Seasonal changes in a cactus‐hoverfly (Diptera: Syrphidae) network

1. We analysed a cacti‐syrphid community focusing on the exploitation of decaying cacti resources by the flies, i.e. if exploitation exhibited a nested topology as a structural pattern, or whether it was temporally random. If availability of cactus resources was predictable during the rainy season, it would then be exploited by a more structured community, or as resource availability would be unpredictable during the dry season, we should expect it to be exploited by a random community. 2. We registered 12 Copestylum species (9 in dry and 11 in rainy season). Four cactus species were recorded per season, three were shared between seasons. 3. The community of Copestylum larvae in the rainy season was not randomly assembled but highly nested, revealing a highly structured pattern of resource use. It exhibited a random organization for the dry season. The high nestedness value obtained for the rainy season suggests that factors along with competition must play a major role in determining community structure. 4. Succession in the cacti‐syrphid community mediated by microorganisms involved in necrosis is an important factor structuring nested subsets. The studied networks were small, which may limit the power of the analysis, and strong conclusions could also be limited.     

Flower resource and land management drives hoverfly communities and bee abundance in seminatural and agricultural grasslands

Pollination is a key ecosystem service, and appropriate management, particularly in agricultural systems, is essential to maintain a diversity of pollinator guilds. However, management recommendations frequently focus on maintaining plant communities, with the assumption that associated invertebrate populations will be sustained. We tested whether plant community, flower resources, and soil moisture would influence hoverfly (Syrphidae) abundance and species richness in floristically‐rich seminatural and floristically impoverished agricultural grassland communities in Wales (U.K.) and compared these to two Hymenoptera genera, Bombus, and Lasioglossum. Interactions between environmental variables were tested using generalized linear modeling, and hoverfly community composition examined using canonical correspondence analysis. There was no difference in hoverfly abundance, species richness, or bee abundance, between grassland types. There was a positive association between hoverfly abundance, species richness, and flower abundance in unimproved grasslands. However, this was not evident in agriculturally improved grassland, possibly reflecting intrinsically low flower resource in these habitats, or the presence of plant species with low or relatively inaccessible nectar resources. There was no association between soil moisture content and hoverfly abundance or species richness. Hoverfly community composition was influenced by agricultural improvement and the amount of flower resource. Hoverfly species with semiaquatic larvae were associated with both seminatural and agricultural wet grasslands, possibly because of localized larval habitat. Despite the absence of differences in hoverfly abundance and species richness, distinct hoverfly communities are associated with marshy grasslands, agriculturally improved marshy grasslands, and unimproved dry grasslands, but not with improved dry grasslands. Grassland plant community cannot be used as a proxy for pollinator community. Management of grasslands should aim to maximize the pollinator feeding resource, as well as maintain plant communities. Retaining waterlogged ground may enhance the number of hoverflies with semiaquatic larvae.     

Influence of season on leaf litter ant (Hymenoptera: Formicidae) diversity in the locality of Minko'o (Southern,Cameroon)

Seasonality is known to influence ant activity in many tropical rain forests in the world such as South America and Africa. We surveyed ant fauna in the leaf litter in the locality of Minko'o. The work aimed to evaluate the effect of seasonal variation on the diversity and composition of litter ants. Ants were sampled from November 2015 to June 2017, using four sampling methods: visual capture, bait, pitfall trap and extraction. Species richness, Shannon diversity index and analysis of similarities were used to characterise diversity of ant communities between seasons. We collected 306 ant species, shared out between 56 genera and 11 subfamilies. Subfamilies Myrmicinae, Ponerinae, Dolichoderinae, Formicinae, Dorylinae, Cerapachyinae and Pseudomyrmecinae occurred in all the seasons. Species richness was highest in major dry season with 243 species followed by minor rainy season with 188, major rainy season with 177 species and finally minor dry season that recorded the lowest with 155 species. Kruskal–Wallis test showed that ant species richness did not differ between seasons (p > 0.05). Species diversity index indicated that diversity was the highest during minor dry season (H′ = 4.24), followed by the major dry season (H′ = 4.23), minor rainy season (H′ = 4.21) and lowest during major rainy season (H′ = 4.06). Eight most frequents ants have been recorded: Axinidris sp.1, Camponotus flavomarginatus, Monomorium guineense, Myrmicaria opaciventris, Odontomachus troglodytes, Carebara perpusilla, Paltothyreus tarsatus and Pheidole megacephala. Assessment of the seasonal effect on diversity reveals that dry season is richer and more diverse than rainy season and the season significantly influence the diversity of litter ants.     

