaginodes punctata were tested in swimming chambers, against one of five flow-through velocities (2, 4, 6, 8 or 10 cm s ⁻¹) for up to a maximum of 120 min. Fish were determined by regression to have an FV₅₀ (50% fatigue velocity) of 6.0 cm s⁻¹. No fish survived the full 120 min at 10 cm s⁻¹. Sixteen individuals were tested in a swimming chamber against a flow-through velocity of 6 cm s ⁻¹ and allowed to swim to exhaustion. Fish swam between 25 and 538 min with a peak at c . 6–8 h. Total swimming time was not correlated with standard length of fish although the size range examined was narrow. Relative to recent studies on the swimming abilities of late-stage larvae of reef fishes, this study indicates that post-larval King George whiting are weak swimmers. The weak swimming ability of post-larval King George whiting is consistent with studies showing passive dispersal and recruitment of this species.     

Land‐use change and the ecological consequences of personality in small mammals

Many plants rely on animals for seed dispersal, but are all individuals equally effective at dispersing seeds? If not, then the loss of certain individual dispersers from populations could have cascade effects on ecosystems. Despite the importance of seed dispersal for forest ecosystems, variation among individual dispersers and whether land‐use change interferes with this process remains untested. Through a large‐scale field experiment conducted on small mammal seed dispersers, we show that an individual's personality affects its choice of seeds, as well as how distant and where seeds are cached. We also show that anthropogenic habitat modifications shift the distribution of personalities within a population, by increasing the proportion of bold, active, and anxious individuals and in‐turn affecting the potential survival and dispersal of seeds. We demonstrate that preserving diverse personality types within a population is critical for maintaining the key ecosystem function of seed dispersal.     

Patterns and drivers of aquatic invertebrate diversity across an arid biome

Managing and restoring faunal diversity across large areas requires an understanding of the roles of connectivity and dispersal in driving community patterns. We sought to determine the influence of connectivity, water regime, water source, geographical location, and dispersal traits on patterns of aquatic invertebrate diversity across a continent‐wide arid biome. We compiled data on freshwater invertebrate assemblages from sites spanning the breadth of arid Australia. Univariate analyses (analysis of variance and rarefaction) revealed that alpha and gamma diversity across sites decreased as latitude increased. Multivariate analyses (ordination and analysis of similarity) revealed that community composition had considerable fidelity to geographic regions. Hydrological connectivity was strongly associated with riverine community composition although water rarely flowed (often less than annually). Hydrologically isolated sites (springs and rockholes) supported communities that were markedly dissimilar to hydrologically connected sites, and to each other. We investigated the influence of dispersal on diversity patterns by examining distance decay relationships for each of four dispersal trait groups (obligate aquatic and passive, weak, and strong aerial dispersers) on the basis of geodesic (shortest path) distances between pairs of sites and Mantel tests. We did not detect clear differences between dispersal traits and distance decay relationships at the continental scale, even for the two groups with the lowest dispersal ability (obligate aquatics and passive dispersers). Our results suggest that the loss of hydrological connectivity from water developments in arid lands (for example, the impoundment of intermittent rivers) is likely to affect macroinvertebrates. However, the exact flow mechanisms underlying such changes remain to be determined.     

Macroalgal species diversity and biomass of subtidal communities of São Miguel (Azores)

The benthic algal communities of two subtidal sites, located on opposite coasts of S?o Miguel Island (S?o Roque in the south and S?o Vicente on the north coast), were studied over a 2 year period (September 1993–September 1995). At both sites the sublittoral region was surveyed from low-tide level down to a depth of 15 m. Qualitative and quantitative changes are described. A depth-related gradient in species diversity and biomass was found at both localities. In general, red algae such as corallines and Pterocladiella capillacea predominated in the shallow sublittoral (5 m) while brown algae such as Zonaria and Stypocaulon were more abundant at 15 m. Multivariate analyses emphasized the existence at each study site of two communities (5 and 15 m depth), separated by a large transition zone. The 15 m community on the south coast site showed the largest number of species (52), whereas the lowest diversity (30 species) also occurred at this site in the 5 m community. A clear seasonal pattern of biomass change could be discerned only at S?o Vicente where the highest biomass was recorded in spring/summer. No major inter-annual variations could be detected, indicating relatively stable communities at least on a short-term basis. Received in revised form: 28 March 2001 Electronic Publication