Evaluation of methods for estimating macroinvertebrate species richness using individual stones in tropical streams

1. The most straightforward way to assess diversity in a site is the species count. However, a relatively large sample is needed for a reliable result because of the presence of many rare species in rich assemblages. The use of richness estimation methods is suggested by many authors as a solution for this problem in many cases.
2. We examined the performance of 13 methods for estimating richness of stream macroinvertebrates inhabiting riffles both at local (stream) and regional (catchment) scales. The evaluation was based on (1) the smallest sub-sample size needed to estimate total richness in the sample, (2) constancy of this size, (3) lack of erratic behaviour in curve shape and (4) similarity in curve shape through different data sets. Samples were from three single stream sites (local) and three from several streams within the same catchment basin (regional). All collections were made from protected forest areas in south-east Brazil.
3. All estimation methods were dependent on sub-sample size, producing higher estimates when using larger sub-sample sizes. The Stout and Vandermeer method estimated total richness in the samples with the smallest sub-sample size, but showed some erratic behaviour at small sub-sample sizes, and the estimated curves were not similar among the six samples. The Bootstrap method was the best estimator in relation to constancy of sub-sample sizes, but needed an unacceptably large sub-sample to estimate total richness in the samples. The second order Jackknife method was the second best estimator both for minimum sub-sample size and constancy of this size and we suggest its use in future studies of diversity in tropical streams. Despite the inferior performance of several other methods, some produced acceptable results. Comments are made on the utility of using these estimators for predicting species richness in an area and for comparative purposes in diversity studies.     

Drivers of beta diversity of macroinvertebrate communities in tropical forest streams

1. There has recently been increasing interest in patterns of beta diversity but we still lack a comprehensive understanding of these patterns in various regions (e.g. the tropics), ecosystems (e.g. streams) and organism groups (e.g. invertebrates). 2. Our aim was to investigate the patterns of beta diversity of stream macroinvertebrates in relation to key environmental (i.e. stream size, pH and habitat degradation) and geographical variables (i.e. latitude, longitude, altitude) in a tropical region. We surveyed a total of 8–10 riffle sites in each of 34 streams (altogether 337 riffle sites were sampled) in Peninsular Malaysia to examine variation in macroinvertebrate community composition at within‐stream and among‐stream scales. 3. Based on test of homogeneity of dispersion, we found that the streams studied differed significantly in within‐stream variation in community composition (i.e. among‐site variation of within stream beta diversity). The patterns were similar based on Bray–Curtis coefficient on abundance data, Sorensen coefficient on presence–absence data and Simpson coefficient on presence–absence data. We also found that within‐stream beta diversity was significantly related to stream size, pH and latitude, with each of these variables individually accounting for around 20% of the variation in beta diversity in simple regressions, while the total variation explained by the three significant variables amounted to around 50% in multiple regressions. By contrast, habitat degradation, longitude and altitude were not significantly related to beta diversity. We also found that the factor drainage basin accounted for much of the variation in beta diversity in general linear models, suppressing the effects of environmental variables. 4. We concluded that within‐stream beta diversity is mainly related to a combination of the identity of a drainage basin and stream environmental factors. Our findings provide important background for stream environmental assessment and conservation planning by emphasising that (i) macroinvertebrate communities within streams are not homogeneous, but show considerable beta diversity, (ii) streams differ in their degree of within‐stream beta diversity, (iii) stream size and water pH should be considered in applied contexts related to within‐stream beta diversity and (iv) historical effects may be different in different drainage basins and may affect present‐day patterns of within‐stream beta diversity.     

Temporal variability in fish assemblages from disturbed and undisturbed streams

Fish assemblages were sampled by electrofishingover a two- to ten-year period in undisturbedand anthropogenically disturbed South Carolinacoastal plain streams. Jaccard similarity,Bray–Curtis similarity, and Spearman rankcorrelations among samples collected from thesame sites over time were significantly greaterat undisturbed sites than at disturbed sites,suggesting greater fish assemblage persistenceand stability at the undisturbed sites. TheIndex of Biotic Integrity (IBI) also exhibitedsignificantly less variation over time atundisturbed sites than at disturbed sites.Physical habitat structure changed more overtime at disturbed sites than at undisturbedsites, and this variability was directlyrelated to temporal variability in fishassemblage structure. Comparisons betweenmultiple and single pass electrofishing samplessuggested that only a small proportion of thetemporal variability observed at the studysites was caused by inefficient sampling.Assessment of temporal variation in fishassemblage structure can serve as an indicatorof environmental disturbance and facilitate thedistinction of substantive ecological changefrom normal background variation.     