Bacterial community structure in the rhizosphere of three cactus species from semi-arid highlands in central Mexico

The nature reserve of Tehuacan-Cuicatlan in central Mexico is known for its diversity and endemism mainly in cactus plants. Although the xerophytic flora is reasonably documented, the bacterial communities associated with these species have been largely neglected. We assessed the diversity and composition of bacterial communities in bulk (non-rhizospheric) soil and the rhizosphere of three cactus plant species: Mammillaria carnea, Opuntia pilifera and Stenocereus stellatus, approached using cultivation and molecular techniques, considering the possible effect of dry and rainy seasons. Cultivation-dependent methods were focused on putative N₂-fixers and heterotrophic aerobic bacteria, in the two media tested the values obtained for dry season samples grouped together regardless of the sample type (rhizospheric or non-rhizospheric), these groups also included the non-rhizospheric sample for rainy season, on each medium. These CFU values were smaller and significantly different from those obtained on rhizospheric samples from rainy season. Genera composition among isolates of the rhizospheric samples was very similar for each season, the most abundant taxa being α-Proteobacteria, Actinobacteria and Firmicutes. Interestingly, the genus Ochrobactrum was highly represented among rhizospheric samples, when cultured in N-free medium. The structure of the bacterial communities was approached with molecular techniques targeting partial 16S rRNA sequences such as denaturing gradient gel electrophoresis and serial analysis of ribosomal sequence tags. Under these approaches, the most represented bacterial phyla were Actinobacteria, Proteobacteria and Acidobacteria. The first two were also highly represented when using isolation techniques.     

Diversity of parasites in Cichlasoma amazonarum Kullander, 1983 during rainy and dry seasons in eastern Amazon (Brazil)

This study investigated the influence of the rainy and dry seasons on the parasite communities of Cichlasoma amazonarum (Kullander, 1983) in a tributary of the Amazon River system, northern Brazil. Of 112 fish examined, 95.5% were parasitized by Ichthyophthirius multifiliis, Piscinoodinium pillulare, Gussevia disparoides, Posthodiplostomum sp., Procamallanus (Spirocamallanus) inopinatus, and Echinorhynchus paranensis, as well as by the Glossiphoniidae leech. Ichthyophthirius multifiliis and P. pillulare were the dominant parasite species, with I. multifiliis the most prevalent and abundant; the leech (Glossiphoniidae) was the least prevalent and least abundant. Parasites presented an aggregate dispersion pattern and seasonal variations in infestation levels, influenced by the environmental conditions in the rainy season. Variation in the parasite dynamics created overall changes in the parasite communities, characterized by greater diversity, species richness and evenness during the rainy season. However, there was a high similarity (99.8%) in the parasite community structure between the rainy and dry seasons. During the dry season the hosts had predominantly 1–3 parasite species compared to 3–4 parasites in the rainy season. Some parasites in the eastern Amazon undergo population changes relating to seasonality. These results thus indicate the correct season to apply adequate prophylactic measures to reduce negative impacts of parasites in this wild ornamental fish when captured.     

