Turnover and nestedness in subtropical dung beetle assemblages along an elevational gradient

Aim

We investigated changes in dung beetle β‐diversity components along a subtropical elevational gradient, to test whether turnover or nestedness‐related processes drive the dissimilarity of assemblages at spatial and temporal scales.

Location

An elevational gradient (200–1,600 m a.s.l.) of the Atlantic Forest in southern Brazil.

Methods

We investigated the extent to which β‐diversity varied along the elevational gradient (six elevations) at both spatial (among sites at different elevations) and temporal (different months at the same site) scales. We compared both the turnover and nestedness‐related dissimilarity of species and genera using multiple‐site or multiple‐month measures and tested whether these measurements were different from random expectations.

Results

A mid‐elevation peak in species richness along the elevational gradient was observed, and the lowest richness occurred at the highest elevations. We found two different groups of species, lowland and highland species, with a mixing of groups at intermediate elevations. The turnover component of β‐diversity was significantly higher for both spatial (i.e. elevational) and temporal changes in species composition. However, when the data for genera by site were considered, the elevational turnover value decreased in relative importance. Nestedness‐related processes are more important for temporal dissimilarity patterns at higher elevation sites.

Main conclusions

Spatial and temporal turnover of dung beetle species is the most important component of β‐diversity along the elevational gradient. High‐elevation assemblages are not subsets of assemblages that inhabit lower elevations, but this relationship ceases when β‐diversity is measured at the generic level. Environmental changes across elevations may be the cause of the differential establishment of distinctive species, but these species typically belong to the same higher taxonomic rank. Conservation strategies should consider elevational gradients in case‐specific scenarios as they may contain distinct species assemblages in lowlands vs. highlands.

     

Edge effects on dung beetle assemblages in an Andean mosaic of forest and coffee plantations

In landscapes dominated by agriculture, conspicuous edges often occur between landscape elements. However, there is disagreement about the existence and intensity of edge effects, and information about species‐specific responses remains scarce. Studying such edge effects can help elucidate functional landscape connectivity and contribute to agricultural management. We, therefore, assessed whether sun‐grown coffee represents a barrier to dung beetles in an Andean agricultural landscape. We also evaluated whether the response to edge effects differs among species. We found that diversity and abundance tend to decrease from forest to sun‐grown coffee and that there are sharp increases in species turnover at the forest–coffee edge. We detected several different species‐specific responses to the forest–coffee edge, suggesting differences in the mobility of the species (or spillover) and in the degree of penetration that takes place from forest patches to sun‐grown coffee plantations. This study demonstrates that the sun‐grown coffee matrix constitutes a barrier to forest species and suggests that the forest–coffee ecotone is more complex than expected. Our results support the notion that the conservation value of native forest patches in agricultural scenarios depends on the functional connectivity of forest units in the landscape to maximize the opportunities species have to disperse through the agricultural matrix.     

The role of habitat and daily activity patterns in explaining the diversity of mountain Neotropical dung beetle assemblages

The interaction between land use and climate change is expected to strongly affect species distributions along high elevation landscapes. We aimed to test the effect of climatic variables on community metrics among five types of land use in a high elevation landscape. We described dung beetle spatial and temporal taxonomic and functional diversity patterns, and partitioned β‐diversity into turnover and nestedness components. The interaction between land use and daily period of activity mostly drives abundance, functional richness and functional diversity, but not dung beetle species richness. Unlike Neotropical lowlands, species richness and abundance in open environments are similar to those existing in forests. Temperature is an important predictor of abundance and functional divergence. There is a higher spatial component of the taxonomic β‐diversity, which is highly driven by species turnover. The temporal component of the taxonomic β‐diversity was strongly driven by nestedness, where night assemblages are sub‐sets, although not entirely, of diurnal assemblages. For functional diversity, the temporal β‐diversity was much higher than the spatial β‐diversity, but both were similarly represented by functional group turnover and nestedness. The composition of nocturnal and diurnal assemblages is clearly different, even more than the differences observed between habitats. However, taxonomic turnover is the dominant force between sampling sites while nestedness dominates the daily pattern. This means that forest habitats are unlikely to act as shelters for grassland species under a scenario of rising temperature.     

