Pleistocene Extinctions and the Perceived Morphofunctional Structure of the Neotropical Felid Ensemble

The felid Neotropical ensemble has experienced important changes from the Pleistocene to the present, the extinction of sabertooth cats being the most significant. Assuming that the morphofunctional structure of the Neotropical felid ensemble was maximally expressed when machairodontines were present, we added specimens of Smilodon fatalis and S. populator to a morphological dataset inclusive of all extant species of Neotropical felids to explore the hypothetical effect of extinction of machairodontines on the perceived morphofunctional structure of the ensemble. We studied 321 specimens of 12 extant species of Neotropical felids plus S. populator and S. fatalis by measuring 123 skull and postcranial morphofunctional variables. We used Principal Component Analysis to find morphofunctional patterns of the skull, postcranium, and both combined (with and without correction for size) of both ensembles, past (with Smilodon) and present (with Smilodon lost to extinction). Canonical Phylogenetic Ordination was performed to assess the degree of phylogenetic influence on this morphospace. We also explored the effect of including homotherines in the Pleistocene guild by including specimens of North American Homotherium serum. Size was the principal factor structuring ensembles. Important morphological characters were associated to ecological performance of species. A pattern of bodily proportions was apparent: more stout species with larger skull, longer rostrum, stronger bite force, and longer pectoral crests, versus gracile, jumping species with opposing characters. The felid morphospace shrank after the extinction of Smilodon and Homotherium, but the configuration of the portion of space containing extant felids remained stable. This pattern is associated with deep phylogenetic roots.     

The Evolution of the Cenozoic Terrestrial Mammalian Predator Guild in South America: Competition or Replacement?

South America was isolated from other continents during most of the Cenozoic, developing a singular mammalian fauna. In contrast to North America, Europe, Asia, and Africa, up to the late Neogene, the carnivore adaptive zone in South America was populated by crocodiles (Sebecidae), large snakes (Madtsoiidae), large birds (Phorusrhacidae), and metatherian mammals (Sparassodonta). Sparassodonta were varied and comprised a wide range of body masses (≈ 2–50 kg) and food habits. Their diversity decreased towards the late Miocene (Huayquerian Stage/Age) and the group became extinct in the “middle” Pliocene (≈ 3 Ma, Chapadmalalan Stage/Age). Several authors have suggested that the cause of this decline and extinction was the ingression of carnivorans to South America (about 6–7 Ma ago), because they competed with the Sparassodonta; although this hypothesis has been criticized in recent years. With the intention of testing the hypothesis of “competitive displacement,” we review the fossil record of South American Sparassodonta and Carnivora, collect data about diversity, estimate size and diet, and determine first and last appearances. The diversity of Sparassodonta is low relative to that of Carnivora throughout the Cenozoic with the early Miocene (Santacrucian Stage/Age) showing the greatest diversity with 11 species. In the late Miocene-middle Pliocene (Huayquerian Stage/Age), the fossil record shows overlap of groups, and the Sparassodonta’s richness curve begins to decline with the first record of Carnivora. Despite this overlap, carnivorans diversity ranged from four or fewer species in the late Miocene-Pliocene to a peak of around 20 species in the early Pleistocene (Ensenadan Stage/Age). Carnivora was initially represented by small-sized, omnivorous species, with large omnivores first appearing in the Chapadmalalan Stage/Age. Over this period, Sparassodonta was represented by large and small hypercarnivores and a single large omnivorous species. From this review of the fossil record, it is suggested that factors other than competitive displacement may have caused the extinction of the Sparassodonta.     