Effects of grazing intensity on pollinator abundance and diversity,and on pollination services

1. Pollinating insects provide important ecosystem services and are influenced by the intensity of grazing. Based on the Intermediate Disturbance Hypothesis (IDH), pollinator diversity is expected to peak at intermediate grazing intensities. However, this hump‐shaped relationship is rarely found. 2. The effect of grazing intensity was tested on flower cover, on the abundance and richness of bees, hoverflies and bee flies, and on pollination services to early‐flowering bee‐pollinated Asphodelus ramosus L. For that, we used data on 11 plant–pollinator phryganic communities from Lesvos Island (Greece) widely differing in grazing intensities. 3. Flower abundance and richness showed hump‐shaped relationships with grazing intensity. Grazing affected the abundance and richness of bees and hoverflies directly and also indirectly, through changes in the flower community. Grazing influenced directly the richness but not the abundance of bee flies. Overall, pollinator abundance and richness showed hump‐shaped relationships with grazing intensity, but variations in strength (hoverfly abundance) and direction (bee community) of the effect appeared along the season. Early in the season, grazing increased bee abundance but decreased richness, resulting in increased pollen limitation in A. ramosus. 4. The effects of grazing on pollinators vary with the intensity of the disturbance, generally supporting the IDH, and the timing of land‐use activities may influence pollination services. Management strategies should include moderate grazing levels to preserve overall diversity in this area, however, the conservation of particular early bee or bee‐pollinated species may benefit from reduced grazing in early spring.     

Abundance of arbuscular mycorrhizal fungi spores in different environments in a tropical rain forest, Veracruz, Mexico

 The aim of this study was to compare mycorrhizal abundance and diversity in sites with different regimes of disturbance in a tropical rain forest at Los Tuxtlas, Veracruz, Mexico. Arbuscular mycorrhizal spores were quantified at two sites: closed canopy and gaps in the forest. Data were recorded during dry, rainy, and windy ("nortes") seasons. Spores of eight Glomus species, sporocarps of three Sclerocystis species, three species of Acaulospora and two of Gigaspora were found. Significant differences in the number of species and spores were found among seasons. The highest numbers of species and spores were observed during the dry season, with a marked decrease during the rainy season. Our results show that disturbance does not but seasonality does affect abundance and richness of mycorrhizal spores in this tropical wet forest. Accepted: 11 October 1998     

Understory spider diversity in two remnants of tropical montane cloud forest in Chiapas, Mexico

We evaluated the spider diversity of a tropical montane cloud forest understory in two nearby sites with different degree of human disturbance at the Biosphere Reserve Volcán Tacaná, Chiapas, Mexico. The study was conducted over a 24 days period distributed in 6 months in 2009, covering dry and rainy seasons. A total of 8,370 spiders (1,208 adults and 7,162 juveniles) were collected. Determined specimens (7,747) represented 112 species and morphospecies, 71 genera and 22 families. The results showed that human disturbance has an influence on spider communities: species richness was significantly higher in the preserved site as regards to the disturbed site. Despite their proximity, the composition of spider communities showed only a moderate similarity between the two sites. No differences in abundance were found among sites when considering the whole sample, but sites differed clearly when seasons were analyzed separately. The rainy season had a negative effect on the abundance of spiders in the preserved site. Although the spider community structure was very similar between sites, there was a trend towards a greater species evenness in the preserved site for the whole sampling period and for the dry season.     

Seasonal variation in abundance and diversity of eavesdropping frog‐biting midges (Diptera,Corethrellidae) in a neotropical rainforest

1. In the tropics, precipitation patterns result in seasonal fluctuations in the abundance and distribution of plant and animal species. Tropical predators and parasites are therefore faced with seasonal changes in prey and host availability. 2. This study investigates the seasonal interaction among a specialised ectoparasite, eavesdropping frog‐biting midges (Corethrella spp.), and their anuran hosts, examining how the abundance and diversity of the frog‐biting midge community fluctuate between the rainy (host abundant) and dry (host sparse) seasons. 3. Midges were captured in both the rainy and dry seasons using acoustic playbacks of calls from a common frog species that breeds during the rainy season, the túngara frog (Engystomops, Physalaemus, pustulosus). During the dry season túngara frog choruses are absent. To explore seasonal shifts in host preference or changes in the midge community due to host specificity, midges were also captured using playbacks of calls from a frog that breeds during the dry season, the pug‐nosed tree frog (Smilisca sila). 4. While the overall abundance of midges decreased in the dry season, only slight differences in the relative abundance between midge species were found. These results suggest that midge populations can shift between hosts as they become available across seasons, allowing adult populations of frog‐biting midges to persist year‐round. To overcome the challenge of detecting and localising different host species, it is proposed that frog‐biting midges have evolved a generalised acoustic template, allowing them to respond to a broad range of available hosts, regardless of seasonal host composition.     

