Spatial pattern, patch dynamics and successional change: chironomid assemblages in a Lake Erie coastal wetland

1. Distribution patterns in 1994 and 1995 of chironomid larvae in the sediments of wetlands on Presque Isle, Pennsylvania were used to determine whether relationships exist among species composition and wetland age, position or environmental characteristics. Canonical correspondence analysis was used to relate wetland age, degree of isolation, vegetation abundance and water chemistry to species composition.
2. Of forty-two chironomid taxa collected, Tanytarsus lugens group, Paratanytarsus sp. and Cladotanytarsus mancus group were the most widespread, but Lauterborniella agrayloides , Stictochironomus sp. and Cryptochironomus sp. were locally abundant in several sampling sites.
3. The canonical ordination based on all site characteristics was statistically significant and accounted for 54.9% of the variance in the species data and 69.5% of the variance in the species–environment relationship. The variables that contributed significantly to the ordination model were macrophyte abundance, conductivity and wetland area.
4. Partial canonical ordination based on environmental conditions with wetland age and position effects as covariables showed that environmental conditions explain a significant proportion of the variance in species composition among sites. Neither wetland age nor position variables explain a significant proportion of the variance in species composition.     

Spatial variation in insect community and species responses to habitat loss and plant community composition

Several experimental studies have examined species responses to manipulations of habitat area and spatial arrangement, but plant composition and spatial variation in species distributions also affect animal responses to habitat alteration. We used an experimental approach to study the combined effects of habitat area, edge, and plant community composition on the spatial structure of insect species richness and composition. The abundance of three guilds (herbivores, predators and parasitoids) and individual species were also analyzed. Habitat patches were created that differed in area and edge by selectively mowing portions of 15 m×15 m plots in a 1.7-ha old field. Spatial and environmental variables were used to predict insect responses in separate multiple regression and ordination models. The variation in species responses due to spatial and environmental variables was then partitioned by combining these variables into an overall regression or ordination. Spatial and environmental variables contributed similar percentages to the total variance in insect species richness, abundance or composition. No significant effects of habitat area were observed in any response variable. Herbivore abundance showed positive responses to legume or grass cover, as well as spatial variation that was unrelated to environmental variables. Predators and parasitoids had greater effects of plant species richness and habitat edge, and less unexplained spatial variation. Individual species differed in their responses to plant variables, depending on host specialization or intraspecific aggregation. Our study highlights the importance of plant community composition and spatial variation apart from environmental variables. Spatial variation stems both from species responses to environmental features as well as species differences in habitat specialization and intraspecific aggregation.     

Environmental and spatial variables influence the composition of frog assemblages in sub-tropical eastern Australia

In community ecology, contrasting theories suggest that the distribution and abundance of species, and thus the composition of assemblages, are influenced by i) environmental gradients, or ii) contagious biotic processes such as predation, competition, dispersal and disease. In the former case, sites with similar environments would tend to support similar assemblages, while in the latter, geographically proximate sites would tend to support more similar assemblages than widely separated sites. I investigated the relative influence of environmental variables and spatial position on the composition of frog assemblages at forest streams in sub-tropical eastern Australia using redundancy analysis (RDA) and partial RDA. Data on the maximum abundance of the frog species at 65 survey sites were transformed such that RDA would yield the Hellinger distance between sites. The following analysis identified 11 environmental variables that explained 45% of the variation in the abundance of species at the survey sites (the species matrix), as a proportion of total variance. The geographic co-ordinates of the survey sites accounted for 12%, while the environmental and spatial variables combined accounted for 47% of the variation in the species matrix. Partial redundancy analysis indicated that of the explained variation, 74% was purely environmental, 5% was purely spatial and 21% was spatial environmental variation. This study is the first to quantify the relative influence of environmental and spatial variables on the composition of amphibian assemblages. It provides support for both the environmental control model and the biotic control model of species' distributions and assemblage composition, although environmental variables appear to have the greater effect at this scale of analysis.     