Clonal ability,height and growth form explain species’ response to habitat deterioration in Fennoscandian wooded meadows

Fennoscandian calcareous wooded meadows have high conservation value due to very high diversity which has been maintained by consistent mowing, but undergo species loss when this management is abandoned. We compared species richness and composition of regularly mown and abandoned wooded meadows in Estonia and established species groups with respect to their response to abandonment. These meadows were very species rich with a maximum of 43 species per 0.25 m². Species whose populations are maintained by mowing constituted >60 % of the floristic diversity of the mown wooded meadows. Abandonment suppressed species with a preference for dry infertile open habitats. Response to abandonment was related to clonality, height and growth form. The greatest negative response was associated with low graminoids, short-lived and non-clonal species, rosette and semi-rosette growth form. Weak competitors with low height and light seeds exhibited a slow decline; perennial life span and clonality enable a delay in local extinction of these species. Response groups had similar persistence at a national level, revealing that earlier assessment of species dynamics at larger spatial scales could underestimate the number of threatened species. Application of restoration measures could avoid local extinctions of many mowing-supported species, whereas some of the mowing-dependent species likely need reintroduction.     

Tree species richness decreases while species evenness increases with disturbance frequency in a natural boreal forest landscape

Understanding species diversity and disturbance relationships is important for biodiversity conservation in disturbance‐driven boreal forests. Species richness and evenness may respond differently with stand development following fire. Furthermore, few studies have simultaneously accounted for the influences of climate and local site conditions on species diversity. Using forest inventory data, we examined the relationships between species richness, Shannon''s index, evenness, and time since last stand‐replacing fire (TSF) in a large landscape of disturbance‐driven boreal forest. TSF has negative effect on species richness and Shannon''s index, and a positive effect on species evenness. Path analysis revealed that the environmental variables affect richness and Shannon''s index only through their effects on TSF while affecting evenness directly as well as through their effects on TSF. Synthesis and applications. Our results demonstrate that species richness and Shannon''s index decrease while species evenness increases with TSF in a boreal forest landscape. Furthermore, we show that disturbance frequency, local site conditions, and climate simultaneously influence tree species diversity through complex direct and indirect effects in the studied boreal forest.     

The Structure of the Mammalian Predator Guild in the Santa Cruz Formation (Late Early Miocene)

The Santa Cruz Formation (late early Miocene, Santacrucian age) registers 11 species of mammalian predators (Metatheria, Sparassodonta). Together with large carnivorous flightless birds, they comprised the terrestrial predator guild. The Santacrucian sparassodonts were diverse in body size, had different locomotory habits, and were primarily hypercarnivores. The objective of this work is to analyze the guild structure of the sparassodonts of the Santa Cruz Formation, using the variables of body mass, diet, and locomotion as proxies. Furthermore, we analyze the interaction with other predators and potential prey. The univariated test V of Poole and Rathcke and the multivariated test of Clark-Evans were used to construct the models. In the multivariate test, we made a Principal Component Analysis to resume and standardize the variables. With body mass and locomotion we obtained an evenly spaced pattern of segregation for the sparassodont species, being non-significant and significant, respectively. The pattern was aggregated and significant only with diet. The analysis of all variables together resulted in an evenly spaced and significant pattern, which is consistent with character displacements (segregation of species throughout the morphospace) that would help to diminish interspecific competition during the Santacrucian age and would allow selection of prey species of different sizes and substrate specializations. When the body size pattern of predator birds and sparassodonts were plotted together, the pattern is evenly spaced and non-significant. Other factors, including locomotion, would differentiate these species and their ecological niches.     

Geometric morphometric analysis of the freshwater prawn Macrobrachium borellii (Decapoda: Palaemonidae) at a microgeographical scale in a floodplain system

Morphometric studies are useful for delineating the shapes of various populations and species over geographical ranges and as evidence of regional differences in crustaceans. Hydrological fluctuations in a floodplain system modulate the dispersal and presence of decapods among habitats and constitute an important macrofactor that regulate other environmental variables and which could explain the richness, distribution and abundance of organisms that live in these systems. Morphological variations among populations of the freshwater prawn Macrobrachium borellii in a floodplain system at a microgeographical scale were studied using geometric morphometrics. Carapace structure was represented using 16 digitised landmarks. Allometry and sexual dimorphism was tested. Variation in shape was explored via Principal Component Analysis. Canonical Variate Analyses was applied to compare the differences in shape between species’ populations. The correlations and covariations among shapes and hydrometric level, current velocity, geographical location and hydrologic distances were analysed. The average carapace shape was different between sexes in all sites. Populations that were near each other in terms of hydrological distance had similar shapes, but all of the populations differed in shape from the farthest population. The environmental variables were not good predictors of the carapace shape. Instead, the shape was strongly related to the hydrologic distance and geographical location. The swimming characteristics of these prawns and their passive movements, together with the dynamics of a floodplain system, explain the low morphological variation between populations in this study. The dynamic characteristics of the system influence the dispersal of the prawns and allow populations to remain connected.     