Spatial and Temporal Patterns of Diversity and Distribution of the Upper Juruá River Fish Community (Brazilian Amazon)

We studied spatial and temporal patterns in fish species composition and diversity at the upper Juruá River located in the west Brazilian Amazon. We collected with gillnet 822 fishes belonging to 90 species in the main Juruá River, its tributaries and the floodplain lakes during wet and dry seasons. Fish abundance and species richness were greater in the dry season. During that season, fishes may be concentrated due to the low water level, being caught more easily by gillnets. There has been a trend towards a greater fish biomass caught in lakes. This might be associated with a greater environmental stability as lakes may be less subject to large variations in water level. The fish communities differed between the two seasons and between lakes and the lotic environments (main river and tributaries). Fish species from the family Curimatidae were most abundant in the lakes, while Pimelodus spp. and Hypostomus spp. predominated in the main Juruá River. Seasonal variations in fish communities may be related to differences in the migratory behavior among fish species. Such spatial and temporal patterns influencing fish community structure at the Upper Juruá Extractive Reserve must be accounted for in management and conservation strategies.     

Multifaceted approach for the analysis of the phototrophic microbial community in a freshwater recreational area of Xochimilco,México

The freshwater microbial community in a recreational area of Xochimilco, México was investigated and compared based on spatial (three different sites) and temporal (dry and rainy seasons) environmental variables. Many of the 16S- and 18S rRNA gene sequences recovered by DGGE fingerprinting analysis were related to phototrophic microbial phylotypes of known identity. Our genetic and morphological analysis indicated the ubiquitous presence of the microeukaryotic green algae Desmodesmus-Scenedesmus spp. and of the unicellular cyanobacteria Cyanobium spp. as the most representative populations in the samples. While 18S rRNA-DGGE fingerprinting analysis revealed a homogeneous community composition across sites and seasons, the 16S rRNA showed significant differences between localities and seasons. None of the cyanobacteria species with potential to produce toxins were identified across the investigated samples. Correlations between biotic and abiotic variables evidenced an important difference between the dry and the rainy season, with a greater consistency in data from the rainy season. According to Principal Component Analysis (PCA), a strong relation between inorganic nitrogen, species richness, and subaquatic irradiance determines environmental variability in Xochimilco. Complementary and relevant data in results obtained from microscopy, fingerprinting, and statistical analysis applied in ecology indicate that a multifaceted approach to the study of microbial communities is necessary to accomplish a comprehensive scientific framework and to generate proper management strategies.     

Temporal dynamics of algal epiphytes on Leathesia marina and Colpomenia sinuosa macrothalli (Phaeophyceae)

This is the first study on the temporal dynamics of the epiphyte communities of Leathesia marina and Colpomenia sinuosa, in terms of richness, diversity and abundance, and their relationships between the hosts and the environmental factors. A total of 31 epiphyte taxa were recorded, with the two hosts sharing 64.5% of the epiphytic species found. The red algal epiphytic group had the highest abundance on both hosts and the diatoms presented the highest species richness. The epiphyte species on L. marina showed a progressive increase in abundance during the warm seasons that was related to the end of the macroscopic life stage of the host. They also showed a greater specific richness on L. marina than on C. sinuosa, but a lower diversity and evenness, and a clear dominance of the species Urospora penicilliformis and Erythrotrichia carnea. Low abundance of epiphytes was found on C. sinuosa, without temporal variation, but high diversity and evenness and without any dominant species. In addition, a clear zonation was observed on this host, where epiphytic abundance decreased on the lateral and upper part of the thallus.     