Co‐declining mammal–dung beetle faunas throughout the Atlantic Forest biome of South America

The millennial–scale evolutionary relationships between mammals and dung beetles have been eroded due to several drivers of contemporary biodiversity loss. Although some evidence of co‐decline has been shown for mammals and dung beetles at some Neotropical sites, a biome‐scale analysis for the entire Atlantic Forest of South America would strengthen our understanding of how relictual sets of mammal species can affect dung beetle co‐occurrences and co‐declines. We therefore collated hundreds of assemblages of both dung beetles and medium‐ to large‐bodied mammals throughout the world's longest tropical forest latitudinal gradient to examine to what extent mammal assemblages may exert a positive influence on dung beetle species composition and functional assembly, and whether this relationship is scale dependent. We also collated several climatic and other environmental variables to examine the degree to which they shape mammal–dung beetle relationships. The relationships between local mammal and dung beetle faunas were examined using regression models, variation partitioning, dissimilarity indices and ecological networks. We found a clear positive relationship between mammal and dung beetle species richness across this forest biome, indicating an ongoing process of mammal–dung beetle niche‐mediated co‐decline. We found a strong relationship between the species composition of both taxa, in which dung beetle species dissimilarity apparently track changes in mammalian dissimilarity, typically in 80% of all cases. Co‐variables such as phytomass and climatic variables also influenced mammal–dung beetle patterns of co‐decline along the Atlantic Forest. We conclude that dung beetle diversity and community assembly are shaped by the remaining co‐occurring mammal assemblages and their functional traits, and both groups were governed by environmental features. We emphasize that ecosystem‐wide effects of mammal population declines remain poorly understood both quantitatively and qualitatively, and curbing large vertebrate defaunation will ensure the persistence of co‐dependent species.     

Stem respiration in tropical forests along an elevation gradient in the Amazon and Andes

Autotrophic respiration involves the use of fixed carbon by plants for their own metabolism, resulting in the release of carbon dioxide as a by‐product. Little is known of how autotrophic respiration components vary across environmental gradients, particularly in tropical ecosystems. Here, we present stem CO₂ efflux data measured across an elevation transect spanning ca. 2800 m in the Peruvian Amazon and Andes. Forest plots from five elevations were studied: 194, 210, 1000, 1500, and 3025 m asl Stem CO₂ efflux (R_s) values from each plot were extrapolated to the 1‐ha plot level. Mean R_s per unit stem surface area declined significantly with elevation, from 1.14±0.12 at 210 m elevation to 0.62±0.09 μmol C m⁻² s⁻¹ at 3025 m elevation. When adjusted for changing forest structure with elevation, this is equivalent to 6.45±1.12 Mg C ha⁻¹ yr⁻¹ at 210 m elevation to 2.94±0.19 Mg C ha⁻¹ yr⁻¹ at 3025 m elevation. We attempted to partition stem respiration into growth and maintenance respiration components for each site. Both growth and maintenance respiration rates per unit stem showed similar, moderately significant absolute declines with elevation, but the proportional decline in growth respiration rates was much greater. Stem area index (SAI) showed little trend along the transect, with declining tree stature at higher elevations being offset by an increased number of small trees. This trend in SAI is sensitive to changes in forest stature or size structure. In the context of rapid regional warming over the 21st century, such indirect, ecosystem‐level temperature responses are likely to be as important as the direct effects of temperature on maintenance respiration rates.     