Patterns in the distribution of fish assemblages in Río Amacuzac,Mexico: influence of abiotic factors and biotic factors

Patterns in spatial and seasonal distribution of fish communities were analyzed in the Río Amacuzac, Mexico, and their relationship to environmental variables and habitat characteristics. The PCA biplot of the study sites and environmental factors showed the first two axes accounting for 52.93% of the variance. The diagram showed the study sites ordination in environmental gradients. The first axis explained variables related to habitat characteristics and temperature (36.30%) and second axis arranged the sites in physicochemical and water quality environmental gradients (conductivity, dissolved oxygen, orthophosphates, ammonium, pH) displaying seasonal variation. Fifteen species were recorded, eight of them are exotic. One new record appeared for the Río Amacuzac: Pterygoplichthys disjunctivus is exotic. Study sites with highest species richness were: 5, 7 and 9 (twelve species each one); while the study sites with low species richness were 1, 2, 3 and 6 (eight species each one). Six of the species were distributed throughout the whole river. Based on the composition of the fish fauna, the study sites form two groups and the analysis of fish species displays three groups according to the Bray–Curtis index. The diagram of the canonical correspondence analysis relates environmental parameters to the abundance of fish species and showed that the first two axes exhibit 78.31% of the explained variance. Species richness had a spatial pattern associated to the introduction of exotic species for ornamental uses. According to the results of the importance value index (IVI), the dominant species were the poecilids Poeciliopsis gracilis and Heterandria bimaculata, small fishes that were introduced in the river. Río Amacuzac has a biotic alteration in the structure of fish communities due to the invasion of exotic species, representing risks to the integrity of the native fish fauna.     

Species co-occurrence, nestedness and guild–environment relationships in stream macroinvertebrates

1. Describing species distribution patterns and the underlying mechanisms is at the heart of ecological research. A number of recent studies have used null model approaches to explore mechanisms behind spatial variation in community structure.
2. However, unexplored questions are the degree to which single guilds of potentially competing stream macroinvertebrate species show: (i) interspecific segregation among-stream sites (i.e. occur together less often than expected by chance), suggesting competitive interactions; (ii) interspecific aggregation (i.e. occur together more often than expected by chance), suggesting similar responses to the environment; (iii) comply with nestedness, suggesting the existence of selective extinctions or colonisations and (iv) show similar environmental relationships.
3. The present analyses showed that guilds of stream macroinvertebrates exhibit non-random co-occurrence patterns that were generally contingent on the weighting of sites by stream size. Despite significant segregation of species, each guild also showed significantly nested patterns. Species richness was correlated with different environmental factors between the guilds, although these correlations were relatively low. By contrast, correlations between the major ordination axes and key environmental variables were slightly stronger in canonical correspondence analysis, and generally the same factors were most strongly correlated with variation in the species composition of each guild.
4. The present findings are the first to show that species within each stream macroinvertebrate guild show significant negative co-occurrence at the among-stream riffle scale. These findings present challenges for future studies that aim to disentangle whether these patterns comply with the habitat checkerboard or the competitive checkerboard explanations.     

Environmental factors and community dynamics at the southernmost part of the North American Graminetum – I. On the contribution of climatic factors to temporal variation in species composition

Compositional patterns of vegetation and their relationship to temporal and spatial environmental variation, with emphasis on climatic factors, were investigated in plant communities located in the southernmost portion of the North American Graminetum, in central México. Data from 353 samples, obtained in four ecologically contrasting plant communities during 11 years, were analyzed by partial canonical correspondence analysis. Eight climatic variables and eighteen covariables (seventeen edaphic and one resource management) were included in the ordination. A relationship between floristic change and weather variation, once covariables effects were fitted, was examined. Despite a strong contrast in ecological conditions among study sites, a set of four climatic variables was finally found in which each variable contributed independently and statistically (P < 0.01) to the total variance in the vegetation data. Thus, environmental variables other than climatic could not conceal the important role of weather as a mediator of floristic change through time. Summer precipitation and summer maximum temperature showed the highest correlations with the first two species axes, 0.77 and –0.39, respectively. Contribution of these two climatic variables to variance in the vegetation data explained by environmental variables was approximately 81%. Annual species were abundant during rainy years, while abundance of perennial grasses and shrubs showed no clear relationship to weather variation. This study explicitly probes the important role of rain patterns in shaping structure and composition of semiarid communities and considers how other environmental factors can affect plant communities at the southernmost part of the North American Graminetum.     

Classification, assessment and trophic reconstruction of Danish lakes using chironomids