Impact of human disturbance on bee pollinator communities in savanna and agricultural sites in Burkina Faso,West Africa

All over the world, pollinators are threatened by land‐use change involving degradation of seminatural habitats or conversion into agricultural land. Such disturbance often leads to lowered pollinator abundance and/or diversity, which might reduce crop yield in adjacent agricultural areas. For West Africa, changes in bee communities across disturbance gradients from savanna to agricultural land are mainly unknown. In this study, we monitored for the impact of human disturbance on bee communities in savanna and crop fields. We chose three savanna areas of varying disturbance intensity (low, medium, and high) in the South Sudanian zone of Burkina Faso, based on land‐use/land cover data via Landsat images, and selected nearby cotton and sesame fields. During 21 months covering two rainy and two dry seasons in 2014 and 2015, we captured bees using pan traps. Spatial and temporal patterns of bee species abundance, richness, evenness and community structure were assessed. In total, 35,469 bee specimens were caught on 12 savanna sites and 22 fields, comprising 97 species of 32 genera. Bee abundance was highest at intermediate disturbance in the rainy season. Species richness and evenness did not differ significantly. Bee communities at medium and highly disturbed savanna sites comprised only subsets of those at low disturbed sites. An across‐habitat spillover of bees (mostly abundant social bee species) from savanna into crop fields was observed during the rainy season when crops are mass‐flowering, whereas most savanna plants are not in bloom. Despite disturbance intensification, our findings suggest that wild bee communities can persist in anthropogenic landscapes and that some species even benefitted disproportionally. West African areas of crop production such as for cotton and sesame may serve as important food resources for bee species in times when resources in the savanna are scarce and receive at the same time considerable pollination service.     

张掖国家湿地公园优势鸟类种群生态位研究

鸟类时空格局影响下鸟类种群生态位的变化是认识鸟类环境适应机制的理论基础。在张掖国家湿地公园保育区,采用样带法研究了冬春季节5种优势鸟类种群生态位宽度和生态位重叠的时间异质性。结果表明:随着月份的更替,5种优势鸟类种群的生态位宽度呈现出先增大后减小的变化趋势,赤麻鸭(Tadornaferruginea)、绿头鸭(Anasplatyrhynchos)、麻雀(Passer montanus)、白骨顶鸡(Fulicaatra)的最高值出现在1月分别为1.601、1.415、1.627、1.771,大白鹭(Ardea alba)的最高值出现在3月为0.556;不同月份间鸟类优势种群生态位重叠存在差异,3月、11月生态位的重叠值最大,1月份最小;优势鸟类种群的生态位宽度和重叠在时间尺度上表现出"此消彼长"的关系。湿地公园时间尺度上的鸟类生态位宽度和重叠的差异与湿地食物资源及隐蔽条件、鸟类群体迁徙规律、鸟类种间的竞争密切相关,反应了湿地鸟类适应生境变化的可塑性机制。     

Spatial and temporal variation in fish community structure of a marine embayment in Zanzibar,Tanzania