Bacterial Community Assemblages Associated with the Phyllosphere,Dermosphere, and Rhizosphere of Tree Species of the Atlantic Forest are Host Taxon Dependent

Bacterial communities associated with tree canopies have been shown to be specific to their plant hosts, suggesting that plant species-specific traits may drive the selection of microbial species that comprise their microbiomes. To further examine the degree to which the plant taxa drive the assemblage of bacterial communities in specific plant microenvironments, we evaluated bacterial community structures associated with the phyllosphere, dermosphere, and rhizosphere of seven tree species representing three orders, four families and four genera of plants from a pristine Dense Ombrophilous Atlantic forest in Brazil, using a combination of PCR-DGGE of 16S rRNA genes and clone library sequencing. Results indicated that each plant species selected for distinct bacterial communities in the phyllosphere, dermosphere, and rhizosphere, and that the bacterial community structures are significantly related to the plant taxa, at the species, family, and order levels. Further characterization of the bacterial communities of the phyllosphere and dermosphere of the tree species showed that they were inhabited predominantly by species of Gammaproteobacteria, mostly related to Pseudomonas. In contrast, the rhizosphere bacterial communities showed greater species richness and evenness, and higher frequencies of Alphaproteobacteria and Acidobacteria Gp1. With individual tree species each selecting for their specific microbiomes, these findings greatly increase our estimates of the bacterial species richness in tropical forests and provoke questions concerning the ecological functions of the microbial communities that exist on different plant parts.     

Local extinction of a coral reef fish explained by inflexible prey choice

While global extinctions of marine species are infrequent, local extinctions are becoming common. However, the role of habitat degradation and resource specialisation in explaining local extinction is unknown. On coral reefs, coral bleaching is an increasingly frequent cause of coral mortality that can result in dramatic changes to coral community composition. Coral-associated fishes are often specialised on a limited suite of coral species and are therefore sensitive to these changes. This study documents the local extinction of a corallivorous reef fish, Oxymonacanthus longirostris, following a mass bleaching event that altered the species composition of associated coral communities. Local extinction only occurred on reefs that also completely lost a key prey species, Acropora millepora, even though coral cover remained high. In an experimental test, fish continued to select bleached A. millepora over the healthy, but less-preferred prey species that resisted bleaching. These results suggest that behavioural inflexibility may limit the ability of specialists to cope with even subtle changes to resource availability.     

Temporal variation of phytophilous Chironomidae over a 11-year period in a shallow Neotropical lake in southern Brazil

Small and shallow lakes make up most of the lake-covered area in the world, and immature Chironomidae generally constitute the dominant group among invertebrates in these systems, exhibiting a wide distribution and tolerance to various levels of water quality. The changes in trophic features associated with the dominance of different major primary producers (macrophyte and phytoplankton) are reflected in the functioning of these ecosystems and likely in their phytophilous Chironomidae species richness and abundance. These were thus studied in a Brazilian shallow lake over a 11-year period characterized by both clear and turbid water conditions in relation to water quality and dominant macrophytes. The results revealed differences in Chironomidae composition but not in richness associated with clear and turbid waters. The Goeldichironomus genera predominated during all years and for different macrophyte types (freefloating, floating leaf, and submerged). We attribute the variability of Chironomidae assemblages to changes in macrophyte dominance and water trophic status, such as the concentration of nutrients and chlorophyll-a, during the 11-year period. Submerged macrophytes were associated with clear waters and harbored the highest diversity, which emphasized the importance of these environmental conditions for preserving the biodiversity of Chironomidae in shallow lakes.     

