The Damage Capacity of Mahanarva spectabilis (Distant, 1909 ...

7
Hindawi Publishing Corporation e Scientific World Journal Volume 2013, Article ID 281295, 6 pages http://dx.doi.org/10.1155/2013/281295 Research Article The Damage Capacity of Mahanarva spectabilis (Distant, 1909) (Hemiptera: Cercopidae) Adults on Brachiaria ruziziensis Pasture Tiago Teixeira Resende, 1 Alexander Machado Auad, 1 Marcy das Graças Fonseca, 1 Fausto Souza Sobrinho, 1 Dayane Ribeiro dos Santos, 1 and Sandra Elisa Barbosa da Silva 2 1 Embrapa Dairy Cattle Research Center, Rua Eugˆ enio de Nascimento, 610 Dom Bosco, 36038-330 Juiz de Fora, MG, Brazil 2 Depatment of Entomology, Federal University of Lavras, Campus Universit´ ario, 37200-000 Lavras, MG, Brazil Correspondence should be addressed to Alexander Machado Auad; [email protected] Received 6 September 2013; Accepted 27 October 2013 Academic Editors: G. E. Brust and C. Dell Copyright © 2013 Tiago Teixeira Resende et al. is is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. e aim of this study was to determine the damage caused by adult Mahanarva spectabilis (Distant, 1909) (Hemiptera: Cercopidae) on Brachiaria ruziziensis (Germain & Evard) under field conditions. A total of 0, 4, 8, 12, or 16 M. spectabilis adults per plot were maintained for 6 days. ereaſter, the insects were removed from the plant, and the following parameters were evaluated: chlorophyll content, damage score, dry as well as fresh weights, percentage of shoots’ dry matter, and the forage’s ability to regrow. e chlorophyll content was significantly reduced; the damage score and percentage of dry matter in plants increased depending on the increased insect infestation density aſter 6 days of exposure. In contrast, no change was observed on the B. ruziziensis fresh and dry weights as well as the regrowth capacity depending on the M. spectabilis infestation densities. Attacks by 8 adult M. spectabilis per clump of B. ruziziensis with an average of 80 tillers for 6 days were sufficient to reduce the chlorophyll content and the functional plant loss index. is density can be a reference for spittlebug integrated management in Brachiaria. 1. Introduction In the 1970s due to the emergence of forage species with a high adaptability to climate and low soil fertility, livestock was expanded in Brazil [1]. Currently, cattle livestock is responsible for over 44% of the national cattle herd, which use grown pastures as their primary food source [2]. Extensive monoculture of such grasses favors development of high spittlebug populations [3]. Spittlebugs from the genus Mahanarva (Hemiptera: Cer- copidae) cause serious damage in pastures and threaten meat production by compromising the forage supply [4, 5]. ey have been reported in several regions and are widely dis- tributed in south and central America [6]. e attacks by spittlebugs may even kill the grass depending on the time of the year and population density [7]. According to ompson [8], the damage caused by spittlebugs generates losses of 840 to 2100 million dollars per year worldwide. Even though nymphs cause damage to forage, adult damage is more severe due to the toxic salivary excretions leſt in the shoots during feeding [914]. According to Byers and Wells [9], the toxic saliva injected during adult feeding interferes in the photosynthetic activity. Val´ erio and Nakano [11] observed a direct relationship between exposure time and damage symptom intensity in Brachiaria decumbens that were under attack by adult Notozu- lia entreriana (Berg 1879). Adopting a simulation model to quantify the economic impact of adult N. entreriana, Holmann and Peck [15] observed a drastic reduction in B. decumbens carrying capacity and estimated that 10 adult spittlebugs can reduce the cultivated pasture stocking rate, which significantly contributes to increased production costs. Research has been directed towards spittlebug control through forage resistance to nymphs. However, L´ opez et al. [14] concluded that adult spittlebugs are a major threat to Brachiaria hybrids with high levels of antibiosis resistance to the nymphs.

Transcript of The Damage Capacity of Mahanarva spectabilis (Distant, 1909 ...

Hindawi Publishing CorporationThe Scientific World JournalVolume 2013 Article ID 281295 6 pageshttpdxdoiorg1011552013281295

Research ArticleThe Damage Capacity of Mahanarva spectabilis(Distant 1909) (Hemiptera Cercopidae) Adults onBrachiaria ruziziensis Pasture

Tiago Teixeira Resende1 Alexander Machado Auad1 Marcy das Graccedilas Fonseca1

Fausto Souza Sobrinho1 Dayane Ribeiro dos Santos1 and Sandra Elisa Barbosa da Silva2

1 Embrapa Dairy Cattle Research Center Rua Eugenio de Nascimento 610 Dom Bosco 36038-330 Juiz de Fora MG Brazil2 Depatment of Entomology Federal University of Lavras Campus Universitario 37200-000 Lavras MG Brazil

Correspondence should be addressed to Alexander Machado Auad alexanderauadembrapabr

Received 6 September 2013 Accepted 27 October 2013

Academic Editors G E Brust and C Dell

Copyright copy 2013 Tiago Teixeira Resende et al This is an open access article distributed under the Creative Commons AttributionLicense which permits unrestricted use distribution and reproduction in any medium provided the original work is properlycited

The aim of this study was to determine the damage caused by adultMahanarva spectabilis (Distant 1909) (Hemiptera Cercopidae)on Brachiaria ruziziensis (Germain amp Evard) under field conditions A total of 0 4 8 12 or 16 M spectabilis adults per plotwere maintained for 6 days Thereafter the insects were removed from the plant and the following parameters were evaluatedchlorophyll content damage score dry as well as fresh weights percentage of shootsrsquo dry matter and the foragersquos ability to regrowThe chlorophyll content was significantly reduced the damage score and percentage of drymatter in plants increased depending onthe increased insect infestation density after 6 days of exposure In contrast no change was observed on the B ruziziensis fresh anddry weights as well as the regrowth capacity depending on theM spectabilis infestation densities Attacks by 8 adultM spectabilisper clump of B ruziziensiswith an average of 80 tillers for 6 days were sufficient to reduce the chlorophyll content and the functionalplant loss index This density can be a reference for spittlebug integrated management in Brachiaria

1 Introduction

In the 1970s due to the emergence of forage species with ahigh adaptability to climate and low soil fertility livestockwas expanded in Brazil [1] Currently cattle livestock isresponsible for over 44of the national cattle herd which usegrown pastures as their primary food source [2] Extensivemonoculture of such grasses favors development of highspittlebug populations [3]

Spittlebugs from the genusMahanarva (Hemiptera Cer-copidae) cause serious damage in pastures and threaten meatproduction by compromising the forage supply [4 5] Theyhave been reported in several regions and are widely dis-tributed in south and central America [6] The attacks byspittlebugs may even kill the grass depending on the time ofthe year and population density [7] According toThompson[8] the damage caused by spittlebugs generates losses of 840to 2100 million dollars per year worldwide Even thoughnymphs cause damage to forage adult damage is more

severe due to the toxic salivary excretions left in the shootsduring feeding [9ndash14] According to Byers and Wells [9] thetoxic saliva injected during adult feeding interferes in thephotosynthetic activity

Valerio and Nakano [11] observed a direct relationshipbetween exposure time and damage symptom intensity inBrachiaria decumbens thatwere under attack by adultNotozu-lia entreriana (Berg 1879) Adopting a simulation modelto quantify the economic impact of adult N entrerianaHolmann and Peck [15] observed a drastic reduction in Bdecumbens carrying capacity and estimated that 10 adultspittlebugs can reduce the cultivated pasture stocking ratewhich significantly contributes to increased production costs

Research has been directed towards spittlebug controlthrough forage resistance to nymphs However Lopez et al[14] concluded that adult spittlebugs are a major threat toBrachiaria hybrids with high levels of antibiosis resistance tothe nymphs

2 The Scientific World Journal

According to Kain and Atkison [16] the main problemassociated with studies that assess loss promoted by insectsin pastures is converting quantitative and qualitative loss ofgrassland into livestock production and consequently estab-lishing economic damage level In this context determiningeffects of adult M spectabilis attacks on B ruziziensis is animportant tool that will assist in an integratedmanagement ofthis insect pest Thus the aim of this study was to determinethe damage caused by adult M spectabilis on B ruziziensisunder field conditions

2 Materials and Methods

21 Plants and Insects The research was conducted at theEmbrapa Dairy Cattle research experimental field which islocated in Coronel Pacheco MG Brazil (21∘3310158402210158401015840 southand 43∘1610158401510158401015840 west) During the experiment the meantemperature was 278∘C (maximum 464∘C and minimum202∘C) with a 769 relative humidity (maximum 982and minimum 243) These parameters were recorded bya data logger every 2 minutes and transferred to software(Hoboware) these values were used to determine the averagefor the experimental period Twenty days before beginningthe experiment the B ruziziensis pasture was cut 15 cm aboveground level to standardize plant height Each experimentalplot included a forage clump with a 30 cm mean height8852 tillers and 3323 SPAD units (chlorophyll content) Mspectabilis adults were collected in a pasture distal to theexperimental area These insects were maintained in cagesuntil they were used in the experiment

22 Experiment The randomized block design included 5infestation densities and 8 repetitions Each plot compriseda B ruziziensis clump protected by a metal frame cage (70 times40 times 40 cm) covered with organza fabric The experimentalplots were installed equidistant (5 times 2m) and covered with400m2 meters of grassland Inside each cage were 0 4 8 12and 16M spectabilis adultsThedead insectswere replenisheddaily to maintain a constantM spectabilis density for 6 daysThereafter the insects were removed and evaluated for thefollowing parameters chlorophyll content damage score dryas well as fresh weights percentage of dry shoot matter andthe forage regrowth ability

The chlorophyll content was measured using the appara-tus Minolta SPAD 502 OL (Konica Minolta Sensing OsakaJapan) before infestation as well as 3 and 6 days after onsetof infestation in three leaf blades from one tiller of the plantAccording to Diaz-Montano et al [17] the chlorophyll meterSPAD-502 is an important device used to measure chloro-phyll loss by sucking insects The mean chlorophyll contentin the three leaf blades from each plant was calculated

After 6 days of infestation the damage percentage score inthe shoots from each plant was assigned by three evaluatorsthat followed the damage scale from 1 through 5 proposedby Cardona et al [18] Furthermore the damage scores wereclassified based on scores developed by Pabon et al [19]as follows scores 1 and 2mdashthe grass tolerates insect attackscores between 21 and 3mdashintermediate tolerance and scoresover 3mdashsusceptible to insect attack

After 6 days the plants with different infestation densitieswere cut at the soil level and their leaves and stems wereweighed for the fresh weight These materials were dried at55∘C for 72 hours and after this period they were weighed torecord the dry weight The percentages were then calculatedfor the dry matter and the functional plant loss index (FPLI)which were proposed by Morgan et al [20] and modifiedby Panda and Heinrichs [21] This index is calculated basedon the damage score (DS) dry weight of uninfested plants(DWUP) and dry weight of infested plants (DWIP) asfollows FPLI () = [1 minus (DWIPDWUP) times (1 minusDS5)] times 100and this method is considered a useful tool for quantifyingtolerance [22]