Spatial and temporal variation in the fish community structure were studied in a tropical non-estuarine embayment in Chwaka Bay, Zanzibar (Tanzania). Fish samples were collected bi-monthly (at each spring low tide) for 1 year (November 2001–October 2002) from a range of bay habitats ranging from mangroves deep inside the bay to seagrass beds close to the mouth of the bay. Additionally, environmental variables were examined to determine their relationship with the fish community structure. Being a non-estuarine embayment, the environmental variables as well as the fish community structure in each habitat remained relatively constant for most part of the year; however, a marked decline was observed during the rainy period (April–May). Significant variations in fish community variables (density, biomass and species richness) and in water temperature and salinity were observed during the rainy season in all habitats, with larger changes in the mangrove and mud/sand flats habitats than in the seagrass beds. Seasonal variations in water clarity and dissolved oxygen were not significant, though. Many species disappeared from the mangrove and mud/sand flats habitats during the rainy season and those which persisted showed a remarkable decrease in density. Moreover, the results indicate that mangroves were the preferred settling habitats for Gerres filamentosus, Gerres oyena, Lethrinus lentjan and Monodactylus argenteus, especially during the dry period (December–February) before the rainy season. This observation is contrary to what has been reported from some other tropical regions where greater abundance and species richness was observed during the rainy season. A significant relationship was found between density of fish and temperature, salinity and turbidity. Since salinity was the most conspicuously changing environmental variable with seasons, we propose that salinity, alone or in combination with low visibility and temperature, was probably the most important environmental factor structuring the fish assemblage in the mangrove and mud/sand flats habitats, particularly during the rainy season. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users. Handling editor: J. A. Cambray     

Impact of exotic invasion on the temporal stability of natural annual plant communities

Much work in ecology has focused on understanding how changes in community diversity and composition will affect the temporal stability of communities (the degree of fluctuations in community abundance or biomass over time). While theory suggests diversity and dominant species can enhance temporal stability, empirical work has tended to focus on testing the effect of diversity, often using synthetic communities created with high species evenness. We use a complementary approach by studying the temporal stability of natural plant communities invaded by a dominant exotic, Erodium cicutarium. Invasion was associated with a significant decline in community diversity and change in the identity of the dominant species allowing us to evaluate predictions about how these changes might affect temporal stability. Community temporal stability was not correlated with community richness or diversity prior to invasion. Following invasion, community stability was again not correlated with community richness but was negatively correlated with community diversity. Before and after invasion, community stability was positively correlated with the stability of the most dominant species in the community, even though the identity of the dominant species changed from a native (prior to invasion) to an exotic species. Our results demonstrate that invasion by a dominant exotic species may reduce diversity without negatively affecting the temporal stability of natural communities. These findings add support to the idea that dominant species can strongly affect temporal stability, independent of community diversity.     

N<Subscript>2</Subscript> fixation in three perennial <Emphasis Type="Italic">Trifolium</Emphasis> species in experimental grasslands of varied plant species richness and composition

This study is the first to investigate quantitative effects of plant community composition and diversity on N₂ fixation in legumes. N₂ fixation in three perennial Trifolium species grown in field plots with varied number of neighbouring species was evaluated with the ¹⁵N natural abundance method (two field sites, several growing seasons, no N addition) and the isotope dilution method (one site, one growing season, 5 g N m⁻²). The proportion of plant N derived from N₂ fixation, pNdfa, was generally high, but the N addition decreased pNdfa, especially in species-poor communities. Also following N addition, the presence of grasses in species-rich communities increased pNdfa in T. hybridum and T. repens L., while legume abundance had the opposite effect. In T. repens, competition for light from grasses appeared to limit growth and thereby the amount of N₂ fixed at the plant level, expressed as mg N₂ fixed per sown seed. We conclude that the occurrence of diversity effects seems to be largely context dependent, with soil N availability being a major determinant, and that species composition and functional traits are more important than species richness regarding how neighbouring plant species influence N₂ fixation in legumes.     

Cumulative annual dung beetle diversity in Mediterranean seasonal environments

Species diversity assessments should consider the dynamic nature of ecological communities, especially in highly seasonal ecosystems. Here we provide a comprehensive framework for analysing seasonal changes in species composition, richness and diversity in two local dung beetle (Coleoptera: Scarabaeoidea: Aphodiidae, Geotrupidae, and Scarabaeidae) communities from Western Tuscany (Italy), in the Mediterranean ecoregion. We test whether, in this highly seasonal region, cumulative annual diversity is an oversimplification of well differentiated seasonal communities. Data were obtained through repeated standardised samples collected regularly over an entire year. We clearly identify different summer and winter communities at each site based on species composition and abundance. Seasonal richness and diversity values are different from the cumulative annual values, as a consequence of beta diversity between seasons, and some dung beetle species are identified as idiosyncratic of each particular season. Both ecological (niche partitioning) and biogeographical factors are suggested as drivers of these temporal variations. Thus, because local inventories of fauna that include records over long time periods actually reflect situations where coexistence and interactions are unlikely to occur, highly seasonal sites must be viewed as having temporally differentiated communities in order to reach feasible and reliable baselines for local diversity assessments.     