Benthic macroinvertebrate assemblages in north Swedish streams: environmental relationships

An analysis of the relationships between lotic macroinvertebrates and environmental variables was earned out on material from 60 riffle sites in streams in northern Sweden The approach involved the use of TWINSPAN classification and canonical correspondence analysis on presence/absence data from two seasons (spring and autumn) Variables most strongly associated with distribution patterns of assemblages were drainage area, elevation, water quality in terms of alkalinity, colour and phosphate and the presence of macrophytes The significance of affinities of different species to these variables are discussed The eight clusters resulting from the TWINSPAN analysis could biologically be interpreted as classes of taxa related to stream size, chemical conditions and algae A multiple regression analysis for predicting species nchness using three independent variables, viz drainage area amount or organic matter, and discharge was constructed The results of the study could be used as a starting point for further work on the community organization of benthie stream assemblages     

From compositional to functional biodiversity metrics in bioassessment: A case study using stream macroinvertebrate communities

While compositional diversity is a common metric for assessing human impacts on aquatic communities, functional diversity is scarcely employed, though highly desirable from the perspective of the European Water Framework Directive. Using abundance data from 99 minimally disturbed sites (i.e., no or very weak anthropogenic impact) from a national survey, we studied the spatial variability of compositional and functional biodiversity metrics across a predefined ecoregional classification. Metrics of compositional diversity comprised taxonomic and EPT richness and Simpson diversity. Functional diversity metrics were based on Rao's Quadratic Entropy (RQE), which described the differences among benthic invertebrate genera in eleven biological traits (e.g., size, life cycle, reproduction types, feeding habits). Using generalized linear models we show that taxonomic richness may vary greatly across ecoregions, contrasting with Simpson diversity and functional metrics that varied weakly in response to natural environmental variability. Functional diversity metrics, because of their stability in response to natural environmental variability, may be useful tools for assessing human impairment to ecosystem function. We further tested the response of functional diversity metrics to a specific human impact (sewage) and demonstrated significant modifications of functional diversity downstream of sewage pollution. Further investigations are required to test the ability of functional diversity metrics to precisely and accurately indicate different types of human impacts.     

Reproductive dynamics of a tropical freshwater crocodilian: Morelet's crocodile in northern Belize

Morelet's crocodile Crocodylus moreletii has not been well-studied and many aspects of its life history are unknown. In particular there is a notable paucity of information on nesting and reproductive ecology. We studied the nesting ecology of Morelet's crocodile in northern Belize from 1992 through 1995. Nesting occurs at the onset of the wet season in mid-June and continues through mid-July (mean oviposition date =1 July±10 days). Eggs hatch from mid-August through mid- to late September. Nesting effort at our primary study site remained relatively constant during 1992, 1993 and 1995, but nearly doubled in 1994; this appeared to reflect a regional trend. Natural and man-made islands are heavily used as nesting sites. Nesting success in 1993 and 1994 was consistently higher on natural islands when compared with man-made islands or shoreline sites. Nest losses were primarily due to flooding and raccoon Procyon lotor predation. Losses from predation were greatest in 1994 when unseasonably low water levels facilitated predator access to nests. Females probably reach sexual maturity in 7–8 years after attaining a total length of 150 cm. Mean clutch size (25.0±7.6; range=9–42; n =73) did not differ among years. Mean clutch size, egg width (EW), egg length, egg mass (EM) and clutch mass were positively correlated with female snout–vent length (SVL). Mean EW was the best predictor of female SVL. A partial correlation analysis of egg and clutch attributes found that independent of female SVL, EM increases with increasing clutch size.     

Comparing the sensitivity of diverse macroinvertebrate metrics to a multiple stressor gradient in Mediterranean streams and its influence on the assessment of ecological status