Elevational gradients in bird diversity in the Eastern Himalaya: an evaluation of distribution patterns and their underlying mechanisms

Background

Understanding diversity patterns and the mechanisms underlying those patterns along elevational gradients is critically important for conservation efforts in montane ecosystems, especially those that are biodiversity hotspots. Despite recent advances, consensus on the underlying causes, or even the relative influence of a suite of factors on elevational diversity patterns has remained elusive.

Methods and Principal Findings

We examined patterns of species richness, density and range size distribution of birds, and the suite of biotic and abiotic factors (primary productivity, habitat variables, climatic factors and geometric constraints) that governs diversity along a 4500-m elevational gradient in the Eastern Himalayan region, a biodiversity hotspot within the world''s tallest mountains. We used point count methods for sampling birds and quadrats for estimating vegetation at 22 sites along the elevational gradient. We found that species richness increased to approximately 2000 m, then declined. We found no evidence that geometric constraints influenced this pattern, whereas actual evapotranspiration (a surrogate for primary productivity) and various habitat variables (plant species richness, shrub density and basal area of trees) accounted for most of the variation in bird species richness. We also observed that ranges of most bird species were narrow along the elevation gradient. We find little evidence to support Rapoport''s rule for the birds of Sikkim region of the Himalaya.

Conclusions and Significance

This study in the Eastern Himalaya indicates that species richness of birds is highest at intermediate elevations along one of the most extensive elevational gradients ever examined. Additionally, primary productivity and factors associated with habitat accounted for most of the variation in avian species richness. The diversity peak at intermediate elevations and the narrow elevational ranges of most species suggest important conservation implications: not only should mid-elevation areas be conserved, but the entire gradient requires equal conservation attention.     

Middle Ottnangian (late Burdigalian) mollusks from the Rott Valley (SE Germany): the ultimate marine fauna of the Western Paratethys

The Bivalvia and Gastropoda from the Upper Marine Molasse deposits of the lower Rott Valley (North Alpine Foreland Basin, Lower Bavaria, SE Germany) comprise the first mollusk fauna of middle Ottnangian age (ca 17.5 Ma) to be documented in detail. Altogether, the assemblages from Brombach, Kainerding and Anzenkirchen yielded 27 species of Bivalvia and 9 species of Gastropoda. The bivalve species Spisula brombachensis n. sp. is described as new to science, and Aequipecten camaretensis is first reported from the Ottnangian of the North Alpine Foreland Basin. Furthermore, the well-preserved material allowed for the taxonomic revision of the Ottnangian index fossil Pecten herrmannseni. The bivalve community is dominated by suspension feeders, mostly shallow, soft bottom dwellers (Glycymeris, Anadara) and cementing epifauna (Ostrea). Gastropods almost exclusively comprise carnivorous browsers and predators. The parautochthonous assemblages are indicative of normal marine, well-oxygenized sandy-silty shoals at a water depth of presumably less than 20 m. Except for the presence of a single estuarine Crassostrea, the middle Ottnangian fauna is not indicative of reduced salinities. Consequently, the paleoenvironmental turnover to the brackish-water fauna of the directly overlying upper Ottnangian deposits of the Rzehakia Lake was abrupt. Glycymeris–Anadara–Ostrea communities similar to the one recorded from the Rott Valley are widespread in European seas and beyond during the Neogene and Quaternary.     

