Paul-André Calatayud

Phylogeography and population genetics of the maize stalk borer Busseola fusca (Lepidoptera, Noctuidae) in sub‐Saharan Africa

The population genetics and phylogeography of African phytophagous insects have received little attention. Some, such as the maize stalk borer Busseola fusca, display significant geographic differences in ecological preferences that may be congruent with patterns of molecular variation. To test this, we collected 307 individuals of this species from maize and cultivated sorghum at 52 localities in West, Central and East Africa during the growing season. For all collected individuals, we sequenced a fragment of the mitochondrial cytochrome b. We tested hypotheses concerning the history and demographic structure of this species. Phylogenetic analyses and nested clade phylogeographic analyses (NCPA) separated the populations into three mitochondrial clades, one from West Africa, and two — Kenya I and Kenya II — from East and Central Africa. The similar nucleotide divergence between clades and nucleotide diversity within clades suggest that they became isolated at about the same time in three different refuges in sub‐Saharan Africa and have similar demographic histories. The results of mismatch distribution analyses were consistent with the demographic expansion of these clades. Analysis of molecular variance (amova) indicated a high level of geographic differentiation at different hierarchical levels. NCPA suggested that the observed distribution of haplotypes at several hierarchical levels within the three major clades is best accounted for by restricted gene flow with isolation by distance. The domestication of sorghum and the introduction of maize in Africa had no visible effect on the geographic patterns observed in the B. fusca mitochondrial genome.

Diversity of lepidopteran stem borers on monocotyledonous plants in eastern Africa and the islands of Madagascar and Zanzibar revisited.

Surveys were completed in Eritrea, Ethiopia, Kenya, Madagascar, Mozambique, Tanzania, Uganda and Zanzibar to assess the lepidopteran stem borer species diversity on wild host plants. A total of 24,674 larvae belonging to 135 species were collected from 75 species of wild host plants belonging to the Poaceae, Cyperaceae and Typhaceae. Amongst them were 44 noctuid species belonging to at least nine genera, 33 crambids, 15 pyralids, 16 Pyraloidea species not yet identified, 25 tortricids and three cossids. The noctuid larvae represented 73.6% of the total number of larvae collected, with 66.3, 3.5 and 3.8% found on Poaceae, Cyperaceae and Typhaceae, respectively. The Crambidae, Pyralidae, Tortricidae and Cossidae represented 19.8, 1.9, 2.5 and 0.1% of the total larvae collected, respectively, with 90.4% of the Crambidae and Pyralidae collected from Poaceae, and 99.7% of the Tortricidae collected from Cyperaceae. The lepidopteran stem borer species diversity in the wild host plants was far more diverse than previously reported.

Geographic distribution and host plant ranges of East African noctuid stem borers

Abstract Surveys were carried out in Kenya, Tanzania, Uganda and Zanzibar to establish geographic distribution in the main vegetation mosaics and ecological (host plant range, feeding behaviour) characteristics of the East African noctuid stem borers. 49 wild plant species belonging to Poaceae, Cyperaceae and Typhaceae were found to harbour stem borers in the six vegetation mosaics surveyed. A total of 36 noctuid species belonging to nine genera were identified from 14,318 larvae collected, out of which 17 were new to science. The species diversity varied among vegetation mosaics and host plants. Most borer species appeared to be specialised feeders with 24 species being monophagous. Species belonging to the same types (named as the Busseola Thurau 1904 and the Sesamia Guenée 1852 types) or genus harboured common ecological characteristics such as pigmentation and feeding site. The Sciomesa Tams and Bowden 1953 genus was an exception as it had a mixture of these characters.

Electrically recorded feeding behaviour of cassava mealybug on host and non‐host plants

The feeding behaviour of cassava mealybug (Phenacoccus manihoti) was analysed by the electrical penetration graph technique (EPG, DC‐system) and by light microscopy. This study confirmed a typical phloem‐feeding behaviour with an aphid‐like predominance of extracellular pathways of stylets. Similarities of EPGs from mealybugs with those of aphids and whiteflies allowed adoption of standard pattern labelling. The main differences with aphid EPGs consisted of fewer but longer intracellular punctures (pd duration of 20 s vs 5–7 s for aphids), longer times to the first phloem‐ingestion period and a lower motility of stylets within the phloem searching process. Comparison of the feeding behaviour on 6 plant types, including two occasional hosts (Talinum and poinsettia), one cassava hybrid and three common cassava cultivars (Manihot esculenta), allowed differentiation of penetration profiles. Intracellular punctures were shorter on occasional hosts, on which phloem finding and subsequent ingestion were markedly delayed although a great variability within plants persisted. Chemical analysis of some secondary plant substances revealed that none of the plants tested contained detectable amounts of alkaloids, and that cyanides were restricted to true hosts (cassavas and Manihot hybrid). Levels of total flavonoids did not differ between hosts and non‐hosts, in contrast to those of phenolic acids. Preliminary correlations with EPG patterns are discussed.

Distribution, pest status and agro-climatic preferences of lepidopteran stem borers of maize in Kenya

Abstract Lepidopterous stem borers are the main field insect pests that attack maize, Zea mays L. in tropical Africa. A survey was carried during the long and short rain cropping seasons of 2002 / 2003 across six main agro-climatic zones (ACZs) to determine the spatial distribution of important stem borer species in Kenya. A total of 474 visits were made in the seventy-eight localities conveniently chosen to represent each of the six ACZs. 189,600 stems were checked for infestation, of which 27,799 infested stems were destructively cut and dissected for stem borer larvae identification. An average of 1.4 stem borer larvae were recovered per infested plant. 54.5% of the recovered larvae were identified as Chilo partellus (Swinhoe) (Crambidae), 39.7% as Busseola fusca (Fuller) (Noctuidae), 4.5% as Sesamia calamistis Hampson (Noctuidae) and 0.8% as Chilo orichalcociliellus (Strand) (Crambidae). Minor species present included Eldana saccharina Walker (Pyralidae), Sesamia nonagrioides (Lefebvre) (Noctuidae), Sesamia cretica Lederer (Noctuidae), Sesamia sp. (Noctuidae), Sciomesa piscator Fletcher (Noctuidae), Busseola sp near phaia (Noctuidae), Chilo sp, Ematheudes sp 1 (Pyralidae) and Ematheudes sp 2 (Pyralidae). Farms were grouped into respective ACZs for statistical analysis and subsequent comparison of dominant species. Results indicated that B. fusca was the dominant stem borer species in high potential zones (highland tropics, moist transitional zone and moist midaltitude) while the exotic C. partellus dominated smallholder farms in low potential zones (dry midaltitude, dry transional and lowland tropical zone). Within each ACZs, there was evidence of variation in species proportions between seasons. These spatio-temporal differences in community structure are discussed in terms of agro-climatic biological adaptations.

Influence of secondary compounds in the phloem sap of cassava on expression of antibiosis towards the mealybug Phenacoccus manihoti

Identification and assay of cyanogenic and phenolic compounds in phloem sap of cassava (Manihot esculenta Crantz, Euphorbiaceae) and in honeydew of the cassava mealybug Phenacoccus manihoti Matt. Ferr. (Homoptera, Pseudococcidae) were realised.

Influence of water‐stressed cassava on Phenacoccus herreni and three associated parasitoids

The influence of cassava Manihot esculenta Crantz grown under condition of water‐stress on development and reproduction of the cassava mealybug, Phenacoccus herreni Cox & Williams, and levels of parasitism of three encyrtid parasitoids, Apoanagyrus diversicornis Howard, Aenasius vexans Kerrich, and Acerophagus coccois Smith, were studied in the laboratory. Two cassava cultivars were used: CM 507‐37 (drought‐tolerant) and CMC 40. A 30 day period of water stress, imposed by reducing the irrigation volume, led to a reduction in shoot development and stomatal conductance of leaves of both cassava genotypes. Phenacoccus herreni development and reproduction were favoured by cassava under water shortage. Parasitism decreased and water stress appeared to enhance the encapsulation of parasitoid eggs or larvae by the mealybug. In the case of the parasitoid A. diversicornis, there was a decrease in size of female progeny, suggesting a lower fitness in this species on cassava plants under water stress. All results indicated that cassava grown under low water availability favoured P. herreni development and reproduction, and affected the success of parasitism and, depending on the species, parasitoid development. The drought‐tolerance characteristic of cassava genotypes and parasitoid species most suitable for controlling P. herreni in drought‐stricken areas are discussed.

Candidate Chemosensory Genes in the Stemborer Sesamia nonagrioides

The stemborer Sesamia nonagrioides is an important pest of maize in the Mediterranean Basin. Like other moths, this noctuid uses its chemosensory system to efficiently interact with its environment. However, very little is known on the molecular mechanisms that underlie chemosensation in this species. Here, we used next-generation sequencing (454 and Illumina) on different tissues from adult and larvae, including chemosensory organs and female ovipositors, to describe the chemosensory transcriptome of S. nonagrioides and identify key molecular components of the pheromone production and detection systems. We identified a total of 68 candidate chemosensory genes in this species, including 31 candidate binding-proteins and 23 chemosensory receptors. In particular, we retrieved the three co-receptors Orco, IR25a and IR8a necessary for chemosensory receptor functioning. Focusing on the pheromonal communication system, we identified a new pheromone-binding protein in this species, four candidate pheromone receptors and 12 carboxylesterases as candidate acetate degrading enzymes. In addition, we identified enzymes putatively involved in S. nonagrioides pheromone biosynthesis, including a ∆11-desaturase and different acetyltransferases and reductases. RNAseq analyses and RT-PCR were combined to profile gene expression in different tissues. This study constitutes the first large scale description of chemosensory genes in S. nonagrioides.

Importance of plant physical cues in host acceptance for oviposition by Busseola fusca

Plant and surrogate stems exhibiting specific combinations of physical cues were used to determine which plant‐related stimuli influence the oviposition of Busseola fusca (Fuller) (Lepidoptera: Noctuidae). The number of eggs and egg batches laid per female increased with an increase in diameter of both natural and artificial stems. Direct observations of the oviposition behaviour (walking, antennating, and sweeping with the ovipositor) indicated that the female moths preferred oviposition supports with a large diameter and non‐pubescent or smooth surfaces over pubescent or rough ones. Pubescence and rough surfaces significantly affected the behavioural steps leading to oviposition by interfering with the ovipositor sweep process necessary to find a suitable oviposition site. Furthermore, more eggs and egg batches were laid on soft than rigid supports. The rigidity of the support affected the proper insertion of the ovipositor for egg deposition. Our results underline the importance of physical stimuli in B. fuscas choice of an oviposition site, which may facilitate the identification of potential host plants or preferred oviposition sites on a plant for this species.

The role of wild host plants in the abundance of lepidopteran stem borers along altitudinal gradient in Kenya

Abstract Presence of wild host plants of stem borers in cereal-growing areas has been considered as reservoirs of lepidopteran stem borers, responsible for attack of crops during the growing season. Surveys to catalogue hosts and borers as well as to assess the abundance of the hosts were carried out during the cropping and non-cropping seasons in different agro-ecological zones along varying altitude gradient in Kenya. A total of 61 stem borer species belonging to families Noctuidae (25), Crambidae (14), Pyralidae (9), Tortricidae (11) and Cossidae (2) were recovered from 42 wild plant species. Two noctuids, Busseola fusca (Fuller), Sesamia calamistis Hampson, and two crambids, Chilo partellus (Swinhoe) and Chilo orichalcociliellus (Strand) were the four main borer species found associated with maize plants. In the wild, B. fusca was recovered from a limited number of host plant species and among them were Sorghum arundinaceum (Desvaux) Stapf, Setaria megaphylla (Steudel) T. Durand & Schinz, Arundo donax L. and Pennisetum purpureum Schumacher. In contrast, the host range of C. partellus was considerably wider [13 for S. calamistis]. However, the number of larvae of these species was lower in the wild compared to cultivated fields, thus the role of natural habitat as a reservoir for cereal stem borers requires further studies. Importance of the wild host plants as well as borer diversity along the altitudinal gradient is discussed.