Habitat modifies the relationship between grass and herbivore species richness in a South African savanna.

The savanna ecosystem is dominated by grasses, which are a key food source for many species of grazing animals. This relationship creates a diverse mosaic of habitats and contributes to the high grass species richness of savannas. However, how grazing interacts with environmental conditions in determining grass species richness and abundance in savannas is still insufficiently understood. In the Kruger National Park, South Africa, we recorded grass species and estimated their covers in 60 plots 50 × 50 m in size, accounting for varying proximity to water and different bedrocks. To achieve this, we located plots (i) near perennial rivers, near seasonal rivers, and on crests that are distant from all water sources and (ii) on nutrient-rich basaltic and nutrient-poor granitic bedrock. The presence and abundance of large herbivores were recorded by 60 camera traps located in the same plots. Grass cover was higher at crests and seasonal rivers than at perennial rivers and on basalts than on granites. The relationship between grass species richness and herbivore abundance or species richness was positive at crests, while that between grass species richness and herbivore species richness was negative at seasonal rivers. We found no support for controlling the dominance of grasses by herbivores in crests, but herbivore-induced microsite heterogeneity may account for high grass species richness there. In contrast, the decrease in grass species richness with herbivore species richness at seasonal rivers indicates that the strong grazing pressure over-rides the resistance of some species to grazing and trampling. We suggest that the relationships between grasses and herbivores may work in both directions, but the relationship is habitat-dependent, so that in less productive environments, the effect of herbivores on vegetation prevails, while in more productive environments along rivers the effect of vegetation and water supply on herbivores is more important.

Magnifying the hotspot: descriptions of nine new species of many-plumed moths (Lepidoptera, Alucitidae), with an identification key to all species known from Cameroon.

This study confirms Mount Cameroon as an unprecedented hotspot for the diversity of many-plumed moths, with the discovery and description of nine new species: Alucitafako Ustjuzhanin & Kovtunovich, sp. nov., Alucitapyrczi Ustjuzhanin & Kovtunovich, sp. nov., Alucitasroczki Ustjuzhanin & Kovtunovich, sp. nov., Alucitapotockyi Ustjuzhanin & Kovtunovich, sp. nov., Alucitasedlaceki Ustjuzhanin & Kovtunovich, sp. nov., Alucitatonda Ustjuzhanin & Kovtunovich, sp. nov., Alucitaerzayi Ustjuzhanin & Kovtunovich, sp. nov., Alucitasokolovi Ustjuzhanin & Kovtunovich, sp. nov., and Alucitahirsuta Ustjuzhanin & Kovtunovich, sp. nov. Additionally, four additional species are reported from the Mount Cameroon area as new for the country: Alucitaagassizi, Alucitadohertyi, Alucitaplumigera, and Alucitarhaptica. Of the 89 Alucitidae known from the Afrotropics, the studied area hosts 36 species, most of which are endemic to the area. This unprecedented level of diversity and endemism within this lepidopteran family highlights Mount Cameroon's significance as a stronghold for specialised insect taxa. Efficient conservation efforts are necessary to protect these ecosystems and their associated unique microlepidopteran diversity.

Bioaccumulation of chemical elements at post-industrial freshwater sites varies predictably between habitats, elements and taxa: A power law approach.

Elevated environmental levels of elements originating from anthropogenic activities threaten natural communities and public health, as these elements can persist and bioaccumulate in the environment. However, their environmental risks and bioaccumulation patterns are often habitat-, species- and element-specific. We studied the bioaccumulation patterns of 11 elements in seven freshwater taxa in post-mining habitats in the Czech Republic, ranging from less polluted mining ponds to highly polluted fly ash lagoons. We found nonlinear, power-law relationships between the environmental and tissue concentrations of the elements, which may explain differences in bioaccumulation factors (BAF) reported in the literature. Tissue concentrations were driven by the environmental concentrations in non-essential elements (Al, As, Co, Cr, Ni, Pb and V), but this dependence was limited in essential elements (Cu, Mn, Se and Zn). Tissue concentrations of most elements were also more closely related to substrate than to water concentrations. Bioaccumulation was habitat specific in eight elements: stronger in mining ponds for Al and Pb, and stronger in fly ash lagoons for As, Cu, Mn, Pb, Se, V and Zn, although the differences were often minor. Bioaccumulation of some elements further increased in mineral-rich localities. Proximity to substrate, rather than trophic level, drove increased bioaccumulation levels across taxa. This highlights the importance of substrate as a pollutant reservoir in standing freshwaters and suggests that benthic taxa, such as molluscs (e.g., Physella) and other macroinvertebrates (e.g., Nepa), constitute good bioindicators. Despite the higher environmental risks in fly ash lagoons than in mining ponds, the observed ability of freshwater biota to sustain pollution supports the conservation potential of post-industrial sites. The power law approach used here to quantify and disentangle the effects of various bioaccumulation drivers may be helpful in additional contexts, increasing our ability to predict the effects of other contaminants and environmental hazards on biota.

Muddying the unexplored post-industrial waters: Biodiversity and conservation potential of freshwater habitats in fly ash sedimentation lagoons.

Deposits of fly ash and other coal combustion wastes are common remnants of the energy industry. Despite their environmental risks from heavy metals and trace elements, they have been revealed as refuges for threatened terrestrial biodiversity. Surprisingly, freshwater biodiversity of fly ash sedimentation lagoons remains unknown despite such lack of knowledge strongly limits the efficient restoration of fly ash deposits. We bring the first comprehensive survey of freshwater biodiversity, including nekton, benthos, zooplankton, phytoplankton, and macrophytes, in fly ash lagoons across industrial regions of the Czech Republic. To assess their conservation potential, we compared their biodiversity with abandoned post-mining ponds, the known strongholds of endangered aquatic species in the region with a shortage of natural ponds. Of 28 recorded threatened species, 15 occurred in the studied fly ash lagoons, some of which were less abundant or even absent in the post-mining ponds. These are often species of nutrient-poor, fishless waters with rich vegetation, although some are specialised extremophiles. Species richness and conservation value of most groups in the fly ash lagoons did not significantly differ from the post-mining ponds, except for species richness of benthos, zooplankton, and macrophytes, which were slightly lower in the fly ash lagoons. Although the concentrations of some heavy metals (mainly Se, V, and As) were significantly higher in the fly ash lagoons, they did not significantly affect species richness or conservation value of the local communities. The differences in species composition therefore does not seem to be caused by water chemistry. Altogether, we have shown that fly ash lagoons are refuges for threatened aquatic species, and we thus suggest maintaining water bodies during site restoration after the cessation of fly ash deposition. Based on our analyses of environmental variables, we discuss suitable restoration practices that efficiently combine biodiversity protection and environmental risk reduction.

Rapid radiation of ant parasitic butterflies during the Miocene aridification of Africa.

Africa has undergone a progressive aridification during the last 20 My that presumably impacted organisms and fostered the evolution of life history adaptations. We test the hypothesis that shift to living in ant nests and feeding on ant brood by larvae of phyto-predaceous Lepidochrysops butterflies was an adaptive response to the aridification of Africa that facilitated the subsequent radiation of butterflies in this genus. Using anchored hybrid enrichment we constructed a time-calibrated phylogeny for Lepidochrysops and its closest, non-parasitic relatives in the Euchrysops section (Poloyommatini). We estimated ancestral areas across the phylogeny with process-based biogeographical models and diversification rates relying on time-variable and clade-heterogeneous birth-death models. The Euchrysops section originated with the emerging Miombo woodlands about 22 million years ago (Mya) and spread to drier biomes as they became available in the late Miocene. The diversification of the non-parasitic lineages decreased as aridification intensified around 10 Mya, culminating in diversity decline. In contrast, the diversification of the phyto-predaceous Lepidochrysops lineage proceeded rapidly from about 6.5 Mya when this unusual life history likely first evolved. The Miombo woodlands were the cradle for diversification of the Euchrysops section, and our findings are consistent with the hypothesis that aridification during the Miocene selected for a phyto-predaceous life history in species of Lepidochrysops, with ant nests likely providing caterpillars a safe refuge from fire and a source of food when vegetation was scarce.

Moth Diversity Increases along a Continent-Wide Gradient of Environmental Productivity in South African Savannahs.

Environmental productivity, i.e., the amount of biomass produced by primary producers, belongs among the key factors for the biodiversity patterns. Although the relationship of diversity to environmental productivity differs among studied taxa, detailed data are largely missing for most groups, including insects. Here, we present a study of moth diversity patterns at local and regional scales along a continent-wide gradient of environmental productivity in southern African savannah ecosystems. We sampled diversity of moths (Lepidoptera: Heterocera) at 120 local plots along a gradient of normalized difference vegetation index (NDVI) from the Namib Desert to woodland savannahs along the Zambezi River. By standardized light trapping, we collected 12,372 specimens belonging to 487 moth species. The relationship between species richness for most analyzed moth groups and environmental productivity was significantly positively linear at the local and regional scales. The absence of a significant relationship of most moth groups' abundance to environmental productivity did not support the role of the number of individuals in the diversity-productivity relationship for south African moths. We hypothesize the effects of water availability, habitat complexity, and plant diversity drive the observed moth diversity patterns.

Water availability, bedrock, disturbance by herbivores, and climate determine plant diversity in South-African savanna.

To identify factors that drive plant species richness in South-African savanna and explore their relative importance, we sampled plant communities across habitats differing in water availability, disturbance, and bedrock, using the Kruger National Park as a model system. We made plant inventories in 60 plots of 50 × 50 m, located in three distinct habitats: (i) at perennial rivers, (ii) at seasonal rivers with water available only during the rainy season, and (iii) on crests, at least ~ 5 km away from any water source. We predicted that large herbivores would utilise seasonal rivers' habitats less intensely than those along perennial rivers where water is available throughout the year, including dry periods. Plots on granite harboured more herbaceous and shrub species than plots on basalt. The dry crests were poorer in herb species than both seasonal and perennial rivers. Seasonal rivers harboured the highest numbers of shrub species, in accordance with the prediction of the highest species richness at relatively low levels of disturbance and low stress from the lack of water. The crests, exposed to relatively low pressure from grazing but stressed by the lack of water, are important from the conservation perspective because they harbour typical, sometimes rare savanna species, and so are seasonal rivers whose shrub richness is stimulated and maintained by the combination of moderate disturbance imposed by herbivores and position in the middle of the water availability gradient. To capture the complexity of determinants of species richness in KNP, we complemented the analysis of the above local factors by exploring large-scale factors related to climate, vegetation productivity, the character of dominant vegetation, and landscape features. The strongest factor was temperature; areas with the highest temperatures reveal lower species richness. Our results also suggest that Colophospermum mopane, a dominant woody species in the north of KNP is not the ultimate cause of the lower plant diversity in this part of the park.

Spatiotemporal variation in the role of floral traits in shaping tropical plant-pollinator interactions.

The pollination syndrome hypothesis predicts that plants pollinated by the same pollinator group bear convergent combinations of specific floral functional traits. Nevertheless, some studies have shown that these combinations predict pollinators with relatively low accuracy. This discrepancy may be caused by changes in the importance of specific floral traits for different pollinator groups and under different environmental conditions. To explore this, we studied pollination systems and floral traits along an elevational gradient on Mount Cameroon during wet and dry seasons. Using Random Forest (Machine Learning) models, allowing the ranking of traits by their relative importance, we demonstrated that some floral traits are more important than others for pollinators. However, the distribution and importance of traits vary under different environmental conditions. Our results imply the need to improve our trait-based understanding of plant-pollinator interactions to better inform the debate surrounding the pollination syndrome hypothesis.

Thirteen moth species (Lepidoptera, Erebidae, Noctuidae) newly recorded in South Africa, with comments on their distribution.

Background: Thanks to the high diversity of ecosystems and habitats, South Africa harbours tremendous diversity of insects. The Kruger National Park, due to its position close to the border between two biogeographic regions and high heterogeneity of environmental conditions, represents an insufficiently studied hotspot of lepidopteran diversity. During our ecological research in the Kruger National Park, we collected abundant moth material, including several interesting faunistic records reported in this study. New information: We reported 13 species of moths which had not yet been recorded in South Africa. In many cases, our records represented an important extension of the species' known distribution, including two species (Ozarbagaedei and O.persinua) whose distribution ranges extended into the Zambezian biogeographic region. Such findings confirmed the poor regional knowledge of lepidopteran diversity.

Differences in Nectar Traits between Ornithophilous and Entomophilous Plants on Mount Cameroon.

Despite a growing number of studies, the role of pollinators as a selection agent for nectar traits remains unclear. Moreover, the lack of data from some biogeographic regions prohibits us from determining their general importance and global patterns. We analyzed nectar carbohydrate traits and determined the main pollinators of 66 plant species in the tropical forests of Mount Cameroon (tropical West Africa). The measured nectar traits included total sugar amounts and proportions of sucrose and hexoses (i.e., glucose and fructose). We report the nectar properties for plants visited by five pollinator groups (bees, butterflies, moths, hoverflies, and specialized birds). Our results indicate that, rather than specific evolution in each of the five plant groups, there was a unique nectar-trait evolution in plants pollinated by specialized birds. The ornithophilous plants had a higher proportion of sucrose and produced larger sugar amounts than the plants pollinated by insects. We also demonstrated a significant phylogenetic signal in the nectar properties in some lineages of the studied plants.

Elevational and seasonal patterns of butterflies and hawkmoths in plant-pollinator networks in tropical rainforests of Mount Cameroon.

Butterflies and moths are conspicuous flower visitors but their role in plant-pollinator interactions has rarely been quantified, especially in tropical rainforests. Moreover, we have virtually no knowledge of environmental factors affecting the role of lepidopterans in pollination networks. We videorecorded flower-visiting butterflies and hawkmoths on 212 plant species (> 26,000 recorded hrs) along the complete elevational gradient of rainforests on Mount Cameroon in dry and wet seasons. Altogether, we recorded 734 flower visits by 80 butterfly and 27 hawkmoth species, representing only ~ 4% of all flower visits. Although lepidopterans visited flowers of only a third of the plant species, they appeared to be key visitors for several plants. Lepidopterans visited flowers most frequently at mid-elevations and dry season, mirroring their local elevational patterns of diversity. Characteristics of interaction networks showed no apparent elevational or seasonal patterns, probably because of the high specialisation of all networks. Significant non-linear changes of proboscis and forewing lengths were found along elevation. A positive relationship between the lengths of proboscis of hesperiid butterflies and tube of visited flowers was detected. Differences in floral preferences were found between sphingids and butterflies, revealing the importance of nectar production, floral size and shape for sphingids, and floral colour for butterflies. The revealed trait-matching and floral preferences confirmed their potential to drive floral evolution in tropical ecosystems.

Effects of disturbances by forest elephants on diversity of trees and insects in tropical rainforests on Mount Cameroon.

Natural disturbances are essential for tropical forests biodiversity. In the Afrotropics, megaherbivores have played a key role before their recent decline. Contrastingly to savanna elephants, forest elephants' impact on ecosystems remains poorly studied. Few decades ago, forests on Mount Cameroon were divided by lava flows, not being crossed by a local population of forest elephants until now. We assessed communities of trees, butterflies and two guilds of moths in the disturbed and undisturbed forests split by the longest lava flow. We surveyed 32 plots, recording 2025 trees of 97 species, and 7853 insects of 437 species. The disturbed forests differed in reduced tree density, height, and high canopy cover, and in increased DBH. Forest elephants' selective browsing and foraging also decreased tree species richness and altered their composition. The elephant disturbance increased butterfly species richness and had various effects on species richness and composition of the insect groups. These changes were likely caused by disturbance-driven alterations of habitats and species composition of trees. Moreover, the abandonment of forests by elephants led to local declines of range-restricted butterflies. The recent declines of forest elephants across the Afrotropics probably caused similar changes in forest biodiversity and should be reflected by conservation actions.

New records of six moth (Lepidoptera: Erebidae, Lasiocampidae) species in south African countries, with comments on their distribution.

BACKGROUND: Southern Africa hosts a high diversity of ecosystems and habitats with a tremendous diversity of Lepidoptera. Although it is one of the most studied parts of the Afrotropics, the knowledge on diversity and distribution of south African moth fauna remains insufficient. To partly fill this gap, we surveyed macromoths by automatic light traps in five localities in two relatively less sampled south African countries. NEW INFORMATION: We reported six species and one genus (Remigioides) of moths which had not yet been recorded in Namibia or Zimbabwe. Although none of these records broadened the known distribution of individual species to a new biogeographical region, they still fill important gaps in their distributions. The known distributional ranges of two species have been substantially extended, although they are still within the same biogeographical regions: ca. 800 km southwards for Remigioides remigina (Mabille, 1884) and ca. 600 km westwards for Haplopacha cinerea Aurivillius, 1905.

Towards unravelling Wolbachia global exchange: a contribution from the Bicyclus and Mylothris butterflies in the Afrotropics.

BACKGROUND: Phylogenetically closely related strains of maternally inherited endosymbiotic bacteria are often found in phylogenetically divergent, and geographically distant insect host species. The interspecies transfer of the symbiont Wolbachia has been thought to have occurred repeatedly, facilitating its observed global pandemic. Few ecological interactions have been proposed as potential routes for the horizontal transfer of Wolbachia within natural insect communities. These routes are however likely to act only at the local scale, but how they may support the global distribution of some Wolbachia strains remains unclear. RESULTS: Here, we characterize the Wolbachia diversity in butterflies from the tropical forest regions of central Africa to discuss transfer at both local and global scales. We show that numerous species from both the Mylothris (family Pieridae) and Bicyclus (family Nymphalidae) butterfly genera are infected with similar Wolbachia strains, despite only minor interclade contacts across the life cycles of the species within their partially overlapping ecological niches. The phylogenetic distance and differences in resource use between these genera rule out the role of ancestry, hybridization, and shared host-plants in the interspecies transfer of the symbiont. Furthermore, we could not identify any shared ecological factors to explain the presence of the strains in other arthropod species from other habitats, or even ecoregions. CONCLUSION: Only the systematic surveys of the Wolbachia strains from entire species communities may offer the material currently lacking for understanding how Wolbachia may transfer between highly different and unrelated hosts, as well as across environmental scales.

Spatial covariance of herbivorous and predatory guilds of forest canopy arthropods along a latitudinal gradient.

In arthropod community ecology, species richness studies tend to be prioritised over those investigating patterns of abundance. Consequently, the biotic and abiotic drivers of arboreal arthropod abundance are still relatively poorly known. In this cross-continental study, we employ a theoretical framework in order to examine patterns of covariance among herbivorous and predatory arthropod guilds. Leaf-chewing and leaf-mining herbivores, and predatory ants and spiders, were censused on > 1000 trees in nine 0.1 ha forest plots. After controlling for tree size and season, we found no negative pairwise correlations between guild abundances per plot, suggestive of weak signals of both inter-guild competition and top-down regulation of herbivores by predators. Inter-guild interaction strengths did not vary with mean annual temperature, thus opposing the hypothesis that biotic interactions intensify towards the equator. We find evidence for the bottom-up limitation of arthropod abundances via resources and abiotic factors, rather than for competition and predation.

Even hotter hotspot: description of seven new species of many-plumed moths (Lepidoptera, Alucitidae) from Mount Cameroon.

Mount Cameroon, SW Cameroon, has already been described as a unique hotspot of the many-plumed moth (Lepidoptera, Alucitidae), with their local diversity unrivalled in the entire Afrotropics. We confirm its importance with description of seven new species: Alucita bakweri Ustjuzhanin & Kovtunovich, sp. nov., Alucita jana Ustjuzhanin & Kovtunovich, sp. nov., Alucita bakingili Ustjuzhanin & Kovtunovich, sp. nov., Alucita tatjana Ustjuzhanin & Kovtunovich, sp. nov., Alucita zuza Ustjuzhanin & Kovtunovich, sp. nov., Alucita deja Ustjuzhanin & Kovtunovich, sp. nov., and Alucita bokwango Ustjuzhanin & Kovtunovich, sp. nov. These descriptions have raised the known local diversity of many-plumed moth species on Mount Cameroon to 22, i.e., over a quarter of the known Afrotropical biodiversity of this group. This study also emphasises the great conservation importance of the area.

Are Reproductive Traits Related to Pollen Limitation in Plants? A Case Study from a Central European Meadow.

The deficiency of pollen grains for ovule fertilization can be the main factor limiting plant reproduction and fitness. Because of the ongoing global changes, such as biodiversity loss and landscape fragmentation, a better knowledge of the prevalence and predictability of pollen limitation is challenging within current ecological research. In our study we used pollen supplementation to evaluate pollen limitation (at the level of seed number and weight) in 22 plant species growing in a wet semi-natural meadow. We investigated the correlation between the pollen limitation index (PL) and floral traits associated with plant reproduction or pollinator foraging behavior. We recorded significant pollen limitation for approximately 41% of species (9 out of 22 surveyed). Seven species had a significant positive response in seed production and two species increased in seed weight after pollen supplementation. Considering traits, PL significantly decreased with the number of pollinator functional groups. The relationship of PL with other examined traits was not supported by our results. The causes of pollen limitation may vary among species with regard to (1) different reproductive strategies and life history, and/or (2) temporary changes in influence of biotic and abiotic factors at a site.

First records of 31 species of butterflies and moths (Lepidoptera) in Cameroon, with remarks on their elevational ranges.

BACKGROUND: The biodiversity of West and Central Africa is understudied, including butterflies and moths (Lepidoptera). Cameroon, through its position in between few biogeographic regions and diversity of habitats, is an important hotspot of lepidopteran diversity. However, the country also ranks low when it comes to local biodiversity knowledge. During our long-term ecological projects in the Cameroonian part of the Gulf of Guinea Highlands, we collected rich material of butterflies and moths, including a number of interesting faunistic records. NEW INFORMATION: In this study, we report 31 species of butterflies and moths which have not yet been recorded in Cameroon. These species comprised eight new genera records for the country. In many cases, our records represented an important extension of the species' known distribution, including ten species whose distribution ranges extended into the Guinean biogeographic region. We also comment on the species' elevational distribution ranges on Mount Cameroon where most of our records originated. Additionally, we confirm the presence of a butterfly Telchinia encedena, after more than a century since its first and so far its only record in Cameroon.

Amerila (Lepidoptera: Erebidae: Arctiinae) of Cameroon with morphological remarks on male and female genitalia.

Amerila is one of the most studied Afrotropical genera of Arctiinae. However, based on a regionally constrained sample of specimens from Mount Cameroon, we show how superficial our knowledge on these tiger moths is. Among six collected Amerila species, A. femina's female is described here for the first time, and A. mulleri and A. roseomarginata had never been recorded before in the country. Moreover, novel biological data are presented, including individual species' elevational ranges. Finally, female reproductive organs of the genus are illustrated here for the first time. The value of such regional studies is highlighted, with some remarks on necessary requirements of such small-scaled field sampling.

Two new species in the genus Andronymus Holland, 1896 (Lepidoptera, Hesperiidae).

During extensive field work in West Africa (Cameroon, Ivory Coast, Liberia, Sierra Leone), the authors collected two skipper species in the genus Andronymus (Lepidoptera, Hesperiidae), which would not fit the descriptions of any existing taxa. Both are described as new, A. magma sp. nov. is known only from Cameroon, while A. fenestra sp. nov. was found in a few localities in the Liberian sub-region of West Africa.