Biological indicators are increasingly being used as integrative measures of the ecosystem health in streams, particularly those using macroinvertebrate assemblage composition. Monitoring biological quality of rivers has not a long tradition in some Mediterranean European countries like Spain. Several macroinvertebrate metrics have been recently proposed to assess ecological status in Mediterranean streams, so it is necessary to compare the use of proposed biological quality metrics to select the most appropriate ones.In the present work, two classic richness metrics (total number of families and number of the Ephemeroptera, Plecoptera and Trichoptera families), three indices (IBMWP, IASPT and t-BMWQ) and two multimetric indices, recently proposed to be used in Mediterranean streams (ICM-9 and ICM-11a or IMMi-L), were compared by the analysis of the sensitivity of these metrics to a multiple stressor gradient which reflected the main pressures present in the study area. For this purpose, data from 193 sites sampled in spring (95 reference sites and 98 disturbed sites) belonging to five different Mediterranean stream types present in 35 basins were studied.The results showed that the adjusted regression coefficients (r²) for all seven metrics in the exponential regression models were higher than linear ones, thus indicating an exponential relationship between metrics and the environmental alteration. The two studied multimetric indices presented higher regression coefficients (r² = 0.590–0.669) than the three indices (r² = 0.524–0.574) and the two metrics (r² = 0.471–0.525), therefore showing a better response to a stressor gradient in Mediterranean streams. Within the multimetric indices group, ICM-11a showed the highest regression coefficients. Based on the results obtained, we suggest using the ICM-11a, apart from the IBMWP, to assess ecological status in Mediterranean streams.     

The origins of species richness in the Hymenoptera: insights from a family-level supertree

Background  

The order Hymenoptera (bees, ants, wasps, sawflies) contains about eight percent of all described species, but no analytical studies have addressed the origins of this richness at family-level or above. To investigate which major subtaxa experienced significant shifts in diversification, we assembled a family-level phylogeny of the Hymenoptera using supertree methods. We used sister-group species-richness comparisons to infer the phylogenetic position of shifts in diversification.     

Influence of a forest remnant on macroinvertebrate communities in a degraded tropical stream

Land use changes have resulted in large environmental impacts, and in agricultural landscapes sometimes only forest fragments remain. Riparian forest remnants can positively influence stream water quality, and serve as refuges for aquatic species. We evaluated whether the presence of a riparian forest remnant influenced the structure and composition of macroinvertebrate communities in a rural stream in southeastern Brazil. We sampled three reaches upstream (within abandoned sugarcane cultivation) and nine downstream the remnant edge, until 600 m inside the forested area, using leaf litter bags. The abundances of Elmidae, Chironomidae, and total macroinvertebrates increased along the forest remnant, whereas the abundance of Baetidae, proportion of Ephemeroptera, Plecoptera, and Trichoptera (EPT), rarefied taxonomic richness, and diversity decreased. Taxon richness and EPT abundance did not vary along the forest remnant. Increases in Chironomidae and total abundance within the forest remnant can be related to moderate increases in nutrient concentrations, or to the availability of high quality leaf litter patches. Forest remnants can influence macroinvertebrate communities, although variation both in temperate and tropical studies can be related to local agricultural practices and land use at the watershed scale. Forest remnants are important in maintaining stream water quality in rural landscapes, and deserve attention in watershed management projects.     

Microhabitat availability in Welsh moorland and forest streams as a determinant of macroinvertebrate distribution

SUMMARY. 1 Eighteen streams in mid-Wales were sampled for macro-invertebrates in both riffles and margins in April 1985–87. Stream macro-flora, substrata and marginal habitats were surveyed in May 1988.
2. TWINSPAN classification of the macroinvertebrate data indicated three major stream groups. One was distinguished by circumneutral pH and had a flora and fauna typical of such conditions. The other two groups consisted of acidic streams with moorland and conifer afforested catchments respectively. The forest streams were the more acidic but the two groups also differed significantly in the composition of their marginal habitats.
3 The acidic moorland streams had more vegetation ('soft' features) in the margins and supported several invertebrate taxa which were relative more abundant there than in the riffles. These taxa may be excluded from forest streams because the margins are 'hard' due to greater erosiveness and shading.
4. In view of the increasing cover by conifer afforestation in Britain, it is clearly necessary to elucidate all its effects on stream ecosystems, which include changes to the physical environment.     

Long-term recovery of macroinvertebrate biota in grossly polluted streams: Re-colonisation as a constraint to ecological quality