Performance of two alpine plant species along environmental gradients in an alpine meadow ecosystem in central Tibet

Understanding how biotic interactions and abiotic conditions affect plant performance is important for predicting changes in ecosystem function and services in variable environments. We tested how performances of Astragalus rigidulus and Potentilla fruticosa change along gradients of biotic interactions (represented by plant species richness, abundance of the dominant plant species Kobresia pygmaea, and herbivory intensity) and abiotic conditions (represented by elevation, aspect, and slope steepness) across a semi-arid landscape in central Tibet. Redundancy analyses showed that the biotic variables explained 30 and 39 % of the variation in overall performance of A. rigidulus (P = 0.03) and P. fruticosa (P = 0.01), respectively. Abiotic variables did not contribute significantly to variation among A. rigidulus populations. Plant size decreased with species richness in both species and was larger on south- rather than north-facing slopes. Reproductive effort for both species was significantly negatively related to the abundance of K. pygmaea and both species had larger reproductive effort on south- rather than north- and west-facing slopes. The proportion of biomass allocated to sexual reproduction in P. fruticosa was negatively correlated with K. pygmaea abundance and herbivory intensity. The population density of P. fruticosa was positively related to elevation, species richness, and K. pygmaea abundance. We conclude that plant performance at a local scale was more strongly related to biotic than abiotic conditions, but different components of plant performance responded differently to predictor variables and the responses were species-specific. These findings have important implications for rangeland management under changing environmental conditions.     

Quantitative historical change in bumblebee (Bombus spp.) assemblages of red clover fields

Background

Flower visiting insects provide a vitally important pollination service for many crops and wild plants. Recent decline of pollinating insects due to anthropogenic modification of habitats and climate, in particular from 1950''s onwards, is a major and widespread concern. However, few studies document the extent of declines in species diversity, and no studies have previously quantified local abundance declines. We here make a quantitative assessment of recent historical changes in bumblebee assemblages by comparing contemporary and historical survey data.

Methodology/Principal Findings

We take advantage of detailed, quantitative historical survey data from the 1930''s on bumblebee (Bombus spp.) abundances and species composition in red clover (Trifolium pratense) fields, an important floral resource and an attractant of all bumblebee species. We used the historical survey data as a pre-industrialization baseline, and repeated the same sampling protocol at nearly the same localities at present, hence setting up a historical experiment. We detected historical changes in abundances (bees/m²) of both workers (the “pollinatory units”) and queens (effective population size), in addition to species composition. In particular, long-tongued bumblebee species showed consistent and dramatic declines in species richness and abundances throughout the flowering season of red clover, while short-tongued species were largely unaffected. Of 12 Bombus species observed in the 1930''s, five species were not observed at present. The latter were all long-tongued, late-emerging species.

Conclusions/Significance

Because bumblebees are important pollinators, historical changes in local bumblebee assemblages are expected to severely affect plant reproduction, in particular long-tubed species, which are pollinated by long-tongued bumblebees.     

Composition and species diversity of pine-wiregrass savannas of the Green Swamp,North Carolina

Fire-maintained, species-rich pine-wiregrass savannas in the Green Swamp, North Carolina were sampled over their natural range of environmental conditions and fire frequencies. Species composition, species richness, diversity (Exp H′, 1/C), and aboveground production were documented and fertilization experiments conducted to assess possible mechanisms for the maintenance of high species diversity in these communities. Although savanna composition varies continuously, DECORANA ordination and TWINSPAN classification of 21 sites facilitated recognition of 3 community types: dry, mesic, and wet savannas. These savannas are remarkably species-rich with up to 42 species/0.25 m² and 84 species/625 m². Maximum richness occurred on mesic, annually burned sites. Aboveground production, reported as peak standing crop, was only 293 g · m^?2 on a frequently burned mesic savanna but was significantly higher (375 g · m^?2) on an infrequently burned mesic site. Production values from fertilized high and low fire frequency sites were equivalent. Monthly harvest samples showed that savanna biomass composition by species groups did not vary seasonally, but within groups the relative importance of species showed clear phenological progressions. The variation in species richness with fire frequency is consistent with non-equilibrium theories of species diversity, while phenological variation in production among similar species and the changing species composition across the moisture gradient suggest the importance of equilibrium processes for maintenance of savanna diversity.     