1. Surface-sediment assemblages of subfossil chironomid head capsules from fifty-four primarily shallow and nutrient-rich Danish lakes were analysed using multivariate numerical techniques. The species data, comprising forty-one chironomid taxa, were compared to environmental monitoring data in order to establish a relationship between chironomid faunal composition and lake trophic state.
2. The subfossil assemblages were compared to the chironomid bathymetric distributions along transects from four lakes. Correspondence analysis and similarity coefficients showed that the subfossil assemblages, sampled in the lake centre, reflect the chironomid communities in the littoral at a depth of 2–7 m.
3. Two-way indicator species analysis (TWINSPAN) was used to classify the Danish lakes into five groups defined by trophic state, lake depth and pH. Eighteen chironomid taxa showed significant differences in abundance among the five groups. Canonical correspondence analysis (CCA) showed the chlorophyll a concentration ([Chl a ]) and Secchi depth to be the variables best correlated to the faunal data, and fourteen taxa were significantly correlated to [Chl a ].
4. The strong correlation between chironomid data and the ln-transformed ([Chl a ]) was used to create a weighted averaging (WA) model to infer lake trophic state. Several models were tested by cross validation (leave-one-out jack-knifing), and a simple WA model using inverse de-shrinking had a RMSEP_jack of 0.65 (ln units) and a r ²_jack of 0.67.
5. The results can be used in the assessment and reconstruction of lake trophic state for long-term monitoring and palaeoecological investigations of shallow, temperate lakes in the mesotrophic to hypertrophic nutrient range.     

The distribution of micro-arthropods in some southern English streams: the influence of physicochemistry

SUMMARY. 1. Micro-arthropods were surveyed during October 1986 at thirty stony, stream riffle sites in the Ashdown Forest, southern England.
2. The importance of a number of physicochemical variables in determining both the distribution of micro-arthropod taxa and community structure was assessed.
3. Acidic sites had an impoverished fauna. Total micro-arthropod species richness and densities were highest under circumneutral conditions. and the same patterns were shown by the Hydrachnellae, Harpucticoida and Cladocera. A number of species seemed indifferent to acidic conditions and were widespread and, as a group, the cyclopoid copepods showed no relationship with pH.
4. Multiple regression showed that other environmental variables, in particular annual mean temperature and maximum discharge, were also important in explaining the between-site distribution of separate micro-arthropod groups and individual species.
5. Other multivariate techniques (ordination, classification and multiple discriminant analysis) showed that impoundment linkage, source distance and conductivity, along with pH, were the most important variables in explaining patterns of species composition between sites.     

Ordination methods and chironomid species in stony bottom streams

Stony bottom streams in Italy were sampled in 369 sites in different years, and 174 chironomid species (or species groups) were identified.The species by sites matrix was submitted to different ordination methods, with the aim of outlining the major factors that are responsible of the observed species composition.The methods considered were principal component analysis (PCA), detrended correspondence analysis (DCA) and non-metric multidimensional scaling (MDSCAL). These methods assume different models describing the species-responses to the environmental gradients.PCA gave poor results, the best species ordination was given by MDSCAL. DCA gave results in agreement with MDSCAL. DCA results were useful as a starting point to perform MDSCAL. MDSCAL was performed with different options. At last Procrustean analysis was carried out to have a single fitted configuration, that summarized the results given by the different MDSCAL methods.All multivariate analyses emphasized that the first ordination axis can be interpreted as a crenonrithron-potamon gradient, probably an oxygen availability and/or a water temperature gradient, whereas the second axis can be considered a water speed gradient, separating lotic from lentic sites.The 3rd and 4th ordination axes had apparently no ecological meaning.Separated clusters of species were never evident, so well defined chironomid species assemblages could not be outlined in this analysis.     

Host diversity and environmental variables as determinants of the species richness of the parasitoids of leaf‐cutting ants

Aim Because of the obligatory relationship between endoparasitoids and their hosts, we presume that hosts exert strong selection pressure on parasitoids. One prediction is a positive relationship between host diversity and parasitoid richness. This relationship could be the product of resource availability which could lead to more opportunities for speciation, or could represent shared responses to the environment by both groups. Location Argentina and Paraguay. Methods We sampled a 1800‐km transect to test for a correlation between the richness of leaf‐cutting ant hosts and their phorid parasitoids. Regression models were used to assess if host and environmental variables could explain phorid species richness at nest, hectare and locality spatial scales. We used canonical correspondence analysis (CCA) to explore if there were similar responses of phorid species to particular host and environmental variables at different spatial scales, and partial CCA to separate the relative importance of both groups of variables. Results Phorid richness was positively correlated with host richness. Host richness/abundance accounted for 20–53% of the variation in parasitoid richness at the hectare and locality scales of analysis, with most of the variation accounted for by ant abundance. We were not able to assess the prediction at the nest scale as only one phorid species was found at most nests. Climatic variables did not explain phorid species richness once host variables were in the models. Partial CCA showed that host‐related variables accounted for most of the variance associated with phorid species ordination at the nest and hectare scales, but not at the largest grain, the locality, where climatic variables were more important. However, most phorid species did not show particular positions along the climatic gradient. Main conclusions The association between parasitoid richness and host richness and abundance, and the overall weak associations with environmental variables, suggest that these host variables are key factors influencing parasitoid speciation.     