To evaluate the forage regrowth ability with adult spit-tlebugs the metal frame of each cage was maintained for 30days without organza fabric covering The number of shootsin each clump was evaluated 15 and 30 days after cutting

23 Statistical Analysis The average chlorophyll content inthe three leaf blades from each plant damage score freshand dry shoot weights dry matter percentage and forageregrowth ability were compared using analysis of varianceand when significant (119875 le 005) regression analyses wereperformed for the spittlebug infestation density The averagechlorophyll content was compared using the Tukey test toassess the effect of 3 and 6 days of forage exposure to adultspittlebugs (119875 le 005)

The analyses were performed using the program SISVAR51 [23] (Federal University of Lavras MG Brazil) ThePearson correlation between the chlorophyll content anddamage score was generated using BioEstat 50 [24] (FederalUniversity of Para Para Brazil)

3 Results

In the first evaluation which was conducted prior to forageinfestation by M spectabilis adults the chlorophyll contentwas the same (119865 = 0191 119875 = 09427) regardless of the futureinfestation level which demonstrates plant standardizationThis standardized plant behavior was reproduced in thesecond evaluation which was performed 3 days after theplants were exposed to insects (119865 = 213 119875 = 008)In the subsequent evaluation which was performed 6 daysafter infestation the forage chlorophyll content significantlydecreased due to an increase in theM spectabilis infestation(119865 = 728 119875 lt 00001) which indicates a quadratic regres-sion curve for this exposure time (Figure 1)

After the exposure the plant chlorophyll content did notchange in plants exposed to 4 adult spittlebugs (119865 = 109 119875 =033) and those with no contact to the insect pest (119865 = 225119875 = 010) In contrast there was a significant reduction inforage chlorophyll content after 6 days of exposure to 8 (119865 =443 119875 lt 001) and 12 insects (119865 = 289 119875 lt 005) as wellas for plants exposed for 3 days to 16 insects (119865 = 742 119875 lt0001) (Figure 2)

For the damage score higher M spectabilis infestationlevels yielded greater damage to plants exposed to spittlebugsfor 6 days (119865 = 1853 119875 lt 00001) (Figure 3) Notably underfield conditions B ruziziensis had an intermediate tolerance

The Scientific World Journal 3

20

24

28

32

36

40

0 4 8 12 16Adults of M spectabilis per plant

Chlo

roph

yll c

onte

nt y = 00652x2 minus 16903x + 36491

R2 = 09354

Figure 1 Relationship between infestation levels of adult Mspectabilis and chlorophyll content (SPAD unit) in B ruziziensis

05

1015

1216

2025303540

4

45

0

0

3 6

8

Time (days)

a a a a a a a ab abb

aa

b b b

Chlo

roph

yll c

onte

nt

Adults of M spectabilis per plant

Figure 2 Relationship between chlorophyll content (SPAD unit)of B ruziziensis and exposure time (0 3 and 6 days) at differentinfestation levels of adultM spectabilis Mean values followed by thesame letter within the levels of infestation did not differ by Tukeytest

to be attacked by 4 8 12 and 16M spectabilis adults for 6 daysbased on converting the damage score parameters into thetolerance established by Pabon et al[19] Further the damagescores inversely correlate with the forage chlorophyll contentupon a day exposure to M spectabilis adults (119903 = minus044119905 = minus3348 119875 = 00018)

For the functional plant loss index there was no signifi-cant difference in this ratio among infestation densities (119865 =1613 119875 = 0208) However only four and eight adult Mspectabilis per plant were sufficient to generate a functionalplant loss index greater than 453 and 60 respectivelyin B ruziziensis however for 16 adults such a loss reached619 Likewise increasing M spectabilis densities did notsignificantly alter the fresh (119865 = 206 119875 = 0113) and dryweights (119865 = 1416119875 = 02546) in plants exposed to the adultinsects for 6 days However the increased infestation densitycaused a significant increase in the dry matter percentage ofplant (119865 = 2798 119875 = 00451) (Figure 4)

The tiller number in plants was equal in the assessmentsperformed 15 (119865 = 0565 119875 = 069) and 30 (119865 = 0432

0

1

2

3

4

5

Dam

age s

core

y = minus00072x2 + 02187x + 11166

R2 = 09376

0 4 8 12 16Adults of M spectabilis per plant

Figure 3 Relationship between infestation levels of M spectabilisadults and damage scores for B ruziziensis over 6 days

20

21

22

23

24

25

26

R2 = 08974

0 4 8 12 16Adults of M spectabilis per plant

Dry

mas

s per

cent

age y = 00122x2 + 00351x + 21824

Figure 4 Relationship between infestation levels of M spectabilisadults and dry mass percentage of the B ruziziensis

119875 = 078) days after cutting the forage shoots that were underdifferent insect densities for 6 days

4 Discussion

The B ruziziensis chlorophyll content decreased dependingon theM spectabilis density and time the plant was exposedto the insect pest Similar results were observed in Niet al [25] wherein attacks by Blissus leucopterus signifi-cantly reduced the chlorophyll content and consequentlythe photosynthetic rate in certain forage millet genotypesunder field conditions those authors reported that such areduction decreased the leaf area and consequently reducedthe accumulated biomass SimilarlyWang et al [26] observedreductions in chlorophyll content due to an attack by her-bivores which may negatively affect plantsrsquo photosyntheticcapacities This observation is concerning for forage becausethe forage rest period is calculated according to its capacity ofregeneration in rotational grazing system where accordingto Martha Junior et al [27] the pasture is subjected toalternating grazing and rest periods According to the resultsherein 8 and 16M spectabilis adults that attack forage clumpfor 6 or 3 days respectively are sufficient to reduce foragechlorophyll content therefore one can infer that a longer restperiod for B ruziziensis is necessary for increasing spittlebuginfestation densities given the plantsrsquo reduced photosynthetic

4 The Scientific World Journal

rates which could jeopardize the fodder supply for animalsThis reduced supply should be even higher because pastureswith damage symptoms from spittlebugs are less palatableaccording to Valerio and Nakano [11]

The plant damage scores herein were lower than reportedby Cardona et al [18] for B ruziziensis (48) Cardona et al[28] for B decumbens (37) infested with 12 and 5 Aeneolamiavaria adults per cage (6 cm in diameter by 24 cm in height)and Resende et al [29] who observed damage scores higherthan 35 for B ruziziensis infested with different adult Mspectabilis densities per cage (040 times 040 times 080m) in agreenhouse Given that this study was conducted in thefield it can be inferred that this difference is related to theconditions of plant development such as the volume of soilavailable to the root system for plants grown in the fieldhigher soil volume may yield lower damage

Despite the observation that infestation densities of 4 812 and 16M spectabilis adults per plant caused intermediatedamage in the fodder (25 to 50 leaf area was damaged)according to Lopez et al [14] damage by adult spittlebugsis irreversible Thus the portion of the forage damaged bythe insect will have a compromised development capacitywhich will reduce the fodder supply for animals over timeand decreasemilk as well asmeat production To alleviate thisproblem the management tactic proposed by Painter [30]and Soares et al [31] is viable it uses harvest anticipationto change the timing between the presence of food andassociated insects which is a form of ecological resistancereferred to as ldquoescape from the hostrdquo Thus the B ruziziensispastures infested by adult M spectabilis may be grazed earlyin the infestation which would target the forage for animaluse before the plants show damage symptoms caused by aninsect attack which may ensure forage regrowth

The inverse relationship observed herein between theforage damage scores and chlorophyll content upon exposureto M spectabilis adults was similar to the results by Resendeet al [29] who observed a high negative correlation betweenthe damage score and chlorophyll content for B ruziziensisinfested withM spectabilis adults in a greenhouse and Lopezet al [14] who reported a high correlation between thedamage score and percentage of chlorophyll loss inBrachiariagenotypes infested with adult spittlebugs

The functional losses observed herein were lower thanthose reported by Resende et al [29] who found that attacksby 12M spectabilis adults generated a functional plant lossindex over 75 of B ruziziensis in a greenhouse Valueshigher than those herein were also demonstrated by Lopez etal [14] in B ruziziensis who recorded index of 934 and 100after attacks byA varia andZ carbonaria adults respectivelyThe functional plant loss indexmeasures the plantsrsquo toleranceto insects [20 21] and according to Lopez et al [14] it isthe best index for estimating Brachiaria tolerance to spittle-bugs

The differences from attacks by adult spittlebugs in theaforementioned trials are attributed to the different condi-tions of plant development The study herein is the onlystudy to assess the impact of adult M spectabilis under fieldconditions with no physical limitations on land area exploredby roots which ensures greater energy reserves in this region

of plants Thus plants tend to better tolerate insect pestattacks under field conditions than in the greenhouse

The increased B ruziziensis dry matter percentage dueto the enhancedM spectabilis infestation density herein wasalso demonstrated by Resende et al [29] in B ruziziensis 10days after exposure to 24M spectabilis adults in a greenhouseThose authors attributed the increased dry matter percent-age to the reduced fresh weight from insect damage Theincreased dry matter percentage was also reported for Bdecumbens infested withZulia entreriana adults [11] in alfalfared clover attacked by Philaenus spumarius [32 33] andDigitaria decumbens infested with Prosapia bicincta [34]

Despite the shoot damage from the M spectabilis attacksuch damage levels did not affect the B ruziziensis regrowthability under field conditions In contrast Resende et al [29]observed a significantly reduced number of B ruziziensisshoots infested with 12 or more M spectabilis adults in agreenhouse This observation is attributed to differences inthe fodder development conditions considering that hereinthe plants were grown directly in the groundwithout physicalconstraints to root system development Because differenceswere not observed in the fresh weight and forage regrowthafter the pest insect attack tactics for using such forage shouldbe adopted early during infestation before themore advancedsymptoms of attack by adults Among these tactics the forageby grazing animals or even cutting and storage of thismaterialin silos to be provided to cattle during the dry season arehighlighted According to Valerio [35] the forage root systemmay be reduced due to frequent spittlebug attacks whichresults in reduced forage persistence The plants used hereinwere grown under favorable conditions for extensive rootdevelopment and were not previously attacked by spittlebugsThese factors were important to maintain the number ofshoots after infestation these data emphasize that properpasture and soil fertility management guarantee forage re-establishment even after insect attacks

5 Conclusion

Thus attacks by 8 adult M spectabilis per clump of B ruz-iziensiswith an average of 80 tillers for six days were sufficientto reduce the chlorophyll content and cause functional plantloss index of 60 but it did not affect the fresh weight norregrowth capability which indicates that tactics for usingthis forage are recommended at the beginning of insectinfestation This population density can be a reference forspittlebug integrated management in Brachiaria

Acknowledgments

The authors thank the Conselho Nacional de Desenvolvi-mento Cientıfico e Tecnologico (CNPq Brazil) and FundacaodeAmparo a Pesquisa doEstado deMinasGerais (FAPEMIGBrazil) for supporting their research

References

[1] A J E Peron and A R Evangelista ldquoDegradacao de pastagensem regioes de cerradordquo Ciencia e Agrotecnologia vol 28 pp655ndash661 2004