Altitudinal variation of dung beetle (Scarabaeidae: Scarabaeinae) assemblages in the Colombian Andes

Aim We describe the changes in species richness, rarity and composition with altitude, and explore whether the differences in Scarabaeinae dung beetle composition along five altitudinal transects of the same mountain range are related to altitude or if there are interregional differences in these altitudinal gradients. Location Field work was carried out on the eastern slope of the eastern Cordillera, Colombian Andes, between Tamá Peak to the north, in the Tamá National Park (07°23′ N, 72°23′ W) and the San Miguel River (00°28′ N, 77°17′ W) to the south. Methods Sampling was carried out between February 1997 and November 1999 in five regions spanning elevation gradients. In each gradient, six sites were chosen at 250 m intervals between 1000 and 2250 m a.s.l. Results We found a curvilinear relationship between altitude and mean species richness, with a peak in richness at middle elevations. However, the diversity of dung beetle assemblages does not seem to be related to the interregional differences in environmental conditions. The number of geographically restricted species is negatively and significantly related to altitude, with geographically restricted species more frequent at low altitude sites. Ordination delimited the two main groups according to altitude: one with all the highest sites (1750–2250 m a.s.l.) and a second group with the remaining sites (< 1750 m a.s.l.). Analysis of species co‐occurrence shows that these dung beetle assemblages seem to be spatially structured when all sites have the same probability of being chosen. In contrast, the spatial structure of species assemblages seems to be random when the probability of choosing any site is proportional to its altitude. Main conclusions The altitude of sites is the main factor that influences the diversity of these dung beetle assemblages. The peak in species richness at middle elevations, the higher number of geographically restricted species at lower altitudinal levels, and the compositional differences along these mountain gradients seem to result from the mixing at these altitudes of dung beetle assemblages that have different environmental adaptations and, probably, different origins. The relevance of altitude in these assemblages is related to the limited role of these Neotropical high altitude environments as centres of refuge and vicariance for a monophyletic group of warm‐adapted species, for which the vertical colonization of these high mountain environments by lineages distributed at lower altitudes would have been very difficult.     

弄岗喀斯特季节性雨林不同群丛物种多样性随海拔的变化

为研究北热带喀斯特季节性雨林不同群丛物种多样性特征及其随相对海拔变化,在对广西弄岗15 hm2北热带喀斯特季节性雨林样地进行群丛划分的基础上,运用丰富度指数、物种多样性指数和均匀度指数对处于不同相对海拔高度群丛的物种多样性以及不同群丛乔木层、亚乔木层和灌木层的物种多样性进行比较。结果表明弄岗喀斯特季节性雨林各群丛的多样性指数存在极显著差异(P0.01),各群丛类型的物种多样性指数的顺序为:枝花流苏树-广西牡荆-蚬木群丛齿叶黄皮-黄梨木-蚬木群丛山石榴-广西牡荆-蚬木群丛南方紫金牛-苹婆-中国无忧花群丛垂茉莉-苹婆-广西棋子豆群丛割舌树-闭花木-蚬木群丛米仔兰-闭花木-蚬木群丛对叶榕-苹婆-董棕群丛;Patrick丰富度指数、Shannon多样性指数和Pielou均匀度指数在中坡以下(弄岗样地相对海拔76.3 m以下)随相对海拔变化规律符合"中间高度膨胀"理论,即处于该地段中部群丛的多样性高,相对海拔较低和较高的群丛的多样性低;而中坡以上(弄岗样地相对海拔76.3 m以上)多样性随相对海拔增高而增大;群丛各层次多样性与热带山地雨林一致,与亚热带喀斯特常绿阔叶林截然相反,呈现出乔木层亚乔木层灌木层。这些结果表明地形空间异质性导致的水热差异影响着北热带喀斯特季节性雨林物种多样性,相对海拔低、水热条件较好的中、下坡乔木多样性高,相对海拔较高、条件恶劣的中、上坡亚乔木、灌木多样性较高,所以在对喀斯特地区水热条件恶劣的中、上坡地带进行植被恢复时,应更多选择亚乔木、灌木。     

The importance of canopy complexity in shaping seasonal spider and beetle assemblages in saltmarsh habitats