Relationships between environmental variables and vegetation across mountain wetland sites,N. Iran

The mountain wetlands studied represent a unique habitat on the southern slopes of the Alborz mountain range, the second largest range in Iran. In comparison with other parts of this range the western section is ecologically and botanically unknown. Floristic and vegetation variation were assessed using diverse environmental variables along a broad altitudinal span (350 m to 3200 m a.s.l.). Using both statistical and ordination analyses floristic variation was assessed on three defined altitudinal belts which were delimited based on Alborz macro-climatic boundaries. The distribution of individual wetland plant species, of phytogeographic elements and of life-forms all differ among altitudinal belts. This result is also shown in both direct and indirect analyses of ordinations. The proportion of geophytes significantly increases with altitude and geophytes are very well represented in the upper altitudinal belt. The number of species of a narrow phytogeographical distribution (e.g. endemics) increases with altitude, soil pH and EC declined with altitude. The first axis of DCA ordination with passively projected environmental variables indicates that, organic matter and concentration of Fe²⁺ are increased toward higher altitude. The second axis of ordination is related to both soil texture and slope inclination. The distribution of species in the CCA species plot is also close to the distribution of those in the DCA ordination. This study indicates that altitude and slope together with other dependent environmental variables (pH, EC, Ca²⁺ and soil texture) are the main ecological factors controlling species distribution across the Western Alborz wetland sites.     

Describing termite assemblage structure in a Peruvian lowland tropical rain forest: a comparison of two alternative methods

Termites are frequently dominant invertebrate decomposers and bioturbators in lowland tropical forests and therefore strongly influence ecosystem processes favouring soil stability, porosity and nutrient retention. In this study, we provide the first spatially replicated dataset on termite assemblage composition, abundance and biomass in a Peruvian rainforest by sampling six separate plots. In addition, two alternative sampling methods (transect method-TM and quadrat method-QM), providing termite species density data, were compared among the plots. The relationships between a range of environmental and spatial variables and species composition were examined using canonical correspondence analysis variation partitioning. We found that the TM captured a higher proportion of the known species in the site (82 %) compared with the QM (66 %). In addition, 56 % of the species sampled by TM were common between the plots while only 18 % of species overlapped using the QM. The QM may therefore potentially have undersampled the species pool. Environmental variables were shown to explain a larger proportion of the species patterns than the spatial variables with elevation, soil temperature and distance to the river being the most important. We discuss the impacts of the environmental and spatial variables on termite species composition.     

Species-environment patterns of forest vegetation on the uplifted reef limestone of Atiu,Mangaia, Ma'uke and Miti'aro,Cook Islands

Abstract. We examined woody species composition and its relation to environmental variables in native forest growing on four limestone islands in the southern Cook Islands: Atiu, Mangaia, Ma'uke, and Miti'aro. Relative dominance (percent basal area) of woody species in 74 sites was sampled using the point-centered quarter method, and the data were analyzed using clustering and ordination techniques. These tropical forests have a relatively low diversity of native woody species (32 native and 10 introduced species occurred in our sites). Four forest types were recognized: Pandanus/Guettarda littoral forest (with several subtypes), Hernandia nymphaeifolia littoral forest, Barringtonia littoral forest, and makatea forest (dominated by Elaeocarpus tonganus and Hernandia moeren-houtiana). These types were related, using canonical correspondence analysis, to geographical attributes (wind-wardness, elevation, and proximity to the coast or roads) that served as surrogates for environmental variables (maritime influence, soil variation, and degree of human disturbance). The eigenvalues for this direct ordination were much lower than for indirect ordination (0.32 vs. 0.71 for the first axis), indicating that the measured geographical attributes could explain only a modest portion of the compositional variation.     

Diversity, distribution and larval habitats of North Swedish blackflies (Diptera: Simuliidae)

1. Blackfly species richness and community structure were analysed at fifty-six sites in northern Sweden in two seasons. The sites were situated in a wide range of streams and rivers from small springbrooks, bog streams and lake-outlet streams to medium-sized forest rivers and large rivers draining montane regions.
2. Thirty-nine blackfly species were found, with between two and thirteen species per site. Neither species richness nor abundance could be related to the environmental variables measured.
3. An analysis of labral fan size of blackflies indicated a clear trend for the prevalence of larvae with small fans in large rivers and larvae with larger fan size in small streams. Similarly, fan size related to current velocities so that large fans were associated with slow current velocities and small fans with high velocities.
4. A strong relationship existed between species composition and habitat, as seen in ordination by non-metric multidimensional scaling. The relationship found between fan size and habitat size-related variables, such as channel width, depth, velocity and substratum particle size, along with longitude and altitude, in partial least squares regression analysis offered an explanation of the species composition–habitat relationship.
5. In addition to testing that distributions of blackfly larvae reflect morphological traits, we tested two general hypotheses pertaining to distribution patterns: (a) that blackfly communities show bimodal distributions; and (b) that their distributions are nested. Neither of these two hypotheses was supported by our observations. However, widespread blackfly species were locally more abundant than those found at relatively few sites, thus showing a positive abundance–occupancy relationship.     