Timing of disturbance,top‐down,and bottom‐up driving on early algal succession patterns in a tropical intertidal community

Herbivory and nutrient enrichment are major drivers of the dynamics of algal communities. However, their effects on algal abundance are under the influence of seasons. This study investigated the effects of herbivory and nutrient enrichment on early algal succession patterns using cages (uncaged and fully caged treatments) and two nutrient levels (ambient and enriched concentrations). To determine seasonal influences, experiment plots on dead coral patches were cleared during both dry and rainy season. Of the 17 algal species recruited in the experiment plots, three were dominant: Ulva paradoxa C. Agardh, Padina in the Vaughaniella stage, and Polysiphonia sphaerocarpa Børgesen. In this succession process, U. paradoxa was the earliest colonizer and occupied the cleared plots within the first month after clearing with the highest percentage of 83.33 ± 1.67% to 88.33 ± 9.28%. Then, it was replaced by the late successional algae, Padina in the Vaughaniella stage, and P. sphaerocarpa. The effects of herbivory and nutrient enrichment on algal abundance varied across algal functional groups and seasons. During the dry season, neither herbivory nor nutrient enrichment affected Ulva cover but during the rainy season, Ulva cover was influenced by nutrient enrichment. However, the abundance of algae in this early stage was not apparently affected by either herbivory or nutrient enrichment. Our results indicated that the timing of disturbance strongly influenced the algal abundance and successional patterns in this tropical intertidal community.     

Change in pattern diversity during secondary succession in Estonian forests

Comparisons of observed variance in species diversity (exp LT) and in species richness with expectation assuming a random and independent distribution of species are used to assess the relative importance during succession of niche limitation (Wilson, Gitay & Agnew 1987; limitation of the abundance or occurrence of species by competitive interactions) and nucleation (Yarranton & Morrison 1974; the development of a community through chance establishment or persistence followed by vegetative expansion from those nuclei). Wilson, Gitay & Agnew (1987) and Palmer (1987) suggest a deficit (i.e. lower than expected) in variance of richness is consistent with a ‘niche limitation’ process where competitive sorting makes patches more similar in species number than would be expected. In contrast, variance that is greater than expected is consistent with and could result from underlying environmental heterogeneity (i.e. ‘waterhole effect’). We extend these ideas to suggest that, in addition, a deficit in variance of diversity (expLT) can be interpreted as indicating niche limitation. Further, we suggest that a deficit of variance in richness need not be interpreted as resulting from niche limitation, but could result from limited dispersal and establishment followed by localized spread that causes quadrats to have less compositional overlap than might be expected if species were distributed at random. However, there is little reason to expect that such nucleation would cause a similar reduction in the variance of diversity. First, expLT is relatively insensitive to the presence or absence of rare species, and second, if local competitive processes do restrict the values of exp LT through some form of niche limitation, this could well be independent of the specific species involved. Thus, a combination of lower than expected variance in richness and high variance in diversity suggests nucleation to be important, particularly if the mean pairwise similarity between samples is lower than expected. We first examine a secondary successional sere with stands representing post-logging and mature forest. The variance of diversity is low in the establishment phase, but not in subsequent phases where the tree canopy is well developed. Significantly low variance of species richness is observed in a young forest just entering the thinning phase. Thus, nucleation may be taking place in the establishment phase of forest development, and is almost certainly important in the early thinning phase. Second, we examine communities in the process of equilibration following fertilization, logging, and paludification. All these communities have a low variance in diversity (exp LT) suggesting niche limitation.