1. Habitat structure, including vegetation structural complexity, largely determines invertebrate assemblages in semi‐natural grasslands. The importance of structural complexity to the saltmarsh invertebrate community, where the interplay between vegetation characteristics and tidal inundation is key, is less well known. 2. It was hypothesised that canopy complexity would be a more important predictor of spider and beetle assemblages than simple vegetation attributes (e.g. height, community type) and environmental variables (e.g. elevation) alone, measured in two saltmarsh regions, south‐east (Essex) and north‐west (Morecambe Bay) U.K. Canopy complexity (number of non‐vegetated ‘gaps’ in canopy ≥ 1 mm wide) was assessed using side‐on photography. Over 1500 spiders and beetles were sampled via suction sampling, winter and summer combined. 3. In summer, saltmarshes with abundant spider and beetle populations were characterised by high scores for canopy complexity often associated with tussocky grass or shrub cover. Simple vegetation attributes (plant cover, height) accounted for 26% of variation in spider abundance and 14% in spider diversity, rising to 46% and 41%, respectively, with the addition of canopy complexity score. Overwintering spider assemblages were associated with elevation and vegetation biomass. Summer beetle abundance, in particular the predatory and zoophagous group, and diversity were best explained by elevation and plant species richness. 4. Summer canopy complexity was identified as a positive habitat feature for saltmarsh spider communities (ground‐running hunters and sheet weavers) with significant ‘added value’ over more commonly measured attributes of vegetation structure.     

Phylogenetic structure and phylogenetic diversity of angiosperm assemblages in forests along an elevational gradient in Changbaishan,China

Aims Understanding what drives the variation in species composition and diversity among local communities can provide insights into the mechanisms of community assembly. Because ecological traits are often thought to be phylogenetically conserved, there should be patterns in phylogenetic structure and phylogenetic diversity in local communities along ecological gradients. We investigate potential patterns in angiosperm assemblages along an elevational gradient with a steep ecological gradient in Changbaishan, China.Methods We used 13 angiosperm assemblages in forest plots (32×32 m) distributed along an elevational gradient from 720 to 1900 m above sea level. We used Faith's phylogenetic diversity metric to quantify the phylogenetic alpha diversity of each forest plot, used the net relatedness index to quantify the degree of phylogenetic relatedness among angiosperm species within each forest plot and used a phylogenetic dissimilarity index to quantify phylogenetic beta diversity among forest plots. We related the measures of phylogenetic structure and phylogenetic diversity to environmental (climatic and edaphic) factors.Important findings Our study showed that angiosperm assemblages tended to be more phylogenetically clustered at higher elevations in Changbaishan. This finding is consistent with the prediction of the phylogenetic niche conservatism hypothesis, which highlights the role of niche constraints in governing the phylogenetic structure of assemblages. Our study also showed that woody assemblages differ from herbaceous assemblages in several major aspects. First, phylogenetic clustering dominated in woody assemblages, whereas phylogenetic overdispersion dominated in herbaceous assemblages; second, patterns in phylogenetic relatedness along the elevational and temperature gradients of Changbaishan were stronger for woody assemblages than for herbaceous assemblages; third, environmental variables explained much more variations in phylogenetic relatedness, phylogenetic alpha diversity and phylogenetic beta diversity for woody assemblages than for herbaceous assemblages.     

Drivers of Collembola assemblages along an altitudinal gradient in northeast China

Altitudinal changes in the diversity of plants and animals have been well documented; however, soil animals received little attention in this context and it is unclear whether their diversity follows general altitudinal distribution patterns. Changbai Mountain is one of few well‐conserved mountain regions comprising natural ecosystems on the Eurasian continent. Here, we present a comprehensive analysis of the diversity and community composition of Collembola along ten altitudinal sites representing five vegetation types from forest to alpine tundra. Among 7834 Collembola individuals, 84 morphospecies were identified. Species richness varied marginally significant with altitude and generally followed a unimodal relationship with altitude. By contrast, the density of Collembola did not change in a consistent way with altitude. Collembola communities changed gradually with altitude, with local habitat‐related factors (soil and litter carbon‐to‐nitrogen ratio, litter carbon content, and soil pH) and climatic variables (precipitation seasonality) identified as major drivers of changes in Collembola community composition. Notably, local habitat‐related factors explained more variation in Collembola assemblages than climatic variables. The results suggest that local habitat‐related factors including precipitation and temperature are the main drivers of changes in Collembola communities with altitude. Specifically, soil and litter carbon‐to‐nitrogen ratio correlated positively with Collembola communities at high altitudes, whereas soil pH correlated positively at low altitudes. This documents that altitudinal gradients provide unique opportunities for identifying factors driving the community composition of not only above‐ but also belowground invertebrates.     