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.     

Structure of chironomid assemblages along environmental and geographical gradients in the Bohemian Forest lakes (Central Europe): An exploratory analysis

Eight glacial lakes of the Bohemian Forest (Czech Republic and Germany) were characterised by the distribution of chironomids collected as pupal exuviae. Twenty-eight taxa were identified, including some faunistically interesting species of the region. Two-way indicator species analysis (TWINSPAN) was used to classify lakes according to their taxonomic composition. Canonical correspondence analysis (CCA) and multiple regression were used to relate the chironomid assemblages to two sets of explanatory variables: (i) local environmental variables, and (ii) broad-scale spatial variables. The TWINSPAN classified the lakes into four groups, whereas presence/absence of three taxa was indicative for this classification. The CCA of assemblage composition on environmental variables showed that chironomids respond significantly to altitude and alkalinity. The ordination of composition data on geographical variables revealed strong longitudinal gradient in chironomid distributions. Altitude and alkalinity accounted for 36.2% of the total variation, while the geographic gradient explained 20.5%. As revealed by the variation partitioning procedure, the significant effect of these variables was, in large part, independent of each other. Overall taxonomic richness appeared to be governed by altitude only. Causal ecological and historical factors underlying these results are discussed. This paper may provide a basis for hypothesis testing in future research of the Bohemian Forest lakes.     

Altitudinal gradients in stream fish diversity and the prevalence of diadromy in the Sixaola River basin,Costa Rica

Landscape-scale patterns of freshwater fish diversity and assemblage structure remain poorly documented in many areas of Central America, while aquatic ecosystems throughout the region have been impacted by habitat degradation and hydrologic alterations. Diadromous fishes may be especially vulnerable to these changes, but there is currently very little information available regarding their distribution and abundance in Central American river systems. We sampled small streams at 20 sites in the Sixaola River basin in southeastern Costa Rica to examine altitudinal variation in the diversity and species composition of stream fish assemblages, with a particular focus on diadromous species. A set of environmental variables was also measured in the study sites to evaluate how changes in fish assemblage structure were related to gradients in stream habitat. Overall, fish diversity and abundance declined steeply with increasing elevation, with very limited species turnover. The contribution of diadromous fishes to local species richness and abundance increased significantly with elevation, and diadromous species dominated assemblages at the highest elevation sites. Ordination of the sampling sites based on fish species composition generally arranged sites by elevation, but also showed some clustering based on geographic proximity. The dominant gradient in fish community structure was strongly correlated with an altitudinal habitat gradient identified through ordination of the environmental variables. The variation we observed in stream fish assemblages over relatively small spatial scales has significant conservation implications and highlights the ecological importance of longitudinal connectivity in Central American river systems.     

Beta diversity of rock-restricted cichlid fishes in Lake Malawi: importance of environmental and spatial factors

The rock-restricted cichlid fish assemblages of Lake Malawi exhibit high spatial diversity in their species composition and relative abundance. However the extent to which this is due to the effects of local environmental differences, dispersal limitation of constituent taxa, and the assignment of allopatric populations to species is uncertain. We examined the factors associated with diversity within an assemblage from the north-western shores, encompassing a spatial scale of 170 km. For both the whole assemblage, and all constituent species-complexes, spatial variance in community structure was significantly dependent upon both geographic distances between locations and local habitat variables. Pronounced effects of distance indicate limited dispersal, but our results also show that that the spatial variance explained by geographic distance alone was strongly linked to proportion of allopatric populations within a species-complex with species status. Thus, the taxonomic status of allopatric populations underlies, at least partially, the biogeographical structure of this assemblage. Substrate composition and habitat depth were also significant determinants of community structure, although spatial variance attributed to these variables was less than that associated with distance alone. Substantial unexplained variance may be a consequence of the effects of unmeasured habitat variables, high ecological similarity between co-occurring species, stochastic influences on population abundance, and the effects of local adaptation. Despite low spatial variance explained by the assessed environmental variables, significant environmental influence on cichlid assemblage structure across a wide spatial scale indicates that even slight future environmental changes may have the capacity to significantly alter species composition.