The Scientific World Journal 5

[2] L Moreira and E D Assad Segmentacao e Classificacao super-visionada Para identificar Pastagens Degradadas Embrapa-CPAC Planaltina Goias 1997

[3] J R Valerio C Cardona D C Peck and G Sotelo SpittlebugsldquoBioecology host plant resistance and advances in IPMrdquo Pro-ceedings of the 19 International Grassland Congress pp 217ndash221 FEALQ Sao Pedro Piracicaba 2001

[4] S V Paula-Moraes Cadeia Produtiva da Carne E ameacadaPor Praga 2006 httpwwwcpacembrapabrmaterias pripagnucleotematicohtml

[5] AM Auad A D SimoesM V Leite S E B Silva D R Santosand P H Monteiro ldquoSeasonal dynamics of egg diapause inMahanarva spectabilis (Distant 1909) (Hemiptera Cercopidae)on elephant grassrdquo Arquivos do Instituto Biologico vol 78 pp325ndash330 2011

[6] D W Fewkes ldquoThe biology of sugar cane froghopperrdquo in Pestsof Sugar Cane J R Williams J R Metcalfe R W Mungomeryand R Mathes Eds pp 281ndash307 Elsevier Amsterdam TheNetherlands 1969

[7] A M Auad A D Simoes A V Pereira et al ldquoSelecao degenotipos de capim-elefante quanto a resistencia a cigarrinha-das-pastagensrdquo Pesquisa Agropecuaria Brasileira vol 42 pp1077ndash1081 2007

[8] V Thompson ldquoAssociative nitrogen fixation C4 photosynthe-sis and the evolution of spittlebugs (Hemiptera Cercopidae)as major pests of neotropical sugarcane and forage grassesrdquoBulletin of Entomological Research vol 94 no 3 pp 189ndash2002004

[9] R A Byers andH DWells ldquoPhytotoxemia of coastal bermuda-grass caused by the two-lined spittlebug Prosapia bicincta(Homoptera Cercopidae)rdquo Annals of the Entomological Societyof America vol 59 pp 1067ndash1071 1966

[10] Z A Ramiro R A Miranda and E A Batista Filho ldquoObser-vacoes sobre a fluctuacao de cigarrinhas (Homoptera Cercop-idae) em pastagem formale em Brachiaria decumbens mantidaem diferentes nıveis de desenvolvimento vegetativordquo Anais daSociedade Entomologica do Brasil vol 13 pp 357ndash370 1984

[11] J R Valerio and O Nakano ldquoDanos causados pelo adulto dacigarrinha-das-pastagens Zulia entreriana (Berg 1879) (Homo-ptera Cercopidae) em plantas de Brachiaria decumbens Stapfmantidas em diferentes nıveis de umidaderdquo Anais da SociedadeEntomologica do Brasil vol 16 pp 341ndash350 1988

[12] S L LapointeG Sotelo andG LArango ldquoImproved techniquefor rearing spittlebugs (Homoptera Cercopi-dae)rdquo Journal ofEconomic Entomology vol 82 pp 1764ndash1766 1989

[13] D C Peck ldquoNatural history of the spittlebug Prosapia bicincta(Homoptera Cercopidae) in association with dairy pastures ofCosta RicardquoAnnals of the Entomological Society of America vol91 no 4 pp 435ndash444 1998

[14] F Lopez C Cardona J W Miles G Sotelo and J Mon-toya ldquoScreening for resistance to adult spittlebugs (HemipteraCercopidae) in Brachiaria spp methods and categories ofresistancerdquo Journal of Economic Entomology vol 102 no 3 pp1309ndash1316 2009

[15] F Holmann and D C Peck ldquoEconomic damage caused by spit-tlebugs (Homoptera Cercopidae) in Colombia a first approxi-mation of impact on animal production in Brachiaria decum-bens pasturesrdquo Neotropical Entomology vol 31 no 2 pp 275ndash284 2002

[16] H F Kain and D S Atkison ldquoProblems of insect pest assess-ment in pasturesrdquo New Zealand Entomologist vol 6 no 1 pp9ndash13 1975

[17] J Diaz-Montano J C Reese W T Schapaugh and L R Camp-bell ldquoChlorophyll loss caused by soybean aphid (HemipteraAphididae) feeding on soybeanrdquo Journal of Economic Entomol-ogy vol 100 no 5 pp 1657ndash1662 2007

[18] C Cardona JWMiles andG Sotelo ldquoAn improvedmethodol-ogy formassive screening of Brachiaria spp genotypes for resis-tance toAeneolamia varia (Homoptera Cercopidae)rdquo Journal ofEconomic Entomology vol 92 no 2 pp 490ndash496 1999

[19] A Pabon C Cardona J W Miles and G Sotelo ldquoResponseof resistant and susceptible Brachiaria spp genotypes tosimultaneous infestation with multiple species of spittlebugs(Hemiptera Cercopidae)rdquo Journal of Economic Entomology vol100 no 6 pp 1896ndash1903 2007

[20] J Morgan G Wilde and D Johnson ldquoGreenbug resistance incommercial sorghum hybrids in the seedling stagerdquo Journal ofEconomic Entomology vol 73 pp 510ndash514 1980

[21] N Panda andEAHeinrichs ldquoLevels of tolerance and antibiosisin rice varieties having moderate resistance to the brown plan-thopper Nilaparvata lugens (Stal) (Hemiptera Delphacidae)rdquoEnvironmental Entomology vol 12 pp 1204ndash1214 1983

[22] C M Smith Z R Khan and E M D Pathak Techniques forEvaluating Insect Resistance in Crop Plants CRC Boca RatonFla USA 1994

[23] SISVAR Versao 53 (Biud 75) Sistemas de Analises de VarianciaPara Dados Balanceados Programa de Analises Estatısticase Planejamento de Experimentos Lavras MG UniversidadeFederal de Lavras 2010

[24] BIOESTAT Sistema Para Analises Estatısticas Versao 50 ParaUniversidade Federal do Para Belem Brazil 2007

[25] X Ni J P Wilson and G D Buntin ldquoDifferential responsesof forage pearl millet genotypes to chinch bug (HeteropteraBlissidae) feedingrdquo Journal of Economic Entomology vol 102no 5 pp 1960ndash1969 2009

[26] T Wang S S Quisenberry N I Xinzhi and V TolmayldquoAphid (Hemiptera Aphididae) resistance in wheat near-isogenic linesrdquo Journal of Economic Entomology vol 97 no 2pp 646ndash653 2004

[27] G BMartha Junior L G Barioni L Vilela and A O BarcellosldquoArea do piquete e taxa de lotacao no pastejo rotacionadordquoComunicado Tecnico 101 2003

[28] C Cardona J W Miles E Zuniga and G Sotelo ldquoIndepen-dence of resistance in Brachiaria spp to nymphs or to adultspittlebugs (Hemiptera Cercopidae) implications for breedingfor resistancerdquo Journal of Economic Entomology vol 103 no 5pp 1860ndash1865 2010

[29] T T Resende A M Auad M G Fonseca T H Santos andT M Vieira ldquoImpact of the spittlebug Mahanarva spectabilison signal grassrdquoTheScientificWorldJournal vol 2012 Article ID926715 6 pages 2012

[30] R H Painter Insect Resistance in Crop Plants MacMillan NewYork NY USA 1951

[31] J J Soares R S Melo C A Almeida and A M C FerreiraldquoProgresso no manejo integrado de pragas do algodoeiro noOeste Baianordquo Comunicado Tecnico 194 Embrapa AlgodaoCampina Grande Brazil 2003

[32] C R Weaver and J W Hibbs ldquoEffects of spittlebug infestationon nutrition value of alfalfa and red cloverrdquo Journal of EconomicEntomology vol 45 pp 626ndash628 1952

[33] R BMarthur andR L Pienkowski ldquoInfluence of adultmeadowspittlebug feeding on forage qualityrdquo Journal of EconomicEntomology vol 60 pp 207ndash209 1967

6 The Scientific World Journal

[34] E B Fagan Bionomics and control of the two-lined spittlebugProsapia bicincta on Florida pastures and notes on Prosapiaplagiata in Costa Rica (Homoptera Cercopidae) [Dissertation]University of Florida 1969

[35] J R Valerio ldquoConsideracoes sobre a morte de pastagensde Brachiaria brizantha cv Marandu em alguns Estados doCentro e Norte do Brasil enfoque entomologicordquo ComunicadoTecnico EmbrapaGado de Corte CampoGrande Brazil 2006

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AgricultureAdvances in

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

PsycheHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BiodiversityInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

ScientificaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Plant GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Biotechnology Research International

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Forestry ResearchInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Journal of BotanyHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

EcologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Veterinary Medicine International

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Cell BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Evolutionary BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

2 The Scientific World Journal

According to Kain and Atkison [16] the main problemassociated with studies that assess loss promoted by insectsin pastures is converting quantitative and qualitative loss ofgrassland into livestock production and consequently estab-lishing economic damage level In this context determiningeffects of adult M spectabilis attacks on B ruziziensis is animportant tool that will assist in an integratedmanagement ofthis insect pest Thus the aim of this study was to determinethe damage caused by adult M spectabilis on B ruziziensisunder field conditions

2 Materials and Methods

21 Plants and Insects The research was conducted at theEmbrapa Dairy Cattle research experimental field which islocated in Coronel Pacheco MG Brazil (21∘3310158402210158401015840 southand 43∘1610158401510158401015840 west) During the experiment the meantemperature was 278∘C (maximum 464∘C and minimum202∘C) with a 769 relative humidity (maximum 982and minimum 243) These parameters were recorded bya data logger every 2 minutes and transferred to software(Hoboware) these values were used to determine the averagefor the experimental period Twenty days before beginningthe experiment the B ruziziensis pasture was cut 15 cm aboveground level to standardize plant height Each experimentalplot included a forage clump with a 30 cm mean height8852 tillers and 3323 SPAD units (chlorophyll content) Mspectabilis adults were collected in a pasture distal to theexperimental area These insects were maintained in cagesuntil they were used in the experiment

22 Experiment The randomized block design included 5infestation densities and 8 repetitions Each plot compriseda B ruziziensis clump protected by a metal frame cage (70 times40 times 40 cm) covered with organza fabric The experimentalplots were installed equidistant (5 times 2m) and covered with400m2 meters of grassland Inside each cage were 0 4 8 12and 16M spectabilis adultsThedead insectswere replenisheddaily to maintain a constantM spectabilis density for 6 daysThereafter the insects were removed and evaluated for thefollowing parameters chlorophyll content damage score dryas well as fresh weights percentage of dry shoot matter andthe forage regrowth ability

The chlorophyll content was measured using the appara-tus Minolta SPAD 502 OL (Konica Minolta Sensing OsakaJapan) before infestation as well as 3 and 6 days after onsetof infestation in three leaf blades from one tiller of the plantAccording to Diaz-Montano et al [17] the chlorophyll meterSPAD-502 is an important device used to measure chloro-phyll loss by sucking insects The mean chlorophyll contentin the three leaf blades from each plant was calculated