Vertical stratification of feeding guilds and body size in beetle assemblages from an Australian tropical rainforest

Abstract The vertical stratification of insect species assemblages inhabiting tropical rainforests is well established but few have examined whether these patterns are reflected in vertical stratification of body size or feeding guilds. We used Malaise and Flight Interception Traps to sample beetle assemblages from five locations, at both canopy and ground zones of a tropical lowland rainforest site near Cape Tribulation, Australia. Beetles from 4 years of sampling were sorted to Family and morphospecies, and allocated to one of five feeding guilds. Within feeding guilds the number of species and individuals, from canopy‐ and ground‐caught traps were compared. The body lengths of species were measure and compared within feeding guilds and families. Herbivores was the dominant guild but was not the majority of all species or individuals. Most beetle species (69%) were less than 5 mm in length and the mean size of canopy‐caught species was greater than that for ground‐caught species. This was probably due to slightly more species of plant feeders (herbivores and xylophages) present in the canopy, which were significantly larger than saprophages, fungivores and predators. Among feeding guilds, there were few overall canopy–ground differences. These results contrast with species composition results presented elsewhere where strong differences between the canopy and the ground were evident. We suggest that our guild groupings may have been too coarse to detect fine‐scale differences and that resource partitioning may have also masked faunal stratification. We propose that fine‐scale differences in resources between the canopy and the ground, together with strong microclimate gradients, are likely to be important in structuring the vertical stratification of insect assemblages at the level of species, but not with respect to functional groups.     

High levels of mercury contamination in a necrophage dung beetle of the Amazon rainforest (Scarabaeidae: Scarabaeinae)

This study represents the first evidence of mercury contamination in the family Scarabaeidae, with a close focus on Coprophanaeus lancifer, the largest copro-necrophagous beetle in South America. This work shows the repartition of total mercury (THg) in the insect body and lays the groundwork for additional future studies. Abstract in Spanish is available with online material.     

Influence of Edge Exposure on Tree Seedling Species Recruitment in Tropical Rain Forest Fragments1

Edge creation has a pronounced influence on the understory vegetation, but the effects of edges on seedling species recruitment are still poorly understood. In Central Amazonia, 9–19 years after fragmentation, we recorded species richness and net seedling recruitment rate in 1 ha blocks exposed to none, one, or multiple edges within forest fragments. One‐hectare blocks were located in the center (no edge), the edge (one edge), the corners (two edges) of 10 and 100 ha fragments, and in a 1 ha fragment (four edges). In 1991, we counted all tree seedlings 5–100 cm tall found within permanent 1 m² plots located within the 1 ha blocks. In May 1993, we manually removed all seedlings that were smaller than 1 m tall from the permanent plots. Six years and five months later (October 1999), all new seedlings recruited into the plots were counted and classified into distinct morphospecies. Species richness of recruited seedlings, scaled by total seedling density, declined from the center to the edge, the corner blocks, and then to the 1 ha fragment. Overall, the four‐edged, 1 ha fragment had the poorest species richness and the non‐edged 100 ha central block the highest. The total number of recruited individuals was 40 percent less than that previously present, with the 100 ha corner having the lowest recruitment. Pairwise comparisons showed that species similarity was related to edge number for the 100 and 1 ha fragments. Species rank/abundance curves showed that a subset of species was common in all blocks within the fragments, and that the 100 ha center held more rare species than any other 1 ha block. This study demonstrated that, in a given fragment patch, the number of tree seedling species recruited varied inversely with the number of edges.     

