Diet of Zonocerus variegatus (Linné, 1758) (Orth., Acrididae) in cassava fields in Bénin

Abstract:  This paper presents the results of a study on the diet of Zonocerus variegatus in cassava agro-ecosystems at Allada, south Benin. The diet was determined by the analysis of plant fragments in the faeces of insects sampled in the fields. The grasshoppers studied consumed 21 plant species among the 81 found in the fields. Cassava, maize, Mallotus oppositifolius and Chromolaena odorata were the major food plants for Z. variegatus in this system. The presence of Z. variegatus in different fields was observed depending on the season and the presence of certain plant species. There was no relationship between plant frequency in the diet and in the field. Cassava was the preferred food resource for mature nymphs and adults. During the dry season, young nymphs stayed near the hatching sites and where good food resources were available. At this time, the grasshoppers were present in a field where cassava was not abundant but where maize and weeds were available. During the dry season, Z. variegatus had a broader diet than during the wet season. This contradicts the current hypothesis that the water content of plants and the floral diversity determines food preference. The dry season is also the time when C. odorata leaves were found in the faeces of both adults and nymphs. These observations made in West Africa are similar to those made in Central Africa, particularly for the number of plant species found in the faeces, 21 (26% of all plant species found in the fields) in Bénin and 17 (23%) in the Congo. The increase of the sample size and the great differences between the ecoregional situation of the study sites do not substantially change the number of plant species included in the diets. In the area covered by this study, C. odorata is not the major substitute for cassava during the dry season, although it remains a major food source and an especially visible food resource.     

Relationships between feeding of Philaenus spumarius (L.) and the amino acid concentration in the xylem sap

Summary. 1. P.spwnarius nymphs selected young, upper leaves of Xanthium strumarium plants, which have a relatively high amino acid concentration in the xylem sap.
2. Nymphs selected or rejected a host leaf as a feeding site after a test ingestion of plant sap.
3. Nymphs fed on detached leaves kept with their petioles in a solution of amino acids, in preference to leaves with petioles in a solution containing no amino acids.
4. Nymphs caged on leaves with relatively high amino acid concentrations in the xylem sap suffered a lower mortality after 7 days than nymphs caged on leaves with low amino acid concentrations in the xylem sap.
5. Mean excretory rate increased with the increase in the amino acid concentration in the xylem sap for third and fourth instar nymphs, and adults, but not in fifth instars.     

Effect of food plants on the volume of repellent secretion obtained in adult Zonocerus variegatus (Orthoptera: Pyrgomorphidae)

The volume of secretion obtained from adult Zonocerus variegatus (Orthoptera: Pyrgomorphidae) was influenced by the type of food plants. Insects fed on leaves of cassava Manihot esculenta, bitter leaves Vernonia amygdalina, and a mixture of M. esculenta and Acalypha wilkesiana gave a good volume of secretion while Chromolaena odorata, Elaeis guinensis, Aspilia africana and Citrus sinensis did not favour secretion production. No significant difference was recorded in the volume of secretion obtained from Z. variegatus from the two seasons irrespective of the food plant. Similarly, food plants gave no significant difference on the volume of secretion between the two seasons.     

A non-specific aggregation pheromone in two Australian reptile ticks

Larvae, nymphs, adult males and adult females of each of the Australian reptile tick species Aponomma hydrosauri and Amblyomma limbatum took up significantly aggregated distributions when left for 12 h in the dark in petri dishes. Nymphs of the two species placed together formed co-aggregations. Pheromonal communication was implicated by the attraction of nymphs to filter paper squares which had been in contact with other nymphs. Each species was attracted to pheromone from the other species and no conspecific preference was detected. A common ancestral pheromone is suggested, which may have been retained after speciation by stabilizing selection for a behaviour of major importance to tick survival.     

Tropidacris collaris (Orthoptera: Romaleidae) expands its damage in orchards of the dwarf cashew,Anacardium occidentale (Anacardiaceae)

The giant grasshopper Tropidacris collaris (Stoll, 1813) (Orthoptera: Romaleidae) is a Neotropical species that causes economic damage to diverse and unrelated crops. We report herein the occurrence of this grasshopper in orchards of dwarf cashew Anacardium occidentale L. (Anacardiaceae) located in the state of Ceará, one of the main cashew‐producing areas of Brazil. Nymphs and adults were observed feeding on cashew leaves during June and July 2016, causing partial or total defoliation of some plants. This report updates the feeding plants of T. collaris and its distribution in Brazil and serves as an early warning of possible future attacks by the pest on cashew orchards in the northeastern region of the country.     

Transmission of potato leafroll virus in relation to the honeydew excretion of Myzus persicae

Honeydew excretion of single Myzus persicae nymphs on potato leafroll virus (PLVR)-infected Physalis floridana was studied during the acquisition access period (AAP) in relation to the efficiency of virus transmission.
With increasing length of the AAP, the percentage of nymphs that transmitted the virus increased. These nymphs produced significantly more honeydew droplets during the AAP on PLRV-infected P. floridana plants than nymphs which failed to transmit the virus. However, the number of honeydew droplets excreted during the AAP by transmitting nymphs did not affect the length of the latency period. Nymphs which infected the first test plant after a short latency period produced a similar amount of honeydew during the AAP to those with a longer latency period.
Honeydew excretion recorded on plants of varied age, showed that nymphs feeding on bottom leaves of infected plants produced more honeydew droplets than on comparable leaves of healthy plants. On infected plants, nymphs produced more honeydew droplets on bottom leaves with pronounced symptoms than on top leaves that hardly showed any symptom of PLRV infection.
The concentration of viral antigen measured by ELISA was lower in top leaves than in bottom leaves of infected plants. Nevertheless, nymphs feeding on top leaves transmitted the virus more efficiently than those which used bottom leaves as virus source. When bottom leaves were used as a virus source, the percentage of viruliferous nymphs decreased with plant age. These results indicate that the availability of virus for acquisition by aphids declines with increasing plant age and symptom severity.     

Pharmacological properties of the repellent secretion of Zonocerus variegatus (Orthoptera: Prygomorphidae)

The odours of the whole body and the secretion of Zonocerus variegatus were easily recognised and perceived by human volunteers. However, the secretion odour is not related to the odour of the food plant consumed by the grasshopper. The repellency of Z. variegatus becomes more pronounced in the 6th. and adult instars whose gland lumens contain an appreciable volume of secretion. The secretion odour is so strong that even dilution does not affect its repulsiveness to humans. The secretion had pharmacological properties: it induced contraction in rat (Rattus rattus) stomach smooth muscle preparations and guinea pig (Cavia porcellus) ileum, and induced oedema formation in the rat hind paw. The secretion was not lethal to the animals used in this study, effects were temporary and recovery occurs after a short time.     

Habitat use and phenology of the large insular endemic grasshopper Acrostira euphorbiae (Orthoptera: Pamphagidae)

The habitat use and the phenology of the large grasshopper Acrostira euphorbiae García and Oromí endemic to La Palma (Canary Islands) are studied. This grasshopper is entirely dependent on the Canarian endemic shrub Euphorbia lamarckii both for food and to avoid predation. Adults stay on subapical branches during the day, probably to reduce the risk of predation, and climb up to the apex at night to feed. While females seem to ensure the genetic diversity of offspring by waiting for visits by different males, the latter have to move to guarantee their reproductive success. Monophagy in this species may be related to the year-round presence of tender shoots, and to the predator-repellent toxic latex found in the host plant. Unlike related species from continental areas, adults and nymphs of A. euphorbiae are present almost all year round, probably in adaptation to the particular climate of the islands. Nymphs are more abundant in winter, when Euphorbia leaves are most available. However, adults are more abundant than nymphs in spring, summer and autumn. Males develop more quickly than females, an apparent reproductive strategy based on achieving sexual maturity to coincide with females undergoing imaginal moulting. Matings start immediately after adult females emerge. Densities oscillated between 73 and 193 individuals ha(-1), which can be considered a low value compared with other continental pamphagid species.     

Host-herbivore studies of Stenoscepa sp. （Orthoptera：Pyrgomorphidae）, a high-Ni herbivore of the South African Ni hyperaccumulator Berkheya coddii （Asteraceae）

Nymphs of Stenoscepa sp. feed on leaves of the Ni hyperaccumulator Berkheya coddii at serpentine sites in Mpumalanga Province, South Africa. These sites contain Ni hyperaccumulators, Ni accumulators, and plants with Ni concentrations in the normal range. We conducted studies to： （i） determine the whole-body metal concentration of nymphs （including those starved to empty their guts）; （ii） compare Stenoscepa sp. nymphs against other grasshoppers in the same habitat for whole-body metal concentrations; and （iii）compare the suitability of Ni hyperaccumulator and Ni accumulator plants as food sources for Stenoscepa sp. and other grasshoppers. Stenoscepa nymphs had extremely high whole-body Ni concentrations （3 500μg Ni/g）. This was partly due to food in the gut, as starved insects contained less Ni （950 pg Ni/g）. Stenoscepa nymphs survived significantly better than other grasshoppers collected from either a serpentine or a non-serpentine site when offered high-Ni plants as food. In a host preference test among four Berkheya species （two Ni hyperaccumulators and two Ni accumulators）, Stenoscepa sp, preferred leaves of the Ni hyperaccumulator species. A preference experiment using leaves of three Senecio species （of which one species, Senecio coronatus, was represented by both a Ni hyperaccumulator and a Ni accumulator population） showed that Stenoscepa sp. preferred Ni accumulator Senecio coronatus leaves to all other choices. We conclude that Stenoscepa sp. is extremely Ni-tolerant. Stenoscepa sp. nymphs prefer leaves of hyperaccumulator Berkheya species, but elevated Ni concentration alone does not determine their food preference. We suggest that the extremely high whole-body Ni concentration of Stenoscepa nymphs may affect food web relationships in these serpentine communities.     

Alarm pheromone eliciting attack and escape responses in the sugar cane woolly aphid,Ceratovacuna lanigera (Homoptera,Pemphigidae)

All nymphal stages and apterous adults ofCeratovacuna lanigera secreted droplets containing an alarm pheromone from their abdominal cornicles when stimulated with a pin. When fresh droplets on piece of filter paper were put near the aphids, 1st-instar nymphs originating from apterous adults attacked it, but the advanced instar nymphs and adults escaped from the spot. Nymphs and adults originating from alate adults showed escaping behavior only. These different responses strongly correlated with morphological differences, such as relative lengths of the frontal horns and legs. When a syrphid larva,Allograpta javana, was placed on a colony of aphids, many 1st-instar nymphs attacked it using their frontal horns. During the attack, these nymphs spontaneously secreted droplets and adhered them to the syrphid body. When these droplets were placed on the body, the syrphid larva was agressively attacked by additional 1st-instar nymphs.     

The probing behaviour of nymphs of Vanduzeea arquata and Enchenopa binotata (Homoptera: Membracidae) on host and non-host plants

Abstract. 1. Nymphs of Vanduzeea arquata Say have been found to be more host-specific in nature and to show a higher degree of selectivity in host discrimination experiments than nymphs of Enchenopa binotata (Say), It was hypothesized that this differential selectivity would be reflected in the probing behaviour of individuals placed on twigs of host and non-host plants. Probing behaviour was examined by direct observation of nymphs and by sectioning and staining the probed plant tissues.
2. All nymphs probed readily and for extended periods on both host and non-host twigs. E.binotuta nymphs showed no consistent differences in probing behaviour on hosts versus non-hosts, but V.atquuta nymphs were more likely to withdraw their stylets within 60 s when on non-host twigs and produced honeydew only when on their host species. V.urquatu nymphs reached the phloem sieve elements only when on host twigs and broke many cells in peripheral plant tissue layers while probing. E.binotata nymphs broke few cells and often reached the phloem of non-host as well as host plants.
3. Nymphs of V.arquata always reject non-host plants, apparently in the course of probing and prior to encountering the phloem sap. Chemical compounds released from ruptured parenchyma cells may act as probing stimulants or inhibitors. E.binotura nymphs often feed on non-host plants in a non-choice situation; their preferential settling on host twigs in discrimination experiments may reflect a tendency to abandon non-host twigs more readily than host twigs.     

Relationship between physical leaf characteristics and growth and survival of polyphagous grasshopper nymphs, <Emphasis Type="Italic">Parapodisma subastris</Emphasis> (Orthoptera: Catantopidae)

Common types of plant defense mechanisms are thought to affect the host ranges of polyphagous herbivorous insects, yet few studies have examined the relationship between host plant suitability for polyphagous insects and defense against them. We investigated the suitability of the 19 plant species growing in the habitat of the polyphagous grasshopper, Parapodisma subastris, to determine the relationship between the physical characteristics of leaves and the growth and survival of grasshopper nymphs. We examined leaf toughness, trichome density, and length. Nymph survival was greater on plants with characteristics ranging from soft leaves and dense trichomes to tough leaves and few trichomes than on plants with soft leaves and few trichomes. The exception was Rorippa indica, a plant with soft leaves and few trichomes that uses biotic defense, on which nymph survival was maximal. Higher-quality plants that share common physical characteristics over families may favor polyphagy by grasshoppers that possess ability to overcome the physical defense easily with their robust mandibles.An erratum to this article can be found at     

Schistosoma mansoni: adult worm chemoattraction with barriers of specific molecular weight exclusions

Chemoattraction studies were done with Schistosoma mansoni adults from mice. To test for attraction or repulsion, some worm pairs were separated mechanically and the individuals placed in polycarbonate chambers. Experiments were done at 37 C and chambers contained dialysis tube chimneys. In all cases, heterosexual attraction occurred when one worm, but not two worms, were placed in the chimneys. Unperforated chimneys with specific molecular weight (Mr) exclusions were compared with perforated chimneys to study heterosexual attraction. Attraction was similar in designs using perforated chimneys and those with 50,000 and 12,000 Mr exclusions, but none was seen in chimney designs with 1000 Mr exclusions.     

On the mimetic association between nymphs of Hyalymenus spp. (Hemiptera: Alydidae) and ants

Nymphs of Hyalymenus , unlike adults, have a highly differentiated ant-like morphology. Both H. tarsatus and H. limbativentris feed mainly on reproductive parts of composites and solanaceous plants, respectively. Mimetic nymphs were observed on plants, together with ants, both day and night; adult Hyalymenus , however, are predominantly nocturnal. Ant-resemblance in nymphs is achieved by several structural adaptations which, when coupled with the mimic's zig-zag locomotion and constantly agitated antennae, produces a striking visual deception. Experiments in captivity showed that mimetic nymphs, but not adult Hyalymenus , are somewhat protected against attacks from the praying mantid Oxyopsis media found on their host plant. Colour and size changes through different nymphal instars of Hyalymenus allow the immature bugs to resemble, during their development, differently sized and coloured ant models. Similar-looking ant species seem to act as Mullerian mimics toward insectivorous vertebrates and invertebrates that avoid ants. It is suggested that nymphs of Hyalymenus gain Batesian protection by resembling available ant models of different Mullerian complexes. Density-dependent selection is thought to be responsible for the observed differences in mimetic morph proportions between populations of mimics, as well as for the mimetic strategy itself employed by nymphs of Hyalymenus.     

A comparison of the attraction of nymphs and adults of the ticksAmblyomma hebraeum andA. variegatum to carbon dioxide and the male-produced aggregation-attachment pheromone

The attraction of nymphs of the ticksAmblyomma hebraeum andA. variegatum to CO₂ alone and CO₂ together with the male-produced aggregation-attachment pheromone (AAP) was investigated. Matching experiments on the attraction of the adults of the two species were run for comparison. Nymphs ofA. hebraeum were strongly attracted to sources of CO₂ and CO₂/AAP at distances of 10 and 15 m. At distances of 20 and 25 m significantly more nymphs were attracted to CO₂/AAP than to CO₂ alone. Adults ofA. hebraeum were not attracted to CO₂ alone at any distance; these ticks were attracted to CO₂ and AAP together for distances of up to 25 m. The patterns of attraction of nymphs and adults ofA. variegatum at 10 m were similar to those recorded inA. hebraeum.     

Recognition of Novel Volatile Cues by the Nymphs of the Glassy-winged Sharpshooter, <Emphasis Type="Italic">Homalodisca vitripennis</Emphasis> (Cicadellidae)

Nymphs of Homalodisca vitripennis are strict xylophages and frequently switch host-plants to obtain a balanced level of nutrients at appropriate tension levels. Associative learning of odors and other semiochemicals may enhance the nymphs’ ability to recognize and locate potential host-plants. We examined whether nymphs could learn to associate a novel olfactory stimulus (vanilla) with a host-plant. Nymphs fed on cowpea plants treated systemically with vanilla extract or on untreated plants. Subsequently, their responses to a pale green target were measured in the presence or absence of vanilla odor. Nymphs were not innately attracted to pale green, and exposure to vanilla odor did not decrease a stereotypical scanning behavior. However, when presented with vanilla odor, significantly more conditioned nymphs selected the target than did control nymphs.     

Assessment ofPaulinia acuminata [Orthoptera: Acrididae] for the biological control ofSalvinia molesta in Australia

The biology and host specificity of the aquatic grasshopper,Paulinia acuminata (De Geer) were studied in quarantine in Australia. Adults and nymphs fed on the leaves of salvinia (Salvinia molesta), water lettuce (Pistia stratiotes) and azolla (Azolla pinnata). Fifty-three plant species representing 38 families were exposed to adults and nymphs ofP. acuminata. Adult feeding occurred on 17 plants but nymphs failed to feed on 9 of these species in the presence ofS. molesta. In starvation trials, 14 plants (excludingS. molesta, P. stratiotes andA. pinnata) were attacked by adults, of which only 5 were attacked by nymphs. Heavy feeding occurred on strawberry (Fragaria xananassa) but no oviposition occurred even when the leaves were held in contact with the water surface. The life cycle ofP. acuminata was completed only onS. molesta, P. stratiotes andA. pinnata. Eggs deposited on water hyacinth (Eichhornia crassipes) failed to hatch. Laboratory evaluation was supplemented with observations on the distribution and abundance ofP. acuminata on the Zambezi River system, Zimbabwe, during October 1984.     

New occurrence of the invasive alien leaf‐footed bug Leptoglossus gonagra (Hemiptera: Coreidae) in South Korea

The invasive alien leaf‐footed bug Leptoglossus gonagra (Fabricius 1775), native to the Americas, is herein recorded from South Korea for the first time. Forty nymphs and adults were collected on 22 September 2016 from arable lands of two separate sites. These individuals were identified based on the analysis of morphological characters and cytochrome oxidase c subunit 1 sequences. A neighbor‐joining tree indicated the existence of two clades: Group A from Brazil, and Group B from Asia and Oceania. Mean genetic distance between clades was 8.11%, range 7.60–8.57%. Within clades, mean genetic distance was 0.42% (0.17–0.83%) in Group A and, 0.45% (0.00–0.83%) in Group B. Colonies of 2nd‐final instar nymphs and adults were found, on top of leaves, flowers, fruits, and between the stems of pumpkins (Cucurbita moschata). Nymphs and adults damaged host plants by sucking their leaves, stems, fruits, and buds. After the first discovery in 2016, we were unable to find the bugs again (in 2017–2019). Therefore, the invasion of this species seems to have occurred only temporarily, during the warm season. We suggest that the species may have failed to overwinter as it is normally distributed in regions warmer than Korea. Nevertheless, possible appearances of this species must be continuously monitored as it is a potential pest of various plants cultivated in South Korea.     

Independent recruitment of a flavin-dependent monooxygenase for safe accumulation of sequestered pyrrolizidine alkaloids in grasshoppers and moths

Several insect lineages have developed diverse strategies to sequester toxic pyrrolizidine alkaloids from food-plants for their own defense. Here, we show that in two highly divergent insect taxa, the hemimetabolous grasshoppers and the holometabolous butterflies, an almost identical strategy evolved independently for safe accumulation of pyrrolizidine alkaloids. This strategy involves a pyrrolizidine alkaloid N-oxygenase that transfers the pyrrolizidine alkaloids to their respective N-oxide, enabling the insects to avoid high concentrations of toxic pyrrolizidine alkaloids in the hemolymph. We have identified a pyrrolizidine alkaloid N-oxygenase, which is a flavin-dependent monooxygenase, of the grasshopper Zonocerus variegatus. After heterologous expression in E. coli, this enzyme shows high specificity for pyrrolizidine alkaloids of various structural types and for the tropane alkaloid atropine as substrates, a property that has been described previously for a pyrrolizidine alkaloid N-oxygenase of the arctiid moth Grammia geneura. Phylogenetic analyses of insect flavin-dependent monooxygenase sequences suggest that independent gene duplication events preceded the establishment of this specific enzyme in the lineages of the grasshoppers and of arctiid moths. Two further flavin-dependent monooxygenase sequences have been identified from Z. variegatus sharing amino acid identities of approximately 78% to the pyrrolizidine alkaloid N-oxygenase. After heterologous expression, both enzymes are also able to catalyze the N-oxygenation of pyrrolizidine alkaloids, albeit with a 400-fold lower specific activity. With respect to the high sequence identity between the three Z. variegatus sequences this ability to N-oxygenize pyrrolizidine alkaloids is interpreted as a relict of a former bifunctional ancestor gene of which one of the gene copies optimized this activity for the specific adaptation to pyrrolizidine alkaloid containing food plants.     

Field biology of Edessa rufomarginata (Hemiptera: Pentatomidae): phenology, behavior, and patterns of host plant use

Pentatomids may cause direct and indirect damage to important crop plants. Biological and ecological features of phytophagous stink bugs in natural environments, however, remain poorly documented. Here, we provide an ecological account of Edessa rufomarginata De Geer on Caryocar brasiliense (Caryocaraceae) in the Brazilian savanna. The phenology of E. rufomarginata matched that of its host plant, with immatures developing in the wet season simultaneously with the production of vegetative and reproductive plant tissue. Females do not exhibit parental care and lay eggs more frequently on larger plants. Oviposition frequency, however, does not differ between plants with and without flowers/fruits. Nymphs and adults usually feed on stem parts and more rarely on flower buds and fruits. First- and second-instar nymphs remain aggregated, but disperse as third-instar nymphs. Adults and nymphs were more abundant on mature stems of C. brasiliense compared with other plant locations. Ants visiting the plant to search for extrafloral nectar occasionally tap the abdomen of E. rufomarginata nymphs with their antennae to obtain honeydew. This is the first record of trophobiotic interactions between Edessa stinkbugs and ants, and one of the few for heteropterans. The interaction of the stink bug with other natural enemies, such as predaceous Heniartes (Reduviidae), was also observed. Given the pest status of Edessa species for crop plants, additional field studies on host plants, interaction with ants, and natural enemies in native habitats are needed for an effective management of these stink bugs in tropical agricultural systems.