After 6 days of infestation the damage percentage score inthe shoots from each plant was assigned by three evaluatorsthat followed the damage scale from 1 through 5 proposedby Cardona et al [18] Furthermore the damage scores wereclassified based on scores developed by Pabon et al [19]as follows scores 1 and 2mdashthe grass tolerates insect attackscores between 21 and 3mdashintermediate tolerance and scoresover 3mdashsusceptible to insect attack

After 6 days the plants with different infestation densitieswere cut at the soil level and their leaves and stems wereweighed for the fresh weight These materials were dried at55∘C for 72 hours and after this period they were weighed torecord the dry weight The percentages were then calculatedfor the dry matter and the functional plant loss index (FPLI)which were proposed by Morgan et al [20] and modifiedby Panda and Heinrichs [21] This index is calculated basedon the damage score (DS) dry weight of uninfested plants(DWUP) and dry weight of infested plants (DWIP) asfollows FPLI () = [1 minus (DWIPDWUP) times (1 minusDS5)] times 100and this method is considered a useful tool for quantifyingtolerance [22]

To evaluate the forage regrowth ability with adult spit-tlebugs the metal frame of each cage was maintained for 30days without organza fabric covering The number of shootsin each clump was evaluated 15 and 30 days after cutting

23 Statistical Analysis The average chlorophyll content inthe three leaf blades from each plant damage score freshand dry shoot weights dry matter percentage and forageregrowth ability were compared using analysis of varianceand when significant (119875 le 005) regression analyses wereperformed for the spittlebug infestation density The averagechlorophyll content was compared using the Tukey test toassess the effect of 3 and 6 days of forage exposure to adultspittlebugs (119875 le 005)

The analyses were performed using the program SISVAR51 [23] (Federal University of Lavras MG Brazil) ThePearson correlation between the chlorophyll content anddamage score was generated using BioEstat 50 [24] (FederalUniversity of Para Para Brazil)

3 Results

In the first evaluation which was conducted prior to forageinfestation by M spectabilis adults the chlorophyll contentwas the same (119865 = 0191 119875 = 09427) regardless of the futureinfestation level which demonstrates plant standardizationThis standardized plant behavior was reproduced in thesecond evaluation which was performed 3 days after theplants were exposed to insects (119865 = 213 119875 = 008)In the subsequent evaluation which was performed 6 daysafter infestation the forage chlorophyll content significantlydecreased due to an increase in theM spectabilis infestation(119865 = 728 119875 lt 00001) which indicates a quadratic regres-sion curve for this exposure time (Figure 1)

After the exposure the plant chlorophyll content did notchange in plants exposed to 4 adult spittlebugs (119865 = 109 119875 =033) and those with no contact to the insect pest (119865 = 225119875 = 010) In contrast there was a significant reduction inforage chlorophyll content after 6 days of exposure to 8 (119865 =443 119875 lt 001) and 12 insects (119865 = 289 119875 lt 005) as wellas for plants exposed for 3 days to 16 insects (119865 = 742 119875 lt0001) (Figure 2)

For the damage score higher M spectabilis infestationlevels yielded greater damage to plants exposed to spittlebugsfor 6 days (119865 = 1853 119875 lt 00001) (Figure 3) Notably underfield conditions B ruziziensis had an intermediate tolerance

The Scientific World Journal 3

20

24

28

32

36

40

0 4 8 12 16Adults of M spectabilis per plant

Chlo

roph

yll c

onte

nt y = 00652x2 minus 16903x + 36491

R2 = 09354

Figure 1 Relationship between infestation levels of adult Mspectabilis and chlorophyll content (SPAD unit) in B ruziziensis

05

1015

1216

2025303540

4

45

0

0

3 6

8

Time (days)

a a a a a a a ab abb

aa

b b b

Chlo

roph

yll c

onte

nt

Adults of M spectabilis per plant

Figure 2 Relationship between chlorophyll content (SPAD unit)of B ruziziensis and exposure time (0 3 and 6 days) at differentinfestation levels of adultM spectabilis Mean values followed by thesame letter within the levels of infestation did not differ by Tukeytest

to be attacked by 4 8 12 and 16M spectabilis adults for 6 daysbased on converting the damage score parameters into thetolerance established by Pabon et al[19] Further the damagescores inversely correlate with the forage chlorophyll contentupon a day exposure to M spectabilis adults (119903 = minus044119905 = minus3348 119875 = 00018)

For the functional plant loss index there was no signifi-cant difference in this ratio among infestation densities (119865 =1613 119875 = 0208) However only four and eight adult Mspectabilis per plant were sufficient to generate a functionalplant loss index greater than 453 and 60 respectivelyin B ruziziensis however for 16 adults such a loss reached619 Likewise increasing M spectabilis densities did notsignificantly alter the fresh (119865 = 206 119875 = 0113) and dryweights (119865 = 1416119875 = 02546) in plants exposed to the adultinsects for 6 days However the increased infestation densitycaused a significant increase in the dry matter percentage ofplant (119865 = 2798 119875 = 00451) (Figure 4)

The tiller number in plants was equal in the assessmentsperformed 15 (119865 = 0565 119875 = 069) and 30 (119865 = 0432

0

1

2

3

4

5

Dam

age s

core

y = minus00072x2 + 02187x + 11166

R2 = 09376

0 4 8 12 16Adults of M spectabilis per plant

Figure 3 Relationship between infestation levels of M spectabilisadults and damage scores for B ruziziensis over 6 days

20

21

22

23

24

25

26

R2 = 08974

0 4 8 12 16Adults of M spectabilis per plant

Dry

mas

s per

cent

age y = 00122x2 + 00351x + 21824

Figure 4 Relationship between infestation levels of M spectabilisadults and dry mass percentage of the B ruziziensis

119875 = 078) days after cutting the forage shoots that were underdifferent insect densities for 6 days

4 Discussion

The B ruziziensis chlorophyll content decreased dependingon theM spectabilis density and time the plant was exposedto the insect pest Similar results were observed in Niet al [25] wherein attacks by Blissus leucopterus signifi-cantly reduced the chlorophyll content and consequentlythe photosynthetic rate in certain forage millet genotypesunder field conditions those authors reported that such areduction decreased the leaf area and consequently reducedthe accumulated biomass SimilarlyWang et al [26] observedreductions in chlorophyll content due to an attack by her-bivores which may negatively affect plantsrsquo photosyntheticcapacities This observation is concerning for forage becausethe forage rest period is calculated according to its capacity ofregeneration in rotational grazing system where accordingto Martha Junior et al [27] the pasture is subjected toalternating grazing and rest periods According to the resultsherein 8 and 16M spectabilis adults that attack forage clumpfor 6 or 3 days respectively are sufficient to reduce foragechlorophyll content therefore one can infer that a longer restperiod for B ruziziensis is necessary for increasing spittlebuginfestation densities given the plantsrsquo reduced photosynthetic

4 The Scientific World Journal

rates which could jeopardize the fodder supply for animalsThis reduced supply should be even higher because pastureswith damage symptoms from spittlebugs are less palatableaccording to Valerio and Nakano [11]

The plant damage scores herein were lower than reportedby Cardona et al [18] for B ruziziensis (48) Cardona et al[28] for B decumbens (37) infested with 12 and 5 Aeneolamiavaria adults per cage (6 cm in diameter by 24 cm in height)and Resende et al [29] who observed damage scores higherthan 35 for B ruziziensis infested with different adult Mspectabilis densities per cage (040 times 040 times 080m) in agreenhouse Given that this study was conducted in thefield it can be inferred that this difference is related to theconditions of plant development such as the volume of soilavailable to the root system for plants grown in the fieldhigher soil volume may yield lower damage

Despite the observation that infestation densities of 4 812 and 16M spectabilis adults per plant caused intermediatedamage in the fodder (25 to 50 leaf area was damaged)according to Lopez et al [14] damage by adult spittlebugsis irreversible Thus the portion of the forage damaged bythe insect will have a compromised development capacitywhich will reduce the fodder supply for animals over timeand decreasemilk as well asmeat production To alleviate thisproblem the management tactic proposed by Painter [30]and Soares et al [31] is viable it uses harvest anticipationto change the timing between the presence of food andassociated insects which is a form of ecological resistancereferred to as ldquoescape from the hostrdquo Thus the B ruziziensispastures infested by adult M spectabilis may be grazed earlyin the infestation which would target the forage for animaluse before the plants show damage symptoms caused by aninsect attack which may ensure forage regrowth

The inverse relationship observed herein between theforage damage scores and chlorophyll content upon exposureto M spectabilis adults was similar to the results by Resendeet al [29] who observed a high negative correlation betweenthe damage score and chlorophyll content for B ruziziensisinfested withM spectabilis adults in a greenhouse and Lopezet al [14] who reported a high correlation between thedamage score and percentage of chlorophyll loss inBrachiariagenotypes infested with adult spittlebugs

The functional losses observed herein were lower thanthose reported by Resende et al [29] who found that attacksby 12M spectabilis adults generated a functional plant lossindex over 75 of B ruziziensis in a greenhouse Valueshigher than those herein were also demonstrated by Lopez etal [14] in B ruziziensis who recorded index of 934 and 100after attacks byA varia andZ carbonaria adults respectivelyThe functional plant loss indexmeasures the plantsrsquo toleranceto insects [20 21] and according to Lopez et al [14] it isthe best index for estimating Brachiaria tolerance to spittle-bugs

The differences from attacks by adult spittlebugs in theaforementioned trials are attributed to the different condi-tions of plant development The study herein is the onlystudy to assess the impact of adult M spectabilis under fieldconditions with no physical limitations on land area exploredby roots which ensures greater energy reserves in this region

of plants Thus plants tend to better tolerate insect pestattacks under field conditions than in the greenhouse

The increased B ruziziensis dry matter percentage dueto the enhancedM spectabilis infestation density herein wasalso demonstrated by Resende et al [29] in B ruziziensis 10days after exposure to 24M spectabilis adults in a greenhouseThose authors attributed the increased dry matter percent-age to the reduced fresh weight from insect damage Theincreased dry matter percentage was also reported for Bdecumbens infested withZulia entreriana adults [11] in alfalfared clover attacked by Philaenus spumarius [32 33] andDigitaria decumbens infested with Prosapia bicincta [34]

Despite the shoot damage from the M spectabilis attacksuch damage levels did not affect the B ruziziensis regrowthability under field conditions In contrast Resende et al [29]observed a significantly reduced number of B ruziziensisshoots infested with 12 or more M spectabilis adults in agreenhouse This observation is attributed to differences inthe fodder development conditions considering that hereinthe plants were grown directly in the groundwithout physicalconstraints to root system development Because differenceswere not observed in the fresh weight and forage regrowthafter the pest insect attack tactics for using such forage shouldbe adopted early during infestation before themore advancedsymptoms of attack by adults Among these tactics the forageby grazing animals or even cutting and storage of thismaterialin silos to be provided to cattle during the dry season arehighlighted According to Valerio [35] the forage root systemmay be reduced due to frequent spittlebug attacks whichresults in reduced forage persistence The plants used hereinwere grown under favorable conditions for extensive rootdevelopment and were not previously attacked by spittlebugsThese factors were important to maintain the number ofshoots after infestation these data emphasize that properpasture and soil fertility management guarantee forage re-establishment even after insect attacks