To meet targets imposed by the European Water Framework Directive (2000/60/EC) it is vital that measures to improve the status of rivers are both effective and economically viable. Achievement of such aims needs robust understanding of biological responses to changes in water quality vis-à-vis mechanisms of and constraints to the colonization of previously polluted sites. This study therefore examined the long-term chemical and biological changes in historically polluted rivers to elucidate the responses of macroinvertebrate biota to improvements in chemical water quality. For three historically polluted sites in the English Midlands, data from surveys over a period of ca. 50 years were analysed. Ammonia (NH₃) and 5-day biochemical oxygen demand (BOD₅) were used as chemical water quality indicators. Variations in the ecological recovery of the study sites were assessed using an average pollution sensitivity score (Average Score Per Taxon) and the number of taxa present (usually to family level) present in hand-net samples. Ecological recovery varied widely and was influenced by the intensity and spatial extent of the pollution and the proximity of available sources of potential colonisers. At the site most isolated from potential sources of colonizing taxa, no clean-water macroinvertebrate taxa were recorded 30 years after the major sources of pollution ceased. Where clean-water colonisers were more readily available, significant improvements in ecological quality followed within 2–5 years of the improvements in chemical quality. Macroinvertebrate communities and hence monitoring data may thus be indicative of long past conditions or of biological isolation rather than contemporaneous chemical conditions. Combined chemical and biological data were used to explore a generic model for predicting recovery rates and success. Neither BOD₅ nor NH₃ were found to provide a consistent and meaningful prediction of either average pollution tolerance of macroinvertebrate taxa or of the number of taxa present. Long-term relationships between macroinvertebrate variables and chemical water quality variables, however, were non-linear, suggesting that water quality thresholds may have to be exceeded before biological recovery can occur. Even when chemical water quality has been improved substantially, the apparent ecological status of macroinvertebrate communities may not reflect reduced pollution levels attained until adequate time to allow for re-colonisation (possibly decades) has elapsed.     

Catchment land cover as a proxy for macroinvertebrate assemblage structure in Carpathian Mountain streams

We compared land cover, riparian vegetation, and instream habitat characteristics with stream macroinvertebrate assemblages in 25 catchments in the Carpathian Mountains in Central Europe. This study area was particularly selected because of its diverse history of forest and agricultural ecosystems linked to geopolitical dynamic, which provide a suite of unique landscape scale, land cover settings in one ecoregion. Canonical Correspondence Analysis (CCA) showed that variation in composition and structure of macroinvertebrate assemblages was primarily related to four land cover types, and not to riparian or instream habitat. These were the portions in the catchment areas of (1) broadleaved forest, (2) fine-grained agricultural landscape mosaic with scattered trees (e.g., pre-industrial cultural landscape), (3) mixed forest, and (4) natural grassland without trees. Principal Component Analysis (PCA) suggested that land cover types and stream channel substrates co-varied. The PCA also showed that chemical variables, including organic carbon, had higher values in the agricultural landscape compared to natural forests. The major source of variation among taxa in streams was higher abundance of Diptera in agricultural landscapes and of Plecoptera, Coleoptera, Trichoptera, and Amphipoda in forests. Gastropoda and Oligochaeta were more abundant in open, fine-grained agricultural landscape mosaics with scattered trees. Ephemeroptera taxa were quite indifferent to these gradients in catchment land cover, but showed a tendency of being more abundant in the pre-industrial cultural landscape. Our findings suggest that land cover can be used as a proxy of the composition and structure of macroinvertebrate assemblages. This means that land use management at the catchment scale is needed for efficient conservation and recovery of stream invertebrate communities.     

Freshwater biodiversity at regional extent: determinants of macroinvertebrate taxonomic richness in headwater streams

We studied spatial variation of macroinvertebrate species richness in headwater streams at two spatial extents, within and across drainage systems, and assessed the relative importance of three groups of variables (local, landscape and regional) at each extent. We specifically asked whether the same variables proposed to control broad‐scale richness patterns of terrestrial organisms (temperature, topographic variability) are important determinants of species richness also in streams, or whether environmental factors effective at mainly local scales (in‐stream heterogeneity, potential productivity) constrain species richness in local communities. We used forward selection with two stopping criteria to identify the key environmental and spatial variables at each study extent. Eigenvector‐based spatial filtering was applied to evaluate spatial patterns in species richness, and variation partitioning was used to assess the amount of variation in richness attributable to purely environmental and spatial components. A prime regulator of richness variation at the bioregion extent was elevation range (increasing richness with higher topographic variability), whereas hydrological stability and temperature were unimportant. Water chemistry variables, particularly water color, exhibited strong spatially‐structured variation across drainage systems. Local environmental variables explained most of the variation in species richness at the drainage‐system extent, reflecting gradients in total phosphorus and water color (negative effect on richness). The importance of the pure spatial component was strongly region‐dependent, with a peak (60%) in one drainage system, suggesting the presence of unmeasured environmental factors. Our results emphasize the need for spatially‐explicit, regional studies to better understand geographical variation of freshwater biodiversity. Future studies need to relate species richness not only to local factors but also to broad‐scale climatic variables, recognizing the presence of spatially‐structured environmental variation.     