Mammalian species richness and morphological complexity along an elevational gradient in the arid south-west

Aim We examined the relationship between species richness and morphological complexity of terrestrial mammal communities along an elevational gradient. Location The gradient is in the Sonoran Desert in Southern California and extends from a sand dune habitat near sea level to coniferous forest ending at >2600 m. Methods Morphological diversity, characterized by both size and shape of coexisting mammal species, was estimated within and between sites from projections of variables on principal components axes. Similarities among species were calculated as Euclidean distances. To tease apart size and shape, we constructed two principal component analyses: one based on log-transformed original measurements, the other on log-transformed proportional shape variables. To test whether species number accounted for the morphological diversity at each site we designed two null models. The models generated were random communities generated from the forty-two-species pool. Indices of morphological diversity for real communities were compared with the results of 500 simulations of each null model. Results Species richness varied along the gradient, peaking in the mid-elevation agave-ocotillo habitat. Morphological diversity of shapes and sizes correlated strongly with species richness. Locomotor, tooth, and skull traits were all important in distinguishing among species. Main conclusions Two important patterns emerged: (1) diversity of both sizes and of shapes of species within communities correlated positively with species number, and both sets of variables behaved similarly across this gradient; (2) the most species rich sites were not composed of specialists on these best places, but rather, a community of species derived from overlapping faunal groups.     

Species turnover, community boundaries and biogeographical composition of dung beetle assemblages across an altitudinal gradient in South Africa

Aim To identify biogeographical boundaries which are obscured by faunal overlap and habitat modification. Location KwaZulu-Natal in south-east, South Africa beyond the southern tip of the Moçambique Coastal Plain. Methods Species abundance data for dung beetles (Scarabaeidae: Scarabaeinae) were collected at six levels from the coastal escarpment (30°16′S – 500 m) to the top of the nearby Drakensberg (29°35′S – 2850 m). Cross-altitudinal boundaries were identified using clustering techniques, beta-diversity indices, and range edge analysis. Biogeographical data for the species were drawn from an extensive reference collection and used to classify the biogeographical affinities of the assemblages. Results Three discrete communities are defined (<10% similarity) from species abundance distributions. These communities occur in coastal forest (500 m), coastal to highveld grassland (500–1500 m), and montane grassland (1900–2850 m). Two of these communities are biogeographically homogeneous comprising >89% east coast endemics (coastal forest) or >84% South African montane endemics (montane grassland) in terms of abundance. The third community in coastal to highveld grassland is biogeographically more heterogeneous. Predominant biota of this community comprise both South African highveld endemics and elements with distributions extending into the tropics. At highveld levels (1500 m), there are proportionately more highveld endemics whereas at lowland levels (500 m), there are proportionately more tropical elements. At 1000 m, there was a change in the balance between these two groups across an anthropogenic gradient due to a decline in the proportion of endemics in favour of temperate/tropical generalists. This gradient from a natural grassland fragment to improved pastures of Kikuyu grass also parallels a decline in species richness and abundance. Species turnover analyses showed three different cross-altitudinal patterns. Range-edge analysis showed a trimodal pattern of species turnover (peaks in forest and the Drakensberg foothills as in the community analysis but also at 1000 m). Five beta-diversity indices showed either a bimodal pattern of turnover (forest/grassland and foothills/middle Drakensberg slopes) or a trimodal turnover pattern (forest/grassland, highveld/Drakensberg foothills, Drakensberg peaks). Main conclusions Clear altitudinal zonation is revealed by community and biogeographical analysis but one natural biogeographical boundary may be obscured by the process of habitat modification. This boundary at 1000 m is revealed by range-edge analysis and is supported by findings for plant communities. Beta-diversity, species turnover patterns diverged slightly from those suggested by the community and range-edge analyses.     