Species’ traits influence ground beetle responses to farm and landscape level agricultural intensification in Europe

Agricultural intensification may result in important shifts in insect community composition and function, but this remains poorly explored. Studying how groups of species with shared traits respond to local and landscape scale land-use management can reveal mechanisms behind such observed impacts. We tested if ground beetles (Coleoptera: Carabidae) divided into trait groups based on body sizes, wing morphologies and dietary preferences respond differently to farming practise (organic and conventional), farming intensity (measured as yield) and landscape complexity (measured as the proportion of arable land within a 1,000 m radius) across Europe. We used data from 143 farms in five regions in northern and central Europe. Organic farms did not differ in abundance or richness of any trait group compared to conventional farms. As farm scale intensity (yield) increased, overall abundance of beetles decreased, but abundances of small and medium sized beetles, as well as that of wingless beetles, were unaffected. Overall species richness was not affected by yield, whereas consideration of traits revealed that phytophagous and omnivorous beetles were less species rich on farms with high yields. Increasing the proportion of arable land in the landscape increased overall beetle abundance. This was driven by an increase in omnivorous beetles. The total species richness was not affected by an increase in the proportion arable land, although the richness of wingless beetles was found to increase. Potential effects on ecosystem functioning need to be taken into account when designing schemes to maintain agricultural biodiversity, because species with different ecological traits respond differently to local management and landscape changes.     

Invertebrate communities delineate hydro-ecoregions and respond to anthropogenic disturbance in East-Amazonian streams

Many tropical regions lack models predicting the biological and environmental conditions expected in any given area, thus precluding the implementation of reference condition-based water policies. We focused on streams of French Guiana, and tested two predictions: geomorphology determines ecological sub-regions that have typical invertebrate communities, and diversity declines as anthropogenic pressure increases. Sixty-five stream sites were sampled for benthic invertebrates and physical–chemical variables across various watersheds. We used the Self-Organizing Map algorithm (neural network) to model relationships between invertebrate communities and environmental variables. Sites characterized by invertebrate communities clustered into two major subsets matching French Guiana’s hydro-ecoregions: the coastal alluvial plain characterized by recent sediment and low elevations, and the Guiana Shield characterized by an eroded rocky substrate and dense rainforests. Changes in community composition, and to a lesser extent taxonomic richness within each sub-region revealed ecological impacts of gold mining and logging, further clustering hydro-ecoregions into subsets of reference and impaired sites. Further analyses would, however, be needed to identify tipping points between natural and disturbed states, especially in remote headwater streams where gold mining had the harsher impact upon freshwater diversity, making upstream communities resembling the most downstream impacted ones.     

Does the Aboveground Herbivore Assemblage Influence Soil Bacterial Community Composition and Richness in Subalpine Grasslands?

Grassland ecosystems support large communities of aboveground herbivores that are known to directly and indirectly affect belowground properties such as the microbial community composition, richness, or biomass. Even though multiple species of functionally different herbivores coexist in grassland ecosystems, most studies have only considered the impact of a single group, i.e., large ungulates (mostly domestic livestock) on microbial communities. Thus, we investigated how the exclusion of four groups of functionally different herbivores affects bacterial community composition, richness, and biomass in two vegetation types with different grazing histories. We progressively excluded large, medium, and small mammals as well as invertebrate herbivores using exclosures at 18 subalpine grassland sites (9 per vegetation type). We assessed the bacterial community composition using terminal restriction fragment length polymorphism (T-RFLP) at each site and exclosure type during three consecutive growing seasons (2009–2011) for rhizosphere and mineral soil separately. In addition, we determined microbial biomass carbon (MBC), root biomass, plant carbon:nitrogen ratio, soil temperature, and soil moisture. Even though several of these variables were affected by herbivore exclusion and vegetation type, against our expectations, bacterial community composition, richness, or MBC were not. Yet, bacterial communities strongly differed between the three growing seasons as well as to some extent between our study sites. Thus, our study indicates that the spatiotemporal variability in soil microclimate has much stronger effects on the soil bacterial communities than the grazing regime or the composition of the vegetation in this high-elevation ecosystem.     