A hierarchical study of macroinvertebrate recolonization of disturbed patches along a longitudinal gradient in a prairie river

1. Macroinvertebrate densities and community composition were examined at three spatial scales after substratum disturbance; among reaches along a longitudinal gradient, within reaches and within plots. Reaches consisted of sandstone outcrops that were separated by approximately 2 km of highly mobile sandy silt substratum. 2. Substrata were disturbed by scraping sandstone plots (0.3 ± 0.3 m). Body-sized depressions created by Trichoptera in the sandstone were removed along with the upper 5 mm of sandstone, resulting in areas of newly exposed, smooth sandstone. 3. The spatial scale of examination determined whether patterns of macroinvertebrate distribution and densities were discernible. Initially there were no significant differences in community composition or total densities among reaches or among upstream/ downstream locations within reaches. Following substratum disturbance and 30 days recolonizarion, total macroinvertebrate densities did not differ significantly between undisturbed plots and disturbed plots. However, densities of Petrophilia (Pyralidae: Lepidoptera) differed along the longitudinal gradient and the Simuliidae had its highest density in the upstream reach. Significant differences were found in total macroinvertebrate densities between the upstream and the downstream halves of disturbed plots, with higher densities occurring in the downstream portions. 4. Recolonized plots had similar macroinvertebrate densities and community composition to undisturbed plots, suggesting that the stream community was highly resilient.     

Length–weight relationships for 36 fish populations from a disturbed tropical estuary in northern Colombia

The study reports length–weight relationships for 36 species belonging to 20 families of the estuarine zone of the Sinú River in northern Colombia. To the best knowledge of the authors, this study presents the first study on length–weight relationships for 10 species worldwide as well as seven new maximum lengths.     

Ticks from a Morelet's crocodile in Belize.

Parasitism of crocodilians by ticks has rarely been reported, and to our knowledge only seven published accounts exist. On 3 July 1999, we collected four ticks from a subadult Morelet's crocodile (Crocodylus moreletii) captured in northern Belize. These were identified as Amblyomma dissimile (one female), and Amblyomma sp. (two nymphs, one larva). The crocodile was captured on land approximately 100 m from water, and all four ticks were attached to loose skin on the lateral surface of the tail. Crocodilians are most susceptible to terrestrial ectoparasites, including ticks, during overland movements. However, most such movements occur in response to drought, when tick questing activity is suppressed, which likely accounts for the small numbers of tick specimens recorded from crocodilians and the absence of any noticeable impact of parasitism on host fitness.     

The effect of fine sedimentation on tropical stream macroinvertebrate assemblages: a comparison using flow-through artificial stream channels and recirculating mesocosms

In-situ artificial stream channels and ex-situ laboratory mesocosms were used to measure the responses of macroinvertebrate assemblages, from upland and lowland tropical streams, to high loads of fine clay sediment. Significant responses were observed mainly in the in-situ channels in the upland stream, where densities and the number of taxa were lower in the treatment channels than in controls. There was no evidence of any taxon being particularly sensitive to sedimentation, with a general decrease in densities across several taxa and differences only detectable for abundant taxa. Animals moved downstream in response to the treatment, but only a short distance within the channels. However, further colonization after the treatment was impeded in the treatment channels. In the mesocosm experiments, the upland macroinvertebrate assemblage demonstrated some negative effects; however, the lowland fauna was very tolerant to sedimentation, even when exposure was extended to 15 days. Together, the mesocosm and in-situ experiments indicate that there is a behavioural response to sedimentation because of a change in the habitat, and that the lowland macroinvertebrate assemblage is more tolerant of sedimentation, at least in the short term. Handling editor: D. M. Harper     

Genetic variation and social structure: a case-study from Africa

Understanding the relations between social and biological factors in determining the evolution of human populations is one of the principal aims of bio-anthropological research. The recent introduction of unilinearly transmitted polymorphisms of mitochondrial DNA and Y-chromosome has disclosed new possibilities for the study of bio-cultural evolutionary processes. In this communication, I present a study on the relationships between social structure and genetic variation in sub-Saharan Africa based on published and unpublished mitochondrial and Y-chromosomal data relative to a total of 40 populations. The results obtained reveal a striking difference in the genetic structure of farming (Bantu speakers) and hunting-gathering (Pygmies and Bushmen) populations. To explain this difference, I propose that asymmetric gene flow, polyginy and patrilocality, and hence the socio-cultural factors underlying them, have had an important role in determining and differentiating the genetic structure of sub-Saharan populations. The limits and the implications of this study are discussed.