Secondary Succession and Indigenous Management in Semideciduous Forest Fallows of the Amazon Basin1

To the discussion on secondary succession in tropical forests, we bring data on three under‐addressed issues: understory as well as overstory changes, continuous as opposed to phase changes, and integration of forest succession with indigenous fallow management and plant uses. Changes in vegetation structure and species composition were analyzed in secondary forests following swidden agriculture in a semideciduous forest of Bolivian lowlands. Twenty‐eight fallows, stratified by four successional stages (early = 1–5 yr, intermediate = 6–10 yr, advanced = 12–20 yr, and older = 22–36 yr), and ten stands of mature forests were sampled. The overstory (plants ≥5 cm diameter at breast height [DBH]) was sampled using a 20 × 50 m plot and the understory (plants <5 cm DBH) in three nested 2 × 5 m subplots. Semistructured interviews provided information on fallow management. Canopy height, basal area, and liana density of the overstory increased with secondary forest age. The early stage had the lowest species density and diversity in the overstory, but the highest diversity in the understory. Species composition and abundance differentiated mature forests and early successional stage from other successional stages; however, species showed individualistic responses across the temporal gradient. A total of 123 of 280 species were useful with edible, medicinal, and construction plants being the most abundant for both over‐ and understories. Most of Los Gwarayo preferred mature forests for making new swidden, while fallows were valuable for crops, useful species, and regenerating timber species.     

Relationships between fish assemblages, macrophytes and environmental gradients in the Amazon River floodplain

During the flood season of 1992–1993, 139 species of fishes were collected from a floodplain lake system in the central Amazon Basin. Fish species distribution was examined relative to abiotic variables in seven vegetation strata on Marchantaria Island, Solimões River. Both environmental variables and species distributions were influenced by a river channel to floodplain-interior gradient. Species diversity was significantly higher in vegetated areas than in unvegetated areas, with deeper water Paspalum repens stands harbouring the highest diversity. As a result, species richness and catches were positively related to habitat complexity, while catch was also negatively related to dissolved oxygen (DO) and water depth. Low DO and shallow waters appeared to act as a refuge from predation. Fish assemblages were related to water chemistry, but species richness was not. Canonical correspondence analysis provided evidence that floodplain fish assemblages formed by the 76 most common species were influenced by physical variables, macrophyte coverage and habitat complexity, which jointly accounted for 67% of the variance of fish species assemblages. Omnivores showed no pattern relative to the river channel to floodplain-interior gradient while detritivores were more likely to be found at interior floodplain sites and piscivores closer to the river. Piscivores could be further separated into three groups, one with seven species associated with free-floating macrophytes in deep water, a second with five species found in shallow waters with rooted grasses and a third with six open water orientated species. The results suggest that fish assemblages in the Amazon floodplain are not random associations of species.     

Variation in freshwater fish assemblages along a regional elevation gradient in the northern Andes,Colombia

Studies on elevation diversity gradients have covered a large number of taxa and regions throughout the world; however, studies of freshwater fish are scarce and restricted to examining their changes along a specific gradient. These studies have reported a monotonic decrease in species richness with increasing elevation, but ignore the high taxonomic differentiation of each headwater assemblage that may generate high β‐diversity among them. Here, we analyzed how fish assemblages vary with elevation among regional elevation bands, and how these changes are related to four environmental clines and to changes in the distribution, habitat use, and the morphology of fish species. Using a standardized field sampling technique, we assessed three different diversity and two structural assemblage measures across six regional elevation bands located in the northern Andes (Colombia). Each species was assigned to a functional group based on its body shape, habitat use, morphological, and/or behavioral adaptations. Additionally, at each sampling site, we measured four environmental variables. Our analyses showed: (1) After a monotonic decrease in species richness, we detected an increase in richness in the upper part of the gradient; (2) diversity patterns vary depending on the diversity measure used; (3) diversity patterns can be attributed to changes in species distribution and in the richness and proportions of functional groups along the regional elevation gradient; and (4) diversity patterns and changes in functional groups are highly correlated with variations in environmental variables, which also vary with elevation. These results suggest a novel pattern of variation in species richness with elevation: Species richness increases at the headwaters of the northern Andes owing to the cumulative number of endemic species there. This highlights the need for large‐scale studies and has important implications for the aquatic conservation of the region.