5 Conclusion

Thus attacks by 8 adult M spectabilis per clump of B ruz-iziensiswith an average of 80 tillers for six days were sufficientto reduce the chlorophyll content and cause functional plantloss index of 60 but it did not affect the fresh weight norregrowth capability which indicates that tactics for usingthis forage are recommended at the beginning of insectinfestation This population density can be a reference forspittlebug integrated management in Brachiaria

Acknowledgments

The authors thank the Conselho Nacional de Desenvolvi-mento Cientıfico e Tecnologico (CNPq Brazil) and FundacaodeAmparo a Pesquisa doEstado deMinasGerais (FAPEMIGBrazil) for supporting their research

References

[1] A J E Peron and A R Evangelista ldquoDegradacao de pastagensem regioes de cerradordquo Ciencia e Agrotecnologia vol 28 pp655ndash661 2004

The Scientific World Journal 5

[2] L Moreira and E D Assad Segmentacao e Classificacao super-visionada Para identificar Pastagens Degradadas Embrapa-CPAC Planaltina Goias 1997

[3] J R Valerio C Cardona D C Peck and G Sotelo SpittlebugsldquoBioecology host plant resistance and advances in IPMrdquo Pro-ceedings of the 19 International Grassland Congress pp 217ndash221 FEALQ Sao Pedro Piracicaba 2001

[4] S V Paula-Moraes Cadeia Produtiva da Carne E ameacadaPor Praga 2006 httpwwwcpacembrapabrmaterias pripagnucleotematicohtml

[5] AM Auad A D SimoesM V Leite S E B Silva D R Santosand P H Monteiro ldquoSeasonal dynamics of egg diapause inMahanarva spectabilis (Distant 1909) (Hemiptera Cercopidae)on elephant grassrdquo Arquivos do Instituto Biologico vol 78 pp325ndash330 2011

[6] D W Fewkes ldquoThe biology of sugar cane froghopperrdquo in Pestsof Sugar Cane J R Williams J R Metcalfe R W Mungomeryand R Mathes Eds pp 281ndash307 Elsevier Amsterdam TheNetherlands 1969

[7] A M Auad A D Simoes A V Pereira et al ldquoSelecao degenotipos de capim-elefante quanto a resistencia a cigarrinha-das-pastagensrdquo Pesquisa Agropecuaria Brasileira vol 42 pp1077ndash1081 2007

[8] V Thompson ldquoAssociative nitrogen fixation C4 photosynthe-sis and the evolution of spittlebugs (Hemiptera Cercopidae)as major pests of neotropical sugarcane and forage grassesrdquoBulletin of Entomological Research vol 94 no 3 pp 189ndash2002004

[9] R A Byers andH DWells ldquoPhytotoxemia of coastal bermuda-grass caused by the two-lined spittlebug Prosapia bicincta(Homoptera Cercopidae)rdquo Annals of the Entomological Societyof America vol 59 pp 1067ndash1071 1966

[10] Z A Ramiro R A Miranda and E A Batista Filho ldquoObser-vacoes sobre a fluctuacao de cigarrinhas (Homoptera Cercop-idae) em pastagem formale em Brachiaria decumbens mantidaem diferentes nıveis de desenvolvimento vegetativordquo Anais daSociedade Entomologica do Brasil vol 13 pp 357ndash370 1984

[11] J R Valerio and O Nakano ldquoDanos causados pelo adulto dacigarrinha-das-pastagens Zulia entreriana (Berg 1879) (Homo-ptera Cercopidae) em plantas de Brachiaria decumbens Stapfmantidas em diferentes nıveis de umidaderdquo Anais da SociedadeEntomologica do Brasil vol 16 pp 341ndash350 1988

[12] S L LapointeG Sotelo andG LArango ldquoImproved techniquefor rearing spittlebugs (Homoptera Cercopi-dae)rdquo Journal ofEconomic Entomology vol 82 pp 1764ndash1766 1989

[13] D C Peck ldquoNatural history of the spittlebug Prosapia bicincta(Homoptera Cercopidae) in association with dairy pastures ofCosta RicardquoAnnals of the Entomological Society of America vol91 no 4 pp 435ndash444 1998

[14] F Lopez C Cardona J W Miles G Sotelo and J Mon-toya ldquoScreening for resistance to adult spittlebugs (HemipteraCercopidae) in Brachiaria spp methods and categories ofresistancerdquo Journal of Economic Entomology vol 102 no 3 pp1309ndash1316 2009

[15] F Holmann and D C Peck ldquoEconomic damage caused by spit-tlebugs (Homoptera Cercopidae) in Colombia a first approxi-mation of impact on animal production in Brachiaria decum-bens pasturesrdquo Neotropical Entomology vol 31 no 2 pp 275ndash284 2002

[16] H F Kain and D S Atkison ldquoProblems of insect pest assess-ment in pasturesrdquo New Zealand Entomologist vol 6 no 1 pp9ndash13 1975

[17] J Diaz-Montano J C Reese W T Schapaugh and L R Camp-bell ldquoChlorophyll loss caused by soybean aphid (HemipteraAphididae) feeding on soybeanrdquo Journal of Economic Entomol-ogy vol 100 no 5 pp 1657ndash1662 2007

[18] C Cardona JWMiles andG Sotelo ldquoAn improvedmethodol-ogy formassive screening of Brachiaria spp genotypes for resis-tance toAeneolamia varia (Homoptera Cercopidae)rdquo Journal ofEconomic Entomology vol 92 no 2 pp 490ndash496 1999

[19] A Pabon C Cardona J W Miles and G Sotelo ldquoResponseof resistant and susceptible Brachiaria spp genotypes tosimultaneous infestation with multiple species of spittlebugs(Hemiptera Cercopidae)rdquo Journal of Economic Entomology vol100 no 6 pp 1896ndash1903 2007

[20] J Morgan G Wilde and D Johnson ldquoGreenbug resistance incommercial sorghum hybrids in the seedling stagerdquo Journal ofEconomic Entomology vol 73 pp 510ndash514 1980

[21] N Panda andEAHeinrichs ldquoLevels of tolerance and antibiosisin rice varieties having moderate resistance to the brown plan-thopper Nilaparvata lugens (Stal) (Hemiptera Delphacidae)rdquoEnvironmental Entomology vol 12 pp 1204ndash1214 1983

[22] C M Smith Z R Khan and E M D Pathak Techniques forEvaluating Insect Resistance in Crop Plants CRC Boca RatonFla USA 1994

[23] SISVAR Versao 53 (Biud 75) Sistemas de Analises de VarianciaPara Dados Balanceados Programa de Analises Estatısticase Planejamento de Experimentos Lavras MG UniversidadeFederal de Lavras 2010

[24] BIOESTAT Sistema Para Analises Estatısticas Versao 50 ParaUniversidade Federal do Para Belem Brazil 2007

[25] X Ni J P Wilson and G D Buntin ldquoDifferential responsesof forage pearl millet genotypes to chinch bug (HeteropteraBlissidae) feedingrdquo Journal of Economic Entomology vol 102no 5 pp 1960ndash1969 2009

[26] T Wang S S Quisenberry N I Xinzhi and V TolmayldquoAphid (Hemiptera Aphididae) resistance in wheat near-isogenic linesrdquo Journal of Economic Entomology vol 97 no 2pp 646ndash653 2004

[27] G BMartha Junior L G Barioni L Vilela and A O BarcellosldquoArea do piquete e taxa de lotacao no pastejo rotacionadordquoComunicado Tecnico 101 2003

[28] C Cardona J W Miles E Zuniga and G Sotelo ldquoIndepen-dence of resistance in Brachiaria spp to nymphs or to adultspittlebugs (Hemiptera Cercopidae) implications for breedingfor resistancerdquo Journal of Economic Entomology vol 103 no 5pp 1860ndash1865 2010

[29] T T Resende A M Auad M G Fonseca T H Santos andT M Vieira ldquoImpact of the spittlebug Mahanarva spectabilison signal grassrdquoTheScientificWorldJournal vol 2012 Article ID926715 6 pages 2012

[30] R H Painter Insect Resistance in Crop Plants MacMillan NewYork NY USA 1951

[31] J J Soares R S Melo C A Almeida and A M C FerreiraldquoProgresso no manejo integrado de pragas do algodoeiro noOeste Baianordquo Comunicado Tecnico 194 Embrapa AlgodaoCampina Grande Brazil 2003

[32] C R Weaver and J W Hibbs ldquoEffects of spittlebug infestationon nutrition value of alfalfa and red cloverrdquo Journal of EconomicEntomology vol 45 pp 626ndash628 1952

[33] R BMarthur andR L Pienkowski ldquoInfluence of adultmeadowspittlebug feeding on forage qualityrdquo Journal of EconomicEntomology vol 60 pp 207ndash209 1967

6 The Scientific World Journal

[34] E B Fagan Bionomics and control of the two-lined spittlebugProsapia bicincta on Florida pastures and notes on Prosapiaplagiata in Costa Rica (Homoptera Cercopidae) [Dissertation]University of Florida 1969

[35] J R Valerio ldquoConsideracoes sobre a morte de pastagensde Brachiaria brizantha cv Marandu em alguns Estados doCentro e Norte do Brasil enfoque entomologicordquo ComunicadoTecnico EmbrapaGado de Corte CampoGrande Brazil 2006

Submit your manuscripts athttpwwwhindawicom

Nutrition and Metabolism

Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Food ScienceInternational Journal of

Agronomy

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Microbiology

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom

Applied ampEnvironmentalSoil Science

Volume 2014

AgricultureAdvances in

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

PsycheHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BiodiversityInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

ScientificaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Plant GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Biotechnology Research International

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Forestry ResearchInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Journal of BotanyHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

EcologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Veterinary Medicine International

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Cell BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Evolutionary BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World Journal 3

20

24

28

32

36

40

0 4 8 12 16Adults of M spectabilis per plant

Chlo

roph

yll c

onte

nt y = 00652x2 minus 16903x + 36491

R2 = 09354

Figure 1 Relationship between infestation levels of adult Mspectabilis and chlorophyll content (SPAD unit) in B ruziziensis

05

1015

1216

2025303540

4

45

0

0

3 6

8

Time (days)

a a a a a a a ab abb

aa

b b b

Chlo

roph

yll c

onte

nt

Adults of M spectabilis per plant

Figure 2 Relationship between chlorophyll content (SPAD unit)of B ruziziensis and exposure time (0 3 and 6 days) at differentinfestation levels of adultM spectabilis Mean values followed by thesame letter within the levels of infestation did not differ by Tukeytest

to be attacked by 4 8 12 and 16M spectabilis adults for 6 daysbased on converting the damage score parameters into thetolerance established by Pabon et al[19] Further the damagescores inversely correlate with the forage chlorophyll contentupon a day exposure to M spectabilis adults (119903 = minus044119905 = minus3348 119875 = 00018)

For the functional plant loss index there was no signifi-cant difference in this ratio among infestation densities (119865 =1613 119875 = 0208) However only four and eight adult Mspectabilis per plant were sufficient to generate a functionalplant loss index greater than 453 and 60 respectivelyin B ruziziensis however for 16 adults such a loss reached619 Likewise increasing M spectabilis densities did notsignificantly alter the fresh (119865 = 206 119875 = 0113) and dryweights (119865 = 1416119875 = 02546) in plants exposed to the adultinsects for 6 days However the increased infestation densitycaused a significant increase in the dry matter percentage ofplant (119865 = 2798 119875 = 00451) (Figure 4)

The tiller number in plants was equal in the assessmentsperformed 15 (119865 = 0565 119875 = 069) and 30 (119865 = 0432

0

1

2

3

4

5

Dam

age s

core

y = minus00072x2 + 02187x + 11166

R2 = 09376

0 4 8 12 16Adults of M spectabilis per plant

Figure 3 Relationship between infestation levels of M spectabilisadults and damage scores for B ruziziensis over 6 days

20

21

22

23

24

25

26

R2 = 08974

0 4 8 12 16Adults of M spectabilis per plant

Dry

mas

s per

cent

age y = 00122x2 + 00351x + 21824

Figure 4 Relationship between infestation levels of M spectabilisadults and dry mass percentage of the B ruziziensis

119875 = 078) days after cutting the forage shoots that were underdifferent insect densities for 6 days

4 Discussion

The B ruziziensis chlorophyll content decreased dependingon theM spectabilis density and time the plant was exposedto the insect pest Similar results were observed in Niet al [25] wherein attacks by Blissus leucopterus signifi-cantly reduced the chlorophyll content and consequentlythe photosynthetic rate in certain forage millet genotypesunder field conditions those authors reported that such areduction decreased the leaf area and consequently reducedthe accumulated biomass SimilarlyWang et al [26] observedreductions in chlorophyll content due to an attack by her-bivores which may negatively affect plantsrsquo photosyntheticcapacities This observation is concerning for forage becausethe forage rest period is calculated according to its capacity ofregeneration in rotational grazing system where accordingto Martha Junior et al [27] the pasture is subjected toalternating grazing and rest periods According to the resultsherein 8 and 16M spectabilis adults that attack forage clumpfor 6 or 3 days respectively are sufficient to reduce foragechlorophyll content therefore one can infer that a longer restperiod for B ruziziensis is necessary for increasing spittlebuginfestation densities given the plantsrsquo reduced photosynthetic

4 The Scientific World Journal

rates which could jeopardize the fodder supply for animalsThis reduced supply should be even higher because pastureswith damage symptoms from spittlebugs are less palatableaccording to Valerio and Nakano [11]

The plant damage scores herein were lower than reportedby Cardona et al [18] for B ruziziensis (48) Cardona et al[28] for B decumbens (37) infested with 12 and 5 Aeneolamiavaria adults per cage (6 cm in diameter by 24 cm in height)and Resende et al [29] who observed damage scores higherthan 35 for B ruziziensis infested with different adult Mspectabilis densities per cage (040 times 040 times 080m) in agreenhouse Given that this study was conducted in thefield it can be inferred that this difference is related to theconditions of plant development such as the volume of soilavailable to the root system for plants grown in the fieldhigher soil volume may yield lower damage

Despite the observation that infestation densities of 4 812 and 16M spectabilis adults per plant caused intermediatedamage in the fodder (25 to 50 leaf area was damaged)according to Lopez et al [14] damage by adult spittlebugsis irreversible Thus the portion of the forage damaged bythe insect will have a compromised development capacitywhich will reduce the fodder supply for animals over timeand decreasemilk as well asmeat production To alleviate thisproblem the management tactic proposed by Painter [30]and Soares et al [31] is viable it uses harvest anticipationto change the timing between the presence of food andassociated insects which is a form of ecological resistancereferred to as ldquoescape from the hostrdquo Thus the B ruziziensispastures infested by adult M spectabilis may be grazed earlyin the infestation which would target the forage for animaluse before the plants show damage symptoms caused by aninsect attack which may ensure forage regrowth

The inverse relationship observed herein between theforage damage scores and chlorophyll content upon exposureto M spectabilis adults was similar to the results by Resendeet al [29] who observed a high negative correlation betweenthe damage score and chlorophyll content for B ruziziensisinfested withM spectabilis adults in a greenhouse and Lopezet al [14] who reported a high correlation between thedamage score and percentage of chlorophyll loss inBrachiariagenotypes infested with adult spittlebugs

The functional losses observed herein were lower thanthose reported by Resende et al [29] who found that attacksby 12M spectabilis adults generated a functional plant lossindex over 75 of B ruziziensis in a greenhouse Valueshigher than those herein were also demonstrated by Lopez etal [14] in B ruziziensis who recorded index of 934 and 100after attacks byA varia andZ carbonaria adults respectivelyThe functional plant loss indexmeasures the plantsrsquo toleranceto insects [20 21] and according to Lopez et al [14] it isthe best index for estimating Brachiaria tolerance to spittle-bugs

The differences from attacks by adult spittlebugs in theaforementioned trials are attributed to the different condi-tions of plant development The study herein is the onlystudy to assess the impact of adult M spectabilis under fieldconditions with no physical limitations on land area exploredby roots which ensures greater energy reserves in this region

of plants Thus plants tend to better tolerate insect pestattacks under field conditions than in the greenhouse

The increased B ruziziensis dry matter percentage dueto the enhancedM spectabilis infestation density herein wasalso demonstrated by Resende et al [29] in B ruziziensis 10days after exposure to 24M spectabilis adults in a greenhouseThose authors attributed the increased dry matter percent-age to the reduced fresh weight from insect damage Theincreased dry matter percentage was also reported for Bdecumbens infested withZulia entreriana adults [11] in alfalfared clover attacked by Philaenus spumarius [32 33] andDigitaria decumbens infested with Prosapia bicincta [34]

Despite the shoot damage from the M spectabilis attacksuch damage levels did not affect the B ruziziensis regrowthability under field conditions In contrast Resende et al [29]observed a significantly reduced number of B ruziziensisshoots infested with 12 or more M spectabilis adults in agreenhouse This observation is attributed to differences inthe fodder development conditions considering that hereinthe plants were grown directly in the groundwithout physicalconstraints to root system development Because differenceswere not observed in the fresh weight and forage regrowthafter the pest insect attack tactics for using such forage shouldbe adopted early during infestation before themore advancedsymptoms of attack by adults Among these tactics the forageby grazing animals or even cutting and storage of thismaterialin silos to be provided to cattle during the dry season arehighlighted According to Valerio [35] the forage root systemmay be reduced due to frequent spittlebug attacks whichresults in reduced forage persistence The plants used hereinwere grown under favorable conditions for extensive rootdevelopment and were not previously attacked by spittlebugsThese factors were important to maintain the number ofshoots after infestation these data emphasize that properpasture and soil fertility management guarantee forage re-establishment even after insect attacks

5 Conclusion

Thus attacks by 8 adult M spectabilis per clump of B ruz-iziensiswith an average of 80 tillers for six days were sufficientto reduce the chlorophyll content and cause functional plantloss index of 60 but it did not affect the fresh weight norregrowth capability which indicates that tactics for usingthis forage are recommended at the beginning of insectinfestation This population density can be a reference forspittlebug integrated management in Brachiaria

Acknowledgments

The authors thank the Conselho Nacional de Desenvolvi-mento Cientıfico e Tecnologico (CNPq Brazil) and FundacaodeAmparo a Pesquisa doEstado deMinasGerais (FAPEMIGBrazil) for supporting their research

References

[1] A J E Peron and A R Evangelista ldquoDegradacao de pastagensem regioes de cerradordquo Ciencia e Agrotecnologia vol 28 pp655ndash661 2004

The Scientific World Journal 5

[2] L Moreira and E D Assad Segmentacao e Classificacao super-visionada Para identificar Pastagens Degradadas Embrapa-CPAC Planaltina Goias 1997

[3] J R Valerio C Cardona D C Peck and G Sotelo SpittlebugsldquoBioecology host plant resistance and advances in IPMrdquo Pro-ceedings of the 19 International Grassland Congress pp 217ndash221 FEALQ Sao Pedro Piracicaba 2001

[4] S V Paula-Moraes Cadeia Produtiva da Carne E ameacadaPor Praga 2006 httpwwwcpacembrapabrmaterias pripagnucleotematicohtml

[5] AM Auad A D SimoesM V Leite S E B Silva D R Santosand P H Monteiro ldquoSeasonal dynamics of egg diapause inMahanarva spectabilis (Distant 1909) (Hemiptera Cercopidae)on elephant grassrdquo Arquivos do Instituto Biologico vol 78 pp325ndash330 2011

[6] D W Fewkes ldquoThe biology of sugar cane froghopperrdquo in Pestsof Sugar Cane J R Williams J R Metcalfe R W Mungomeryand R Mathes Eds pp 281ndash307 Elsevier Amsterdam TheNetherlands 1969

[7] A M Auad A D Simoes A V Pereira et al ldquoSelecao degenotipos de capim-elefante quanto a resistencia a cigarrinha-das-pastagensrdquo Pesquisa Agropecuaria Brasileira vol 42 pp1077ndash1081 2007

[8] V Thompson ldquoAssociative nitrogen fixation C4 photosynthe-sis and the evolution of spittlebugs (Hemiptera Cercopidae)as major pests of neotropical sugarcane and forage grassesrdquoBulletin of Entomological Research vol 94 no 3 pp 189ndash2002004

[9] R A Byers andH DWells ldquoPhytotoxemia of coastal bermuda-grass caused by the two-lined spittlebug Prosapia bicincta(Homoptera Cercopidae)rdquo Annals of the Entomological Societyof America vol 59 pp 1067ndash1071 1966

[10] Z A Ramiro R A Miranda and E A Batista Filho ldquoObser-vacoes sobre a fluctuacao de cigarrinhas (Homoptera Cercop-idae) em pastagem formale em Brachiaria decumbens mantidaem diferentes nıveis de desenvolvimento vegetativordquo Anais daSociedade Entomologica do Brasil vol 13 pp 357ndash370 1984

[11] J R Valerio and O Nakano ldquoDanos causados pelo adulto dacigarrinha-das-pastagens Zulia entreriana (Berg 1879) (Homo-ptera Cercopidae) em plantas de Brachiaria decumbens Stapfmantidas em diferentes nıveis de umidaderdquo Anais da SociedadeEntomologica do Brasil vol 16 pp 341ndash350 1988

[12] S L LapointeG Sotelo andG LArango ldquoImproved techniquefor rearing spittlebugs (Homoptera Cercopi-dae)rdquo Journal ofEconomic Entomology vol 82 pp 1764ndash1766 1989

[13] D C Peck ldquoNatural history of the spittlebug Prosapia bicincta(Homoptera Cercopidae) in association with dairy pastures ofCosta RicardquoAnnals of the Entomological Society of America vol91 no 4 pp 435ndash444 1998

[14] F Lopez C Cardona J W Miles G Sotelo and J Mon-toya ldquoScreening for resistance to adult spittlebugs (HemipteraCercopidae) in Brachiaria spp methods and categories ofresistancerdquo Journal of Economic Entomology vol 102 no 3 pp1309ndash1316 2009

[15] F Holmann and D C Peck ldquoEconomic damage caused by spit-tlebugs (Homoptera Cercopidae) in Colombia a first approxi-mation of impact on animal production in Brachiaria decum-bens pasturesrdquo Neotropical Entomology vol 31 no 2 pp 275ndash284 2002

[16] H F Kain and D S Atkison ldquoProblems of insect pest assess-ment in pasturesrdquo New Zealand Entomologist vol 6 no 1 pp9ndash13 1975

[17] J Diaz-Montano J C Reese W T Schapaugh and L R Camp-bell ldquoChlorophyll loss caused by soybean aphid (HemipteraAphididae) feeding on soybeanrdquo Journal of Economic Entomol-ogy vol 100 no 5 pp 1657ndash1662 2007

[18] C Cardona JWMiles andG Sotelo ldquoAn improvedmethodol-ogy formassive screening of Brachiaria spp genotypes for resis-tance toAeneolamia varia (Homoptera Cercopidae)rdquo Journal ofEconomic Entomology vol 92 no 2 pp 490ndash496 1999

[19] A Pabon C Cardona J W Miles and G Sotelo ldquoResponseof resistant and susceptible Brachiaria spp genotypes tosimultaneous infestation with multiple species of spittlebugs(Hemiptera Cercopidae)rdquo Journal of Economic Entomology vol100 no 6 pp 1896ndash1903 2007

[20] J Morgan G Wilde and D Johnson ldquoGreenbug resistance incommercial sorghum hybrids in the seedling stagerdquo Journal ofEconomic Entomology vol 73 pp 510ndash514 1980

[21] N Panda andEAHeinrichs ldquoLevels of tolerance and antibiosisin rice varieties having moderate resistance to the brown plan-thopper Nilaparvata lugens (Stal) (Hemiptera Delphacidae)rdquoEnvironmental Entomology vol 12 pp 1204ndash1214 1983

[22] C M Smith Z R Khan and E M D Pathak Techniques forEvaluating Insect Resistance in Crop Plants CRC Boca RatonFla USA 1994

[23] SISVAR Versao 53 (Biud 75) Sistemas de Analises de VarianciaPara Dados Balanceados Programa de Analises Estatısticase Planejamento de Experimentos Lavras MG UniversidadeFederal de Lavras 2010

[24] BIOESTAT Sistema Para Analises Estatısticas Versao 50 ParaUniversidade Federal do Para Belem Brazil 2007

[25] X Ni J P Wilson and G D Buntin ldquoDifferential responsesof forage pearl millet genotypes to chinch bug (HeteropteraBlissidae) feedingrdquo Journal of Economic Entomology vol 102no 5 pp 1960ndash1969 2009

[26] T Wang S S Quisenberry N I Xinzhi and V TolmayldquoAphid (Hemiptera Aphididae) resistance in wheat near-isogenic linesrdquo Journal of Economic Entomology vol 97 no 2pp 646ndash653 2004

[27] G BMartha Junior L G Barioni L Vilela and A O BarcellosldquoArea do piquete e taxa de lotacao no pastejo rotacionadordquoComunicado Tecnico 101 2003

[28] C Cardona J W Miles E Zuniga and G Sotelo ldquoIndepen-dence of resistance in Brachiaria spp to nymphs or to adultspittlebugs (Hemiptera Cercopidae) implications for breedingfor resistancerdquo Journal of Economic Entomology vol 103 no 5pp 1860ndash1865 2010

[29] T T Resende A M Auad M G Fonseca T H Santos andT M Vieira ldquoImpact of the spittlebug Mahanarva spectabilison signal grassrdquoTheScientificWorldJournal vol 2012 Article ID926715 6 pages 2012

[30] R H Painter Insect Resistance in Crop Plants MacMillan NewYork NY USA 1951

[31] J J Soares R S Melo C A Almeida and A M C FerreiraldquoProgresso no manejo integrado de pragas do algodoeiro noOeste Baianordquo Comunicado Tecnico 194 Embrapa AlgodaoCampina Grande Brazil 2003

[32] C R Weaver and J W Hibbs ldquoEffects of spittlebug infestationon nutrition value of alfalfa and red cloverrdquo Journal of EconomicEntomology vol 45 pp 626ndash628 1952

[33] R BMarthur andR L Pienkowski ldquoInfluence of adultmeadowspittlebug feeding on forage qualityrdquo Journal of EconomicEntomology vol 60 pp 207ndash209 1967

6 The Scientific World Journal

[34] E B Fagan Bionomics and control of the two-lined spittlebugProsapia bicincta on Florida pastures and notes on Prosapiaplagiata in Costa Rica (Homoptera Cercopidae) [Dissertation]University of Florida 1969

[35] J R Valerio ldquoConsideracoes sobre a morte de pastagensde Brachiaria brizantha cv Marandu em alguns Estados doCentro e Norte do Brasil enfoque entomologicordquo ComunicadoTecnico EmbrapaGado de Corte CampoGrande Brazil 2006

Submit your manuscripts athttpwwwhindawicom

Nutrition and Metabolism

Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Food ScienceInternational Journal of

Agronomy

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Microbiology

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom

Applied ampEnvironmentalSoil Science

Volume 2014

AgricultureAdvances in

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

PsycheHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BiodiversityInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

ScientificaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Plant GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Biotechnology Research International

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Forestry ResearchInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Journal of BotanyHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

EcologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Veterinary Medicine International

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Cell BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Evolutionary BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

4 The Scientific World Journal

rates which could jeopardize the fodder supply for animalsThis reduced supply should be even higher because pastureswith damage symptoms from spittlebugs are less palatableaccording to Valerio and Nakano [11]

The plant damage scores herein were lower than reportedby Cardona et al [18] for B ruziziensis (48) Cardona et al[28] for B decumbens (37) infested with 12 and 5 Aeneolamiavaria adults per cage (6 cm in diameter by 24 cm in height)and Resende et al [29] who observed damage scores higherthan 35 for B ruziziensis infested with different adult Mspectabilis densities per cage (040 times 040 times 080m) in agreenhouse Given that this study was conducted in thefield it can be inferred that this difference is related to theconditions of plant development such as the volume of soilavailable to the root system for plants grown in the fieldhigher soil volume may yield lower damage

Despite the observation that infestation densities of 4 812 and 16M spectabilis adults per plant caused intermediatedamage in the fodder (25 to 50 leaf area was damaged)according to Lopez et al [14] damage by adult spittlebugsis irreversible Thus the portion of the forage damaged bythe insect will have a compromised development capacitywhich will reduce the fodder supply for animals over timeand decreasemilk as well asmeat production To alleviate thisproblem the management tactic proposed by Painter [30]and Soares et al [31] is viable it uses harvest anticipationto change the timing between the presence of food andassociated insects which is a form of ecological resistancereferred to as ldquoescape from the hostrdquo Thus the B ruziziensispastures infested by adult M spectabilis may be grazed earlyin the infestation which would target the forage for animaluse before the plants show damage symptoms caused by aninsect attack which may ensure forage regrowth

The inverse relationship observed herein between theforage damage scores and chlorophyll content upon exposureto M spectabilis adults was similar to the results by Resendeet al [29] who observed a high negative correlation betweenthe damage score and chlorophyll content for B ruziziensisinfested withM spectabilis adults in a greenhouse and Lopezet al [14] who reported a high correlation between thedamage score and percentage of chlorophyll loss inBrachiariagenotypes infested with adult spittlebugs

The functional losses observed herein were lower thanthose reported by Resende et al [29] who found that attacksby 12M spectabilis adults generated a functional plant lossindex over 75 of B ruziziensis in a greenhouse Valueshigher than those herein were also demonstrated by Lopez etal [14] in B ruziziensis who recorded index of 934 and 100after attacks byA varia andZ carbonaria adults respectivelyThe functional plant loss indexmeasures the plantsrsquo toleranceto insects [20 21] and according to Lopez et al [14] it isthe best index for estimating Brachiaria tolerance to spittle-bugs

The differences from attacks by adult spittlebugs in theaforementioned trials are attributed to the different condi-tions of plant development The study herein is the onlystudy to assess the impact of adult M spectabilis under fieldconditions with no physical limitations on land area exploredby roots which ensures greater energy reserves in this region

of plants Thus plants tend to better tolerate insect pestattacks under field conditions than in the greenhouse

The increased B ruziziensis dry matter percentage dueto the enhancedM spectabilis infestation density herein wasalso demonstrated by Resende et al [29] in B ruziziensis 10days after exposure to 24M spectabilis adults in a greenhouseThose authors attributed the increased dry matter percent-age to the reduced fresh weight from insect damage Theincreased dry matter percentage was also reported for Bdecumbens infested withZulia entreriana adults [11] in alfalfared clover attacked by Philaenus spumarius [32 33] andDigitaria decumbens infested with Prosapia bicincta [34]

Despite the shoot damage from the M spectabilis attacksuch damage levels did not affect the B ruziziensis regrowthability under field conditions In contrast Resende et al [29]observed a significantly reduced number of B ruziziensisshoots infested with 12 or more M spectabilis adults in agreenhouse This observation is attributed to differences inthe fodder development conditions considering that hereinthe plants were grown directly in the groundwithout physicalconstraints to root system development Because differenceswere not observed in the fresh weight and forage regrowthafter the pest insect attack tactics for using such forage shouldbe adopted early during infestation before themore advancedsymptoms of attack by adults Among these tactics the forageby grazing animals or even cutting and storage of thismaterialin silos to be provided to cattle during the dry season arehighlighted According to Valerio [35] the forage root systemmay be reduced due to frequent spittlebug attacks whichresults in reduced forage persistence The plants used hereinwere grown under favorable conditions for extensive rootdevelopment and were not previously attacked by spittlebugsThese factors were important to maintain the number ofshoots after infestation these data emphasize that properpasture and soil fertility management guarantee forage re-establishment even after insect attacks

5 Conclusion

Thus attacks by 8 adult M spectabilis per clump of B ruz-iziensiswith an average of 80 tillers for six days were sufficientto reduce the chlorophyll content and cause functional plantloss index of 60 but it did not affect the fresh weight norregrowth capability which indicates that tactics for usingthis forage are recommended at the beginning of insectinfestation This population density can be a reference forspittlebug integrated management in Brachiaria

Acknowledgments

The authors thank the Conselho Nacional de Desenvolvi-mento Cientıfico e Tecnologico (CNPq Brazil) and FundacaodeAmparo a Pesquisa doEstado deMinasGerais (FAPEMIGBrazil) for supporting their research

References

[1] A J E Peron and A R Evangelista ldquoDegradacao de pastagensem regioes de cerradordquo Ciencia e Agrotecnologia vol 28 pp655ndash661 2004

The Scientific World Journal 5

[2] L Moreira and E D Assad Segmentacao e Classificacao super-visionada Para identificar Pastagens Degradadas Embrapa-CPAC Planaltina Goias 1997

[3] J R Valerio C Cardona D C Peck and G Sotelo SpittlebugsldquoBioecology host plant resistance and advances in IPMrdquo Pro-ceedings of the 19 International Grassland Congress pp 217ndash221 FEALQ Sao Pedro Piracicaba 2001

[4] S V Paula-Moraes Cadeia Produtiva da Carne E ameacadaPor Praga 2006 httpwwwcpacembrapabrmaterias pripagnucleotematicohtml

[5] AM Auad A D SimoesM V Leite S E B Silva D R Santosand P H Monteiro ldquoSeasonal dynamics of egg diapause inMahanarva spectabilis (Distant 1909) (Hemiptera Cercopidae)on elephant grassrdquo Arquivos do Instituto Biologico vol 78 pp325ndash330 2011

[6] D W Fewkes ldquoThe biology of sugar cane froghopperrdquo in Pestsof Sugar Cane J R Williams J R Metcalfe R W Mungomeryand R Mathes Eds pp 281ndash307 Elsevier Amsterdam TheNetherlands 1969

[7] A M Auad A D Simoes A V Pereira et al ldquoSelecao degenotipos de capim-elefante quanto a resistencia a cigarrinha-das-pastagensrdquo Pesquisa Agropecuaria Brasileira vol 42 pp1077ndash1081 2007

[8] V Thompson ldquoAssociative nitrogen fixation C4 photosynthe-sis and the evolution of spittlebugs (Hemiptera Cercopidae)as major pests of neotropical sugarcane and forage grassesrdquoBulletin of Entomological Research vol 94 no 3 pp 189ndash2002004

[9] R A Byers andH DWells ldquoPhytotoxemia of coastal bermuda-grass caused by the two-lined spittlebug Prosapia bicincta(Homoptera Cercopidae)rdquo Annals of the Entomological Societyof America vol 59 pp 1067ndash1071 1966

[10] Z A Ramiro R A Miranda and E A Batista Filho ldquoObser-vacoes sobre a fluctuacao de cigarrinhas (Homoptera Cercop-idae) em pastagem formale em Brachiaria decumbens mantidaem diferentes nıveis de desenvolvimento vegetativordquo Anais daSociedade Entomologica do Brasil vol 13 pp 357ndash370 1984

[11] J R Valerio and O Nakano ldquoDanos causados pelo adulto dacigarrinha-das-pastagens Zulia entreriana (Berg 1879) (Homo-ptera Cercopidae) em plantas de Brachiaria decumbens Stapfmantidas em diferentes nıveis de umidaderdquo Anais da SociedadeEntomologica do Brasil vol 16 pp 341ndash350 1988

[12] S L LapointeG Sotelo andG LArango ldquoImproved techniquefor rearing spittlebugs (Homoptera Cercopi-dae)rdquo Journal ofEconomic Entomology vol 82 pp 1764ndash1766 1989

[13] D C Peck ldquoNatural history of the spittlebug Prosapia bicincta(Homoptera Cercopidae) in association with dairy pastures ofCosta RicardquoAnnals of the Entomological Society of America vol91 no 4 pp 435ndash444 1998

[14] F Lopez C Cardona J W Miles G Sotelo and J Mon-toya ldquoScreening for resistance to adult spittlebugs (HemipteraCercopidae) in Brachiaria spp methods and categories ofresistancerdquo Journal of Economic Entomology vol 102 no 3 pp1309ndash1316 2009

[15] F Holmann and D C Peck ldquoEconomic damage caused by spit-tlebugs (Homoptera Cercopidae) in Colombia a first approxi-mation of impact on animal production in Brachiaria decum-bens pasturesrdquo Neotropical Entomology vol 31 no 2 pp 275ndash284 2002

[16] H F Kain and D S Atkison ldquoProblems of insect pest assess-ment in pasturesrdquo New Zealand Entomologist vol 6 no 1 pp9ndash13 1975

[17] J Diaz-Montano J C Reese W T Schapaugh and L R Camp-bell ldquoChlorophyll loss caused by soybean aphid (HemipteraAphididae) feeding on soybeanrdquo Journal of Economic Entomol-ogy vol 100 no 5 pp 1657ndash1662 2007

[18] C Cardona JWMiles andG Sotelo ldquoAn improvedmethodol-ogy formassive screening of Brachiaria spp genotypes for resis-tance toAeneolamia varia (Homoptera Cercopidae)rdquo Journal ofEconomic Entomology vol 92 no 2 pp 490ndash496 1999

[19] A Pabon C Cardona J W Miles and G Sotelo ldquoResponseof resistant and susceptible Brachiaria spp genotypes tosimultaneous infestation with multiple species of spittlebugs(Hemiptera Cercopidae)rdquo Journal of Economic Entomology vol100 no 6 pp 1896ndash1903 2007

[20] J Morgan G Wilde and D Johnson ldquoGreenbug resistance incommercial sorghum hybrids in the seedling stagerdquo Journal ofEconomic Entomology vol 73 pp 510ndash514 1980

[21] N Panda andEAHeinrichs ldquoLevels of tolerance and antibiosisin rice varieties having moderate resistance to the brown plan-thopper Nilaparvata lugens (Stal) (Hemiptera Delphacidae)rdquoEnvironmental Entomology vol 12 pp 1204ndash1214 1983

[22] C M Smith Z R Khan and E M D Pathak Techniques forEvaluating Insect Resistance in Crop Plants CRC Boca RatonFla USA 1994

[23] SISVAR Versao 53 (Biud 75) Sistemas de Analises de VarianciaPara Dados Balanceados Programa de Analises Estatısticase Planejamento de Experimentos Lavras MG UniversidadeFederal de Lavras 2010

[24] BIOESTAT Sistema Para Analises Estatısticas Versao 50 ParaUniversidade Federal do Para Belem Brazil 2007

[25] X Ni J P Wilson and G D Buntin ldquoDifferential responsesof forage pearl millet genotypes to chinch bug (HeteropteraBlissidae) feedingrdquo Journal of Economic Entomology vol 102no 5 pp 1960ndash1969 2009

[26] T Wang S S Quisenberry N I Xinzhi and V TolmayldquoAphid (Hemiptera Aphididae) resistance in wheat near-isogenic linesrdquo Journal of Economic Entomology vol 97 no 2pp 646ndash653 2004

[27] G BMartha Junior L G Barioni L Vilela and A O BarcellosldquoArea do piquete e taxa de lotacao no pastejo rotacionadordquoComunicado Tecnico 101 2003

[28] C Cardona J W Miles E Zuniga and G Sotelo ldquoIndepen-dence of resistance in Brachiaria spp to nymphs or to adultspittlebugs (Hemiptera Cercopidae) implications for breedingfor resistancerdquo Journal of Economic Entomology vol 103 no 5pp 1860ndash1865 2010

[29] T T Resende A M Auad M G Fonseca T H Santos andT M Vieira ldquoImpact of the spittlebug Mahanarva spectabilison signal grassrdquoTheScientificWorldJournal vol 2012 Article ID926715 6 pages 2012

[30] R H Painter Insect Resistance in Crop Plants MacMillan NewYork NY USA 1951

[31] J J Soares R S Melo C A Almeida and A M C FerreiraldquoProgresso no manejo integrado de pragas do algodoeiro noOeste Baianordquo Comunicado Tecnico 194 Embrapa AlgodaoCampina Grande Brazil 2003

[32] C R Weaver and J W Hibbs ldquoEffects of spittlebug infestationon nutrition value of alfalfa and red cloverrdquo Journal of EconomicEntomology vol 45 pp 626ndash628 1952

[33] R BMarthur andR L Pienkowski ldquoInfluence of adultmeadowspittlebug feeding on forage qualityrdquo Journal of EconomicEntomology vol 60 pp 207ndash209 1967

6 The Scientific World Journal

[34] E B Fagan Bionomics and control of the two-lined spittlebugProsapia bicincta on Florida pastures and notes on Prosapiaplagiata in Costa Rica (Homoptera Cercopidae) [Dissertation]University of Florida 1969

[35] J R Valerio ldquoConsideracoes sobre a morte de pastagensde Brachiaria brizantha cv Marandu em alguns Estados doCentro e Norte do Brasil enfoque entomologicordquo ComunicadoTecnico EmbrapaGado de Corte CampoGrande Brazil 2006

Submit your manuscripts athttpwwwhindawicom

Nutrition and Metabolism

Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Food ScienceInternational Journal of

Agronomy

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Microbiology

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom

Applied ampEnvironmentalSoil Science

Volume 2014

AgricultureAdvances in

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

PsycheHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BiodiversityInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

ScientificaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Plant GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Biotechnology Research International

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Forestry ResearchInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Journal of BotanyHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

EcologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Veterinary Medicine International

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Cell BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Evolutionary BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

The Scientific World Journal 5

[2] L Moreira and E D Assad Segmentacao e Classificacao super-visionada Para identificar Pastagens Degradadas Embrapa-CPAC Planaltina Goias 1997

[3] J R Valerio C Cardona D C Peck and G Sotelo SpittlebugsldquoBioecology host plant resistance and advances in IPMrdquo Pro-ceedings of the 19 International Grassland Congress pp 217ndash221 FEALQ Sao Pedro Piracicaba 2001

[4] S V Paula-Moraes Cadeia Produtiva da Carne E ameacadaPor Praga 2006 httpwwwcpacembrapabrmaterias pripagnucleotematicohtml

[5] AM Auad A D SimoesM V Leite S E B Silva D R Santosand P H Monteiro ldquoSeasonal dynamics of egg diapause inMahanarva spectabilis (Distant 1909) (Hemiptera Cercopidae)on elephant grassrdquo Arquivos do Instituto Biologico vol 78 pp325ndash330 2011

[6] D W Fewkes ldquoThe biology of sugar cane froghopperrdquo in Pestsof Sugar Cane J R Williams J R Metcalfe R W Mungomeryand R Mathes Eds pp 281ndash307 Elsevier Amsterdam TheNetherlands 1969

[7] A M Auad A D Simoes A V Pereira et al ldquoSelecao degenotipos de capim-elefante quanto a resistencia a cigarrinha-das-pastagensrdquo Pesquisa Agropecuaria Brasileira vol 42 pp1077ndash1081 2007

[8] V Thompson ldquoAssociative nitrogen fixation C4 photosynthe-sis and the evolution of spittlebugs (Hemiptera Cercopidae)as major pests of neotropical sugarcane and forage grassesrdquoBulletin of Entomological Research vol 94 no 3 pp 189ndash2002004

[9] R A Byers andH DWells ldquoPhytotoxemia of coastal bermuda-grass caused by the two-lined spittlebug Prosapia bicincta(Homoptera Cercopidae)rdquo Annals of the Entomological Societyof America vol 59 pp 1067ndash1071 1966

[10] Z A Ramiro R A Miranda and E A Batista Filho ldquoObser-vacoes sobre a fluctuacao de cigarrinhas (Homoptera Cercop-idae) em pastagem formale em Brachiaria decumbens mantidaem diferentes nıveis de desenvolvimento vegetativordquo Anais daSociedade Entomologica do Brasil vol 13 pp 357ndash370 1984

[11] J R Valerio and O Nakano ldquoDanos causados pelo adulto dacigarrinha-das-pastagens Zulia entreriana (Berg 1879) (Homo-ptera Cercopidae) em plantas de Brachiaria decumbens Stapfmantidas em diferentes nