Susceptibility of twolined spittlebug (Hemiptera: Cercopidae) life stages to entomophagous arthropods in turfgrass

Prosapia bicincta (Say) (Hemiptera: Cercopidae), the twolined spittlebug, is an economic pest of turfgrass in the southeastern United States. No data concerning natural enemies of P. bicincta in turfgrass have been reported previously. We compared predation of spittlebug eggs, nymphs, and adults in the laboratory by potential generalist predators commonly found in turfgrass: bigeyed bugs Geocoris uliginosus Say and Geocoris punctipes Say; red imported fire ant, Solenopsis invicta Buren; wolf spiders (Lycosa sp. Walckenaer); carabid beetles Harpalaus pensylvanicus DeGeer and Calosoma sayi Dejean; and tiger beetles Megacephala carolina carolina L. Eggs were readily consumed by generalist predators. S. invicta consumed 100% of the eggs offered. H. pensylvanicus and C. sayi were also significant predators of P. bicincta eggs. Nymphs live in spittlemasses that protect them from attack by predators, but exposed nymphs were susceptible to attack when mechanically removed from their spittlemasses. S. invicta and M. carolina carolina caused significant mortality of exposed nymphs. P. bicincta adults are aposematic and have the ability to reflex bleed; however, reflex bleeding did not prevent attack by predators. S. invicta and M. carolina carolina killed 100% of the adult spittlebugs offered in laboratory bioassays. Lycosa sp. are less voracious predators of adults. Sound background knowledge about P. bicincta and its potential natural enemy complex is important for the development and implementation of a detailed, site-specific, biologically based pest management program in turfgrass.     

Bioactivity of neem, Azadirachta indica, against spittlebug Mahanarva fimbriolata (Hemiptera: Cercopidae) on sugarcane

The effect of neem, Azadirachta indica A. Juss, on some biological parameters of Mahanarva fimbriolata (Stil) (Hemiptera: Cercopidae) was studied in the laboratory by using NeemAzal-T/S, Nimkol-LS, and an aqueous neem seed extract. Initially, the LC,, was estimated for nymphs. Later, nymphs fed on sugarcane, Saccharum officinarum L., roots were sprayed with the respective LC,, for each product. The adults were maintained in cages on sugarcane plants sprayed at the base with the maximum rate recommended commercially for the crop (3 liter/ha). Moistened cotton discs surrounding the base of the plant were used as oviposition substrates. The LCso values estimated for NeemAzal, Nimkol, and aqueous extract were 0.014, 0.225, and 0.611%, respectively. There was a reduction in spittlebug longevity, regardless of sex, in relation to the control. Males exposed to the neem products, and aqueous extract showed longevity reductions of approximately 50%, whereas for females the reductions were 55-60%. The neem products and extract reduced fecundity by 75-85%. Morphological and physiological changes were observed in 9% of the eggs from individuals submitted to NeemAzal. Neem-based products, especially NeemAzal, have potential for the control of M. fimbriolata.     

Ventral pigmentation variation in the meadow spittlebug Philaenus spumarius (L.) (Hemiptera,Cicadomorpha: Cercopidae)

The meadow spittlebug Philaenus spumarius (L.) (Hemiptera, Cicadomorpha: Cercopidae) exhibits heritable colour/pattern variation on the dorsal and ventral sides of the adults. Eleven principal dorsal morphs are grouped as melanics and non-melanics according to the pigmentation of the dorsal surface. An investigation of ventral pigmentation variation in Philaenus spumarius was carried out with laboratory breeding stock obtained from Llysdinam, Wales. Analyses of the 1552 individuals, 753 females and 799 males, indicated that the pigmentation on the ventral surface of the adult individuals which vary significantly is associated with dorsal morph. Ventral parts of the dorsal melanics are usually darker than in non-melanics in both females (Kruskal-Wallis one-way ANOVA: H = 372.70, d.f. = 7, P < 0.001) and males (H = 407.36, d.f. = 7, P < 0.001). The combined “C” group morphs (flavicollis + gibbus + leuco- cephalus) are always darker than all other morphs in both sexes.     

Evaluation of turfgrass species and cultivars for potential resistance to twolined spittlebug (Hemiptera: Cercopidae)

Potential resistance to the twolined spittlebug, Prosapia bicincta (Say), was evaluated among 56 turfgrass genotypes. Greenhouse, laboratory, and field bioassays identified differences in spittlebug survival and development, host preference and damage levels, and turfgrass tolerance to and ability to recover from pest induced injury. All centipede grasses demonstrated high levels of susceptibility, followed by bermudagrasses, seashore paspalums, and zoysiagrasses. Average nymphal survival to the adult stage ranged from 1.5 to 78.1%. Development required 38.1-62.0 d under greenhouse conditions, depending on plant taxa. Among seashore paspalums, nymphal survival to the adult stage was lowest and duration of development was longest on HI-1, 'Sea Isle 2000', 561-79, and 'Mauna Kea'. Reduced spittlebug survival and increased developmental times were also observed on the bermudagrasses BERPC 91-15 and 'Tifway'. Although zoysiagrasses supported spittlebug development and survival to the adult stage, developmental times were extended on the zoysiagrass cultivars 'Emerald' and 'El Toro'. Spittlebug preference varied with generation evaluated. First-generation spittlebugs inflicted the greatest damage on TC201 (centipede grass), 'Primavera' (bermudagrass), and 'Emerald' (zoysiagrass) in choice tests. In the fall, second-generation spittlebugs damaged TC201 (centipedegrass) and 'Sea Isle 1' (paspalum) most severely, whereas 561-79 (paspalum) and 'Emerald'(zoysiagrass) were less severely affected. Among taxa included in field trials, HI-1, 'Mauna Kea', 'Sea Isle 2000',and AP-14 paspalums, 'Tifway' bermudagrass, and 'Emerald' zoysiagrass were most tolerant (demonstrated the best regrowth potential following twolined spittlebug feeding).     

Risk predictive model for damage caused by the spittlebug Aeneolamiapostica (Walker) Fennah (Hemiptera: Cercopidae)

This paper evaluated the risk that Aenolamia postica (Walker) Fennah populations reach the economic threshold in sugar cane fields in Veracruz, México. A risk deductive model was constructed to include the sequence of events leading to damaging populations, considered the top event or critical failure in the crop. Model events were identified and quantified, and model was validated on field conditions. The model components and their state values were identified as: temperature e" 28 degrees C, precipitation e" 45% during June and July, soil clay content e" 40%, infested adjoining fields, deficient weed control, wind dominance, crop phenology and variety, deficient chemical and biological control, and irrigation. Sensitivity analysis showed that the most important events triggering high densities of A. postica were high temperatures and precipitation, previous field infestation, nymph and weed presence. Event probability estimates were combined using Boolean algebra to compute the minimum, mean and maximum probabilities for the top event, yielding values of 0.417, 0.563, y 0.734 respectively. Model was tested in field, by selecting sugar cane fields having the model properties and compared to fields without these features. Fields were sampled in both conditions during 2004 year and high-risk fields had significantly (F = 13, 4, gl = 1, 18, P = 0,0018) higher densities (2.4 adults m(-1)) than low-risk plots (0.4 adults m(-1)) thus agreeing with the model forecast.     

The role of biofoam in shielding spittlebug nymphs (Insecta,Hemiptera, Cercopidae) against bright light

1. Biofoam in spittlebugs has traditionally been seen as a defence against predation and a microclimate that reduces the risks of overheating and drying out. This study addresses the possible role of the foam as a light attenuator. 2. Nymphs exhibit higher mortalities when reared under brighter light (1800–2000 lux) than under less bright light (600–800 lux). At all developmental stages, photoavoidance is strongest when the nymphs are depleted of foam covers. First‐ and second‐instar nymphs appear to be the most vulnerable to exposures by bright light. 3. Smaller bubbles are more effective as light attenuators than are larger ones. As younger instars possess smaller canals from which bubbles are released and, furthermore, exhibit higher concentrations of proteins in bubble liquid, they can produce smaller‐sized bubbles and their foams are more effective at reducing light than are those of older nymphs. 4. The findings of this study show that most of the visible and UV radiation is reflected by the foam: transmittance of visible light was 15% at 600 nm and 12% at 350 nm. 5. These results demonstrate that spittlebug foam also possesses properties that render it an effective barrier against potentially damaging solar radiation.     

Sublethal effects of antibiosis resistance on the reproductive biology of two spittlebug (Hemiptera: Cercopidae) species affecting Brachiaria spp

Several greenhouse experiments were used to measure how high levels of antibiosis resistance to nymphs in two interspecific Brachiaria (brachiariagrass) hybrids affect life history parameters of the spittlebugs Aeneolamia varia (F.) and Zulia carbonaria (Lallemand), two of the most important spittlebug (Hemiptera: Cercopidae) species affecting Brachiaria production in Colombia. The A. varia-resistant hybrid CIAT 36062, the Z. carbonaria-resistant hybrid SX01NO/0102, and the susceptible accession CIAT 0654 were used to compare the effect of all possible combinations of food sources for nymphs and adults. Calculation of growth indexes showed a significant impact of antibiosis resistance on the biology of immature stages of both species. Median survival times of adults feeding on resistant genotypes did not differ from those recorded on the susceptible genotype, suggesting that factors responsible for high mortality of nymphs in the resistant hybrids did not affect adult survival. Rearing nymphs of A. varia on CIAT 36062 and of Z. carbonaria on SX01NO/0102 had deleterious sublethal effects on the reproductive biology of resulting adult females. It is concluded that high nymphal mortality and subsequent sublethal effects of nymphal antibiosis on adults should have a major impact on the demography of the two spittlebug species studied.     

Functional response of the tiger beetle Megacephala carolina carolina (Coleoptera: Carabidae) on twolined spittlebug (Hemiptera: Cercopidae) and fall armyworm (Lepidoptera: Noctuidae)

The functional response of the tiger beetle Megacephala carolina carolina L. (Coleoptera: Carabidae) was determined on adult twolined spittlebug, Prosapia bicincta (Say) (Hemiptera: Cercopidae), and fourth instars of fall armyworm, Spodoptera frugiperda (J.E. Smith) (Lepidoptera: Noctuidae), in single-prey and two-prey systems. In the laboratory, M. carolina carolina demonstrated a type II functional response for P. bicincta and S. frugiperda in both single- and two-prey systems. Search efficiency of M. carolina declined for both prey as the initial number of prey increased. Of the total prey consumed, M. carolina carolina killed significantly more S. frugiperda than P. bicincta in the single-prey system (8.0 and 4.5, respectively) and the two-prey system (5.0 and 2.0, respectively). Estimates of attack coefficient, a, were not significantly different for P. bicincta and S. frugiperda in the single-prey (0.07 and 0.02) and two-prey systems (0.04 and 0.06), respectively. The handling time, T(h), was significantly greater for P. bicincta (5.02 and 10.64 h) than for S. frugiperda (2.66 and 4.41 h) in single- and two-prey systems, respectively. Estimations of attack coefficient and handling time in the single-prey system were used to predict prey preference of M. carolina carolina. No strong prey switching response was observed. M. carolina carolina showed no preference for either prey. However, in the presence of S. frugiperda, the functional response of the predator for P. bicincta was reduced. M. carolina carolina is a potential predator of one or more turfgrass pests and should be considered in conservation efforts.     

Application methods of entomopathogenic nematodes for control of Mahanarva spectabilis (Hemiptera: Cercopidae)

The pathogenicity of entomopathogenic nematodes (EPNs) against nymphs of the pasture spittlebug Mahanarva spectabilis (Distant, 1909) (Hemiptera: Cercopidae) was studied under four application methods. Nymphs of the fourth or fifth instar were placed on the roots of elephant grass plants and submitted to four EPN isolates (Steinernema carpocapsae, S. feltiae, S. riobravis and Heterorhabditis amazonensis RSC1), at two concentrations (2000 and 4000 infective juveniles/mL), with four application methods (pipetting, spraying on the nymphs after froth formation, spraying before froth formation and placement of infected host cadaver method). There was no significant difference in the nymph mortality in function of the concentration and/or the isolate used. However, the efficacy was influenced by the application method, with the most efficient being sprayed on nymphs after froth formation and infected host cadavers. Steinernema riobravis applied by spraying on nymphs with froth, at a concentration of 2000 EPNs/mL, and H. amazonensis RSC1 applied by infected host cadavers caused 71% of the nymphs to die. The use of infected host cadavers and spraying in an aqueous solution on nymphs after froth formation were found to be the most efficient methods to control Mahanarva spectabilis.     

Morphological and molecular identification of Prosapia simulans (Walker) (Hemiptera: Cercopidae), and screening and mechanisms of resistance to this spittlebug in Brachiaria hybrids

Prosapia simulans (Walker) is an important spittlebug species that attacks forage grasses of the genus Brachiaria (Trin.) Griseb. from Mexico to Colombia. This, and several other species of spittlebugs, cause important economic losses to the livestock production industry. Development of resistant cultivars is regarded as the best method of control. In the present study we used taxonomic keys, dissection of male genitalia and RAPD-PCR patterns to reconfirm the identity of P. simulans specimens collected in Colombia and Mexico. We were able to reconfirm that P. simulans occurs as a pest of Brachiaria from Mexico to Colombia. We also studied the levels and mechanisms of resistance present in 34 Brachiaria hybrids developed by CIAT. Infestations were made with six eggs per plant. We used 10 replications (plants) per genotype in a completely randomized design. Seven hybrids were found to be susceptible, 16 showed intermediate resistance and 11 were resistant. Antibiosis was the mechanism of resistance expressed in resistant hybrids as well as in the resistant checks CIAT 6294 and CIAT 36062. Tolerance was absent. The genotypes BRX 4402 and CIAT 0606 were classified as highly susceptible.     

Mating success in the spittlebug Cercopis sanguinolenta (Scopoli, 1763) (Homoptera, Cercopidae): the role of body size and mobility

In insects, a sexual size dimorphism commonly occurs, with larger females. However, as a deviation from this general rule, larger males are found in some species. In these species often sexual selection for large males has been presumed. The spittlebug Cercopis sanguinolenta exhibits a distinct sexual size dimorphism with larger males. Mating behaviour was studied in a field population in respect to mating success of males and females. The aim of this study was to examine the mechanisms that lead to the observed non-random mating pattern. The results showed a mating pattern without size-assortative mating. A correlation was found between mating success and body size in males. In females no such correlation was found. The mobility of males depends on their body size and mobility is high only when females are present. However, in an analysis of covariance it was found that male mating success is not correlated with mobility, when controlled for body size. The mating system of the spittlebug was classified as scramble competition polygyny. Electronic Publication     

Mating Behavior of Mahanarva spectabilis (Hemiptera: Cercopidae) Under Laboratory Conditions

The mating behavior of Mahanarva spectabilis (Distant) was studied under laboratory conditions. Virgin adults were individually maintained in transparent cages for observation of the mating sequence. Mating behavior of 23 couples was monitored during the photophase portion of the day. Copulation was observed in 17 of the 23 pairs. The mating sequence includes male approaching the female, mounting of the female, and copulation. Of the 17 mating pairs, 70.6 % mated on the second day after emergence. Most of the copulations began between 8 and 9 h after the onset of photophase. The average duration of copulation was 268?±?24.9 min, and most pairs mated once or twice during their lifetimes. The mating behavior of M. spectabilis is important to understand because it determines the age and ideal time for further behavioral testing, which is essential for determining the cues involved in the communication system of the species.     