Mating Competition,Promiscuity, and Life History Traits as Predictors of Sexually Transmitted Disease Risk in Primates

Competition among males influences the distribution of copulations and should therefore influence the spread of sexually transmitted diseases (STDs). We developed a model to investigate STDs in the mating and social systems found in primates, and we tested predictions using comparative methods. In the model, groups were distributed on a square lattice in which males or females disperse and males undergo characteristic dominance trajectories at maturity (challenge vs. queuing). We investigated the impact of mating rate, mating skew, migration rate of males or females, and group size on disease spread and prevalence. The model generated several predictions: 1) STD prevalence is higher in females than males; 2) STD risk increases with copulation rate; 3) high skew is negatively associated with STD risk; 4) STD risk is higher for all individuals when females disperse and 5) when mortality rates are lower; and 6) reproductive skew and later age of male dominance (queuing) produce more strongly female-biased STD prevalence. In comparative tests, we quantified STD risk as prevalence and richness of sexually transmitted organisms at the host species level. We found positive associations between host longevity and higher STD richness, and only (nonsignificant) weak trends for females to have higher STD prevalence. Mating skew showed a weakly positive association with STD richness, contrary to predictions of our model but consistent with predictions from a previous model. In some tests, we also found that female dispersal resulted in greater STD infection risk. Collectively, these results demonstrate that mating competition and demography influence patterns of STD infection, with mortality rates having the strongest effects in comparative tests.     

An experimental test of diversity maintenance mechanisms,by a species removal experiment in a species-rich wooded meadow

Most theories of plant species coexistence assume the presence of diversity maintenance mechanisms, i.e. mechanisms enhancing species richness in a community. We wished to determine whether such a mechanism was operating by establishing a field experiment in the species-rich wooded meadow of Laelatu, western Estonia. Ten to seventeen subordinate species were removed periodically (for 4 years) from 10 permanent plots of 1×1 m (each plot had its specific list of excluded species; 10 plots served as control). Since the removed species were all subordinate ones, very little biomass was removed, but at the same time richness was reduced by 25–33%. If some diversity maintenance mechanism was operating, we would expect that immigration of other subordinate species would restore the original species richness. It was not possible to reject the null hypothesis of an identical immigration rate of new species into manipulated and control plots. The rate of small-scale species turnover was not affected by the removal of subordinate species. Interrelations of five richness characteristics were studied, by comparing empirical correlations among them, with those expected from a null model of random migration of species. The immigration rate of new species appeared to be related to the number of constant species, and immigration/extinction balance related to initial richness, more strongly than predicted by the null model. In the manipulated series these relationships matched the expectation from the null model. While the results generally support the so-called species pool hypothesis (and the carousel model), it seems that species small-scale turnover depends on the richness pattern in the studied grassland. In the case of plots with artificially reduced richness no departures can be detected from the random migration hypothesis.     

West African rivers as biogeographic islands: species richness of fish communities

Summary Some factors influencing the species richness of West African fish communities were studied in a sample of 26 rivers using four habitat and hydrologic variables. Analysis of a larger sample of 39 rivers showed that species richness was positively related to area. A power function with an exponent of 0.32 gave the best fit. As the surface area used was that of the catchment area and not that (unknown) of the river, the biological significance of this relationship and the possibilities of comparison were limited. Ridge regression analysis and forward stepwise selection indicated that a model that explained ln(species richness) as a function of ln(mean annual discharge) and ln-(catchment surface area) was best, accounting for 90% of the variance of the dependent variable. The combination of surface area and discharge was presumed to act through the volume of water available for the fishes and habitat productivity. Habitat diversity, measured by the diversity of the terrestrial vegetation covering the catchment area, had no significant positive effect when surface area was used in the regression. Rivers (islands) should have fewer species than tributaries of similar size since, for fishes within a river system (continent), there is free circulation between all its branches. The model derived from the river data underestimated the species richness of a sample of 11 tributaries. This was compatible with the hypothesis of higher population extinction rates in insular biotopes. The residuals of the linear model did not show random geographical distribution; the rivers in some areas had more species than expected. The possibility that historical factors, especially Quaternary climatic variations, might cause this distribution is discussed.