Mitogenomics and phylogenetics of twelve species of African Saturniidae (Lepidoptera).

African Saturniidae (Lepidoptera) include numerous species consumed at the caterpillar stage throughout the continent, and their importance to local communities as a source of nutrition and seasonal income cannot be overestimated. However, baseline genetic data with utility for the characterization of their diversity, phylogeography and phylogenetic relationships have remained scarce compared to their Asian counterparts. To bridge this gap, we sequenced the mitochondrial genomes of 12 species found in southern Africa for comparative mitogenomics and phylogenetic reconstruction of the family, including the first representatives of the tribes Eochroini and Micragonini. Mitochondrial gene content and organization were conserved across all Saturniidae included in the analyses. The phylogenetic positions of the 12 species were assessed in the context of publicly available mitogenomes using Bayesian inference and maximum likelihood (ML) methods. The monophyly of the tribes Saturniini, Attacini, Bunaeini and Micragonini, the sister relationship between Saturniini and Attacini, and the placement of Eochroa trimenii and Rhodinia fugax in the tribes Eochroini and Attacini, respectively, were strongly supported. These results contribute to significantly expanding genetic data available for African Saturniidae and allow for the development of new mitochondrial markers in future studies.

Fragmented landscapes affect honey bee colony strength at diverse spatial scales in agroecological landscapes in Kenya.

Landscape fragmentation and habitat loss at multiple scales directly affect species abundance, diversity, and productivity. There is a paucity of information about the effect of the landscape structure and diversity on honey bee colony strength in Africa. Here, we present new insights into the relationship between landscape metrics such as patch size, shape, connectivity, composition, and configuration and honey bee (Apis mellifera) colony strength characteristics. Remote-sensing-based landscape variables were linked to honey bee colony strength variables in a typical highly fragmented smallholder agroecological region in Kenya. We examined colonies in six sites with varying degrees of land degradation during the period from 2017 to 2018. Landscape structure was first mapped using medium resolution bitemporal Sentinel-1 and Sentinel-2 satellite imagery with an optimized random forest model. The influence of the surrounding landscape matrix was then constrained to two buffer distances, i.e., 1 km representing the local foraging scale and 2.5 km representing the wider foraging scale around each investigated apiary and for each of the six sites. The results of zero-inflated negative binomial regression with mixed effects showed that lower complexity of patch geometries represented by fractal dimension and reduced proportions of croplands were most influential at local foraging scales (1 km) from the apiary. In addition, higher proportions of woody vegetation and hedges resulted in higher colony strength at longer distances from the apiary (2.5 km). Honey bees in moderately degraded landscapes demonstrated the most consistently strong colonies throughout the study period. Efforts towards improving beekeeper livelihoods, through higher hive productivity, should target moderately degraded and heterogeneous landscapes, which provide forage from diverse land covers.

Spatio-Environmental Analysis of Vespula germanica Nest Records Explains Slow Invasion in South Africa.

Investigating the distributions of invasive species in marginal habitats can give clues to the factors constraining invasive spread. Vespula germanica is the most widely distributed of all the invasive Vespids, which in the Southern Hemisphere typically have large extensive invasive populations. In contrast, the invasion into South Africa has been slow and is still confined to a small geographic area. Here we analyse the distribution of all recent nest records in South Africa (n = 405). The distance to main rivers, mean annual rainfall, summer normalised difference moisture index (NDMI) values, and mean annual temperatures (average, minimum, maximum, and summer maximum temperature) was measured for every nest. We find that value ranges of these variables are different between the value ranges recorded for nests, the general distribution area of the wasp, and the area of absence. Optimised Hot Spot Analysis was used to quantify spatial structure in the measured climatic variables. Generally, factors related to moisture stress set the environmental limits of V. germanica's landscape distribution. Due to the strong preference of nesting sites close to river courses, for higher rainfall conditions, medium to medium-high NDMI values, and lower mean annual temperatures, it is unlikely that V. germanica will be able to spread uniformly where it is currently found in South Africa.

Horizon scanning for South African biodiversity: A need for social engagement as well as science.

A horizon scan was conducted to identify emerging and intensifying issues for biodiversity conservation in South Africa over the next 5-10 years. South African biodiversity experts submitted 63 issues of which ten were identified as priorities using the Delphi method. These priority issues were then plotted along axes of social agreement and scientific certainty, to ascertain whether issues might be "simple" (amenable to solutions from science alone), "complicated" (socially agreed upon but technically complicated), "complex" (scientifically challenging and significant levels of social disagreement) or "chaotic" (high social disagreement and highly scientifically challenging). Only three of the issues were likely to be resolved by improved science alone, while the remainder require engagement with social, economic and political factors. Fortunately, none of the issues were considered chaotic. Nevertheless, strategic communication, education and engagement with the populace and policy makers were considered vital for addressing emerging issues.

Seed survival of Australian Acacia in the Western Cape of South Africa in the presence of biological control agents and given environmental variation.

Studies of invasive Australian Acacia have shown that many seeds are still produced and accumulate in soil stored seed banks regardless of the presence of seed-targeting biological control agents. This is despite claims of biological control success, although there is generally a lack of data on the seed production of invasive Australian Acacia before and after the release of the respective agents. We aimed to quantify seed production and seed survival of invasive Australian Acacia currently under biological control. The seed production and survival (proportion of aborted, predated and surviving seeds) of A. longifolia, A. pycnantha and A. saligna were each studied at four to five sites in the Western Cape of South Africa. The relationships between seed production and stand characteristics were determined and the relative effects of seed predation and abortion on seed survival were established. The investigated invasive Australian Acacia produced many seeds that survived the pre-dispersal stage despite long-term presence of released biological control agents. It was shown that seed crop size is the only significant factor influencing seed survival of the studied Australian Acacia species. Furthermore, the seeds surviving per tree and per square meter were related to tree size. No quantitative evidence was found to suggest that seed-reducing biological control agents are having an impact on the population dynamics of their Australian Acacia hosts. This study illustrates the importance of studying the seed ecology of invasive plants before biological control agents are selected and released.

Invasive Australian Acacia seed banks: Size and relationship with stem diameter in the presence of gall-forming biological control agents.

Australian Acacia are invasive in many parts of the world. Despite significant mechanical and biological efforts to control their invasion and spread, soil-stored seed banks prevent their effective and sustained removal. In response South Africa has had a strong focus on employing seed reducing biological control agents to deal with Australian Acacia invasion, a programme that is considered as being successful. To provide a predictive understanding for their management, seed banks of four invasive Australian acacia species (Acacia longifolia, A. mearnsii, A. pycnantha and A. saligna) were studied in the Western Cape of South Africa. Across six to seven sites for each species, seed bank sizes were estimated from dense, monospecific stands by collecting 30 litter and soil samples. Average estimated seed bank size was large (1017 to 17261 seed m-2) as was annual input into the seed bank, suggesting that these seed banks are not residual but are replenished in size annually. A clear relationship between seed bank size and stem diameter was established indicating that mechanical clearing should be conducted shortly after fire-stimulated recruitment events or within old populations when seed banks are small. In dense, monospecific stands seed-feeding biological control agents are not effective in reducing seed bank size.

Including irrigation in niche modelling of the invasive wasp Vespula germanica (Fabricius) improves model fit to predict potential for further spread.

The European wasp, Vespula germanica (Fabricius) (Hymenoptera: Vespidae), is of Palaearctic origin, being native to Europe, northern Africa and Asia, and introduced into North America, Chile, Argentina, Iceland, Ascension Island, South Africa, Australia and New Zealand. Due to its polyphagous nature and scavenging behaviour, V. germanica threatens agriculture and silviculture, and negatively affects biodiversity, while its aggressive nature and venomous sting pose a health risk to humans. In areas with warmer winters and longer summers, queens and workers can survive the winter months, leading to the build-up of large nests during the following season; thereby increasing the risk posed by this species. To prevent or prepare for such unwanted impacts it is important to know where the wasp may be able to establish, either through natural spread or through introduction as a result of human transport. Distribution data from Argentina and Australia, and seasonal phenology data from Argentina were used to determine the potential distribution of V. germanica using CLIMEX modelling. In contrast to previous models, the influence of irrigation on its distribution was also investigated. Under a natural rainfall scenario, the model showed similarities to previous models. When irrigation is applied, dry stress is alleviated, leading to larger areas modelled climatically suitable compared with previous models, which provided a better fit with the actual distribution of the species. The main areas at risk of invasion by V. germanica include western USA, Mexico, small areas in Central America and in the north-western region of South America, eastern Brazil, western Russia, north-western China, Japan, the Mediterranean coastal regions of North Africa, and parts of southern and eastern Africa.

Potential supply of floral resources to managed honey bees in natural mistbelt forests.

Honey bees play a vital role in the pollination of flowers in many agricultural systems, while providing honey through well managed beekeeping activities. Managed honey bees rely on the provision of pollen and nectar for their survival and productivity. Using data from field plot inventories in natural mistbelt forests, we (1) assessed the diversity and relative importance of honey bee plants, (2) explored the temporal availability of honey bee forage (nectar and pollen resources), and (3) elucidated how plant diversity (bee plant richness and overall plant richness) influenced the amount of forage available (production). A forage value index was defined on the basis of species-specific nectar and pollen values, and expected flowering period. Up to 50% of the overall woody plant richness were found to be honey bee plant species, with varying flowering period. As expected, bee plant richness increased with overall plant richness. Interestingly, bee plants' flowering period was spread widely over a year, although the highest potential of forage supply was observed during the last quarter. We also found that only few honey bee plant species contributed 90 percent of the available forage. Surprisingly, overall plant richness did not significantly influence the bee forage value. Rather, bee plant species richness showed significant and greater effect. The results of this study suggest that mistbelt forests can contribute to increase the spatial and temporal availability of diverse floral resources for managed honey bees. Conservation efforts must be specifically oriented towards honey bee plant species in mistbelt forests to preserve and enhance their potential to help maintain honey bee colonies. The implications for forest management, beekeeping activities and pollination-based agriculture were discussed.

Mutually beneficial pollinator diversity and crop yield outcomes in small and large farms.

Ecological intensification, or the improvement of crop yield through enhancement of biodiversity, may be a sustainable pathway toward greater food supplies. Such sustainable increases may be especially important for the 2 billion people reliant on small farms, many of which are undernourished, yet we know little about the efficacy of this approach. Using a coordinated protocol across regions and crops, we quantify to what degree enhancing pollinator density and richness can improve yields on 344 fields from 33 pollinator-dependent crop systems in small and large farms from Africa, Asia, and Latin America. For fields less than 2 hectares, we found that yield gaps could be closed by a median of 24% through higher flower-visitor density. For larger fields, such benefits only occurred at high flower-visitor richness. Worldwide, our study demonstrates that ecological intensification can create synchronous biodiversity and yield outcomes.

Tree species diversity promotes aboveground carbon storage through functional diversity and functional dominance.

The relationship between biodiversity and ecosystem function has increasingly been debated as the cornerstone of the processes behind ecosystem services delivery. Experimental and natural field-based studies have come up with nonconsistent patterns of biodiversity-ecosystem function, supporting either niche complementarity or selection effects hypothesis. Here, we used aboveground carbon (AGC) storage as proxy for ecosystem function in a South African mistbelt forest, and analyzed its relationship with species diversity, through functional diversity and functional dominance. We hypothesized that (1) diversity influences AGC through functional diversity and functional dominance effects; and (2) effects of diversity on AGC would be greater for functional dominance than for functional diversity. Community weight mean (CWM) of functional traits (wood density, specific leaf area, and maximum plant height) were calculated to assess functional dominance (selection effects). As for functional diversity (complementarity effects), multitrait functional diversity indices were computed. The first hypothesis was tested using structural equation modeling. For the second hypothesis, effects of environmental variables such as slope and altitude were tested first, and separate linear mixed-effects models were fitted afterward for functional diversity, functional dominance, and both. Results showed that AGC varied significantly along the slope gradient, with lower values at steeper sites. Species diversity (richness) had positive relationship with AGC, even when slope effects were considered. As predicted, diversity effects on AGC were mediated through functional diversity and functional dominance, suggesting that both the niche complementarity and the selection effects are not exclusively affecting carbon storage. However, the effects were greater for functional diversity than for functional dominance. Furthermore, functional dominance effects were strongly transmitted by CWM of maximum plant height, reflecting the importance of forest vertical stratification for diversity-carbon relationship. We therefore argue for stronger complementary effects that would be induced also by complementary light-use efficiency of tree and species growing in the understory layer.

Valuation of pollinator forage services provided by Eucalyptus cladocalyx.

We assess the monetary value of forage provisioning services for honeybees as provided by an alien tree species in the Western Cape province of South Africa. Although Eucalyptus cladocalyx is not an officially declared invader, it is cleared on a regular basis along with other invasive Eucalyptus species such as Eucalyptus camaldulensis, and Eucalyptus conferruminata (which have been prioritised for eradication in South Africa). We present some of the trade-offs associated with the clearing of E. cladocalyx by means of a practical example that illustrates a situation where the benefits of the species to certain stakeholders could support the containment of the species in demarcated areas, while allowing clearing outside such areas. Given the absence of market prices for such forage provisioning services, the replacement cost is used to present the value of the loss in forage as provided by E. cladocalyx if the alien tree species is cleared along with invasive alien tree species. Two replacement scenarios formed the basis for our calculations. The first scenario was an artificial diet as replacement for the forage provisioning service, which yielded a direct cost estimate of US$7.5 m per year. The second was based on a Fynbos cultivation/restoration initiative aimed at substituting the forage provisioning service of E. cladocalyx, which yielded a direct cost of US$20.2 m per year. These figures provide estimates of the potential additional cost burden on the beekeeping industry if E. cladocalyx is completely eradicated from the Western Cape. The cost estimates should be balanced against the negative impacts of E. cladocalyx on ecosystem services in order to make an informed decision with regard to appropriate management strategies for this species. The findings therefore serve as useful inputs to balance trade-offs for alien species that are considered as beneficial to some, but harmful to other.

Correlations between physical and chemical defences in plants: tradeoffs, syndromes, or just many different ways to skin a herbivorous cat?

Most plant species have a range of traits that deter herbivores. However, understanding of how different defences are related to one another is surprisingly weak. Many authors argue that defence traits trade off against one another, while others argue that they form coordinated defence syndromes. We collected a dataset of unprecedented taxonomic and geographic scope (261 species spanning 80 families, from 75 sites across the globe) to investigate relationships among four chemical and six physical defences. Five of the 45 pairwise correlations between defence traits were significant and three of these were tradeoffs. The relationship between species' overall chemical and physical defence levels was marginally nonsignificant (P = 0.08), and remained nonsignificant after accounting for phylogeny, growth form and abundance. Neither categorical principal component analysis (PCA) nor hierarchical cluster analysis supported the idea that species displayed defence syndromes. Our results do not support arguments for tradeoffs or for coordinated defence syndromes. Rather, plants display a range of combinations of defence traits. We suggest this lack of consistent defence syndromes may be adaptive, resulting from selective pressure to deploy a different combination of defences to coexisting species.

Putting plant resistance traits on the map: a test of the idea that plants are better defended at lower latitudes.

• It has long been believed that plant species from the tropics have higher levels of traits associated with resistance to herbivores than do species from higher latitudes. A meta-analysis recently showed that the published literature does not support this theory. However, the idea has never been tested using data gathered with consistent methods from a wide range of latitudes. • We quantified the relationship between latitude and a broad range of chemical and physical traits across 301 species from 75 sites world-wide. • Six putative resistance traits, including tannins, the concentration of lipids (an indicator of oils, waxes and resins), and leaf toughness were greater in high-latitude species. Six traits, including cyanide production and the presence of spines, were unrelated to latitude. Only ash content (an indicator of inorganic substances such as calcium oxalates and phytoliths) and the properties of species with delayed greening were higher in the tropics. • Our results do not support the hypothesis that tropical plants have higher levels of resistance traits than do plants from higher latitudes. If anything, plants have higher resistance toward the poles. The greater resistance traits of high-latitude species might be explained by the greater cost of losing a given amount of leaf tissue in low-productivity environments.

Natural and within-farmland biodiversity enhances crop productivity.

Ongoing expansion of large-scale agriculture critically threatens natural habitats and the pollination services they offer. Creating patches with high plant diversity within farmland is commonly suggested as a measure to benefit pollinators. However, farmers rarely adopt such practice, instead removing naturally occurring plants (weeds). By combining pollinator exclusion experiments with analysis of honeybee behaviour and flower-visitation webs, we found that the presence of weeds allowed pollinators to persist within sunflower fields, maximizing the benefits of the remaining patches of natural habitat to productivity of this large-scale crop. Weed diversity increased flower visitor diversity, hence ameliorating the measured negative effects of isolation from natural habitat. Although honeybees were the most abundant visitors, diversity of flower visitors enhanced honeybee movement, being the main factor influencing productivity. Conservation of natural patches combined with promoting flowering plants within crops can maximize productivity and, therefore, reduce the need for cropland expansion, contributing towards sustainable agriculture.

Dissecting the plant-insect diversity relationship in the Cape.

It has been argued that insect diversity in the Cape is disproportionately low, considering the unusually high plant diversity in this region. Recent studies have shown that this is not the case, but the precise mechanisms linking plant diversity and insect diversity in the Cape are still poorly understood. Here we use a dated genus-level phylogenetic tree of the Cape plants to assess how plant phylogenetic diversity compares with taxonomic diversity at various levels in predicting insect diversity. We find that plant phylogenetic diversity (PD) is a better predictor of insect species diversity that plant species diversity, but the number of plant genera is overall as good a predictor as PD, and much easier to calculate. The relationship is strongest between biomes, suggesting that the relationship between plant diversity and insect diversity is to a large extent indirect, both variables being driven by the same abiotic factors and possibly by common diversification, immigration and extinction histories. However, a direct relationship between plant diversity and insect diversity can be detected at fine scales, at least within certain biomes. Diversity accumulation curves also indicate that the way plant phylogenetic diversity and the number of plant genera increase over spatial scales is most similar to that for insect species; plant species show a greater increase at large spatial scales due to high numbers of local endemics.

Valuing insect pollination services with cost of replacement.

Value estimates of ecosystem goods and services are useful to justify the allocation of resources towards conservation, but inconclusive estimates risk unsustainable resource allocations. Here we present replacement costs as a more accurate value estimate of insect pollination as an ecosystem service, although this method could also be applied to other services. The importance of insect pollination to agriculture is unequivocal. However, whether this service is largely provided by wild pollinators (genuine ecosystem service) or managed pollinators (commercial service), and which of these requires immediate action amidst reports of pollinator decline, remains contested. If crop pollination is used to argue for biodiversity conservation, clear distinction should be made between values of managed- and wild pollination services. Current methods either under-estimate or over-estimate the pollination service value, and make use of criticised general insect and managed pollinator dependence factors. We apply the theoretical concept of ascribing a value to a service by calculating the cost to replace it, as a novel way of valuing wild and managed pollination services. Adjusted insect and managed pollinator dependence factors were used to estimate the cost of replacing insect- and managed pollination services for the Western Cape deciduous fruit industry of South Africa. Using pollen dusting and hand pollination as suitable replacements, we value pollination services significantly higher than current market prices for commercial pollination, although lower than traditional proportional estimates. The complexity associated with inclusive value estimation of pollination services required several defendable assumptions, but made estimates more inclusive than previous attempts. Consequently this study provides the basis for continued improvement in context specific pollination service value estimates.

Spatially explicit analyses unveil density dependence.

Density-dependent processes are fundamental in the understanding of species population dynamics. Whereas the benefits of considering the spatial dimension in population biology are widely acknowledged, the implications of doing so for the statistical detection of spatial density dependence have not been examined. The outcome of traditional tests may therefore differ from those that include ecologically relevant locational information on both the prey species and natural enemy. Here, we explicitly incorporate spatial information on individual counts when testing for density dependence between an insect herbivore and its parasitoids. The spatially explicit approach used identified significant density dependence more frequently and in different instances than traditional methods. The form of density dependence detected also differed between methods. These results demonstrate that the explicit consideration of patch location in density-dependence analyses is likely to significantly alter current understanding of the prevalence and form of spatial density dependence in natural populations.