Effect of temperature on the life history and demographic parameters of Spalgis epius (Lepidoptera: Lycaenidae), a candidate biological control agent of mealybugs (Hemiptera: Pseudococcidae)

Many species of mealybugs (Hemiptera: Pseudococcidae) are serious pests of economically important crops worldwide. We evaluated the influence of constant temperatures: 14, 16, 18, 20, 22, 24, 26, 28, 30, 32 and 34°C on the life history and demographic parameters of Spalgis epius (Lepidoptera: Lycaenidae), a candidate biological control agent of various species of mealybugs. No eggs completed their development at 14 and 34°C. Egg-to-adult developmental time significantly decreased from 89.9 days at 16°C to 20.4 days at 32°C. The estimated lower temperature threshold of 10.2°C and 416.6 degree-days were required to complete egg-to-adult development. The mortality of immature stages was maximum at 16 and 32°C and minimum at 28°C. The highest lifetime fecundity was recorded at 28°C and it significantly decreased at 32°C. The longevity of adults was about three times more at 16°C than at 30 and 32°C. The net reproductive rate (R ₀) significantly increased with increased temperatures up to 28°C and significantly decreased at 32°C. The mean generation time (T) significantly decreased with increased temperature up to 30°C, but it significantly increased at 32°C. The intrinsic rate of population increase (r _m ) was highest at 30°C. The finite rate of increase (λ) was significantly greater at 30°C than at other temperatures. These data suggest that S. epius can develop, reproduce and survive in a wide range of temperatures and thus could be regarded a potential biological control agent of mealybugs.     

Effect of temperature and cultivar on pea aphid, Acyrthosiphon pisum (Hemiptera: Aphididae) life history

Life history parameters of the pea aphid, Acyrthosiphon pisum were studied at five constant temperatures on two cultivars of peas, Scout and Sancho. The development and mortality of juveniles and the life-span, age-specific fecundity and survivorship of adult aphids were recorded and used to construct life tables. The juvenile development period (from birth to adulthood) was longest at 11.9 degrees C (16.8 days on cv. Scout and 16.2 days on cv. Sancho) and shortest at 26.7 degrees C (8.5 days on cv. Scout and 8.8 days on cv. Sancho). At all temperatures, except 26.7 degrees C, juveniles developed faster on cv. Sancho than on cv. Scout. On both pea varieties juvenile mortality was highest at temperatures above 19.6 degrees C and lowest at 19.6 degrees C. Highest cumulative juvenile mortality was recorded on cv. Scout at 26.7 degrees C when only 9% of aphids survived from birth to reproductively mature adults. Fecundity rates were unaffected by temperature in the range tested on cv. Sancho but increased with increasing temperatures between 11.9 and 19.6 degrees C on cv. Scout. These differences in life history parameters were reflected in the population growth (rm) of aphids on both pea cultivars which increased with increasing temperatures between 11.9 and 23.1 degrees C on cv. Sancho and 11.9 and 19.6 degrees C on cv. Scout, declining thereafter. Population growth was consistently greater at all temperatures for aphids reared on cv. Sancho than those reared on cv. Scout.     

Life history,diagnosis, and biological aspects of Engytatus varians (Hemiptera: Miridae), a predator of Bactericera cockerelli (Hemiptera: Triozidae)

The life cycle and several life parameters of the zoophytophagous predator Engytatus varians (Distant) (Heteroptera: Miridae), including nymphal growth according to Dyar’s rule, were examined in the laboratory. The egg, nymph (five instars), and adult stages were 9.20, 17.36, and 19.02?d in length, respectively. The growth ratio for nymphs was consistent with Dyar’s rule based on the lengths of the femora of the forelegs, the tibiae and femora of the middle legs, and the antennae. Some biological characteristics of E. varians were also evaluated when the mirid was fed three different diets (B. cockerelli third instars, Sitotroga cerealella Olivier [Lepidoptera: Gelechiidae] eggs, and a mixture of both instars and eggs). The length of the nymphal stage was three days longer on a diet of only S. cerealella eggs than when the mirids were fed the third instars of B. cockerelli only or a mixture of both. The sex ratio was not affected by the type of diet. Nymphs of E. varians consumed B. cockerelli nymphs (80–85) when fed third instars only and third instars?+?S. cerealella eggs, respectively. The potential use of this predator as a biological control agent of B. cockerelli is discussed.     

Morphology of the mouthparts of the spittlebug Philagra albinotata Uhler (Hemiptera: Cercopoidea: Aphrophoridae)

Mouthparts associated with feeding behavior and feeding habits are important sensory and feeding structures in insects. To obtain a better understanding of feeding in Cercopoidea, the morphology of mouthparts of the spittlebug, Philagra albinotata Uhler was examined using scanning electron microscopy. The mouthparts of P. albinotata are of the typical piercing–sucking type found in Hemiptera, comprising a cone-shaped labrum, a tube-like, three-segmented labium with a deep groove on the anterior side, and a stylet fascicle consisting of two mandibular and two maxillary stylets. The mandibles consist of a dorsal smooth region and a ventral serrate region near the apical half of the external convex region, and bear five nodules or teeth on the dorsal external convex region on the distal extremity; these are regarded as unique features that distinguish spittlebugs from other groups of Hemiptera. The externally smooth maxillary stylets, interlocked to form a larger food canal and a smaller salivary canal, are asymmetrical only in the internal position of longitudinal carinae and grooves. One dendritic canal is found in each maxilla and one in each mandible. Two types of sensilla trichodea, three types of sensilla basiconica and groups of multi-peg structures occur in different locations on the labium, specifically the labial tip with two lateral lobes divided into anterior sensory fields with ten small peg sensilla arranged in a 5 + 4 + 1 pattern and one big peg sensillum, and posterior sensory fields with four sensilla trichodea. Compared with those of previously studied Auchenorrhyncha, the mouthparts of P. albinotata may be distinguished by the shape of the mandibles, the multi-peg structures and a tooth between the salivary canal and the food canal on the extreme end of the stylets. The mouthpart morphology is illustrated using scanning electron micrographs, and the taxonomic and putative functional significance of the different structures is briefly discussed.     

Spittlebugs (Hemiptera: Cercopidae) and their host plants: a strategy for pasture diversification

Information about the biology of spittlebugs on different species of forage is necessary for the development of tactics for their control, and is also essential for evaluating, selecting and launching new forage cultivars. The aims of this study were to compare the nutritional suitability of different forage species–Brachiaria ruziziensis (Congo grass), Pennisetum purpureum (elephant grass), Panicum maximum (Guinea grass), and Cynodon dactylon (Bermuda grass)–for the spittlebug species Mahanarva spectabilis (Distant) (Hemiptera: Cercopidae), Deois schach (Fabricius) (Hemiptera: Cercopidae) and Notozulia entreriana (Berg) (Hemiptera: Cercopidae), and to evaluate the performance of each spittlebug species on forage monocultures. B. ruziziensis was the most suitable host plant for D. schach. Since N. entreriana did not show differences in the evaluated parameters, it was not possible to determine its ideal forage species. For M. spectabilis, P. purpureum was the most suitable forage species. The evaluation of the performance of the different spittlebugs on a single forage species also showed which spittlebugs are a problem. M. spectabilis and D. schach developed better on a monoculture of B. ruziziensis. M. spectabilis was a major problem when it fed on P. purpureum, but on C. dactylon and P. maximum, none of the spittlebugs developed properly, indicating that these plants are less suitable for them. These results suggest that the forage diversification of pastures may be a strategy for the integrated management of spittlebugs.     

Antifungal effect and chitinase activities of the froth of spittlebug Poophilus costalis (Walker) (Hemiptera: Cercopoidea: Aphrophoridae)

Nymphs of xylophagous spittlebug Poophilus costalis (Walker, 1851) produce froth that covers their entire body surface. However, no mold has ever been found on this froth under natural conditions. This study has shown that the froth of P. costalis can repress the growth of Fusarium oxysporum f. sp. pisi. Three chitinases, i.e., β-N-acetylglucosaminidase, chitobiosidase, and endochitinase, with specific activities of 1.759, 0.365, and 1.172 U/mg protein, respectively, and a protease with minimal activity have been detected in this froth. β-N-actylglucosaminidase activity was also detected in the source of the froth, the nymphal Malpighian tubules and the host plant Commelina diffusa Burm. f. These findings suggest that the antifungal effect of the froth from P. costalis may be attributed to the chitinase produced by its Malpighian tubules and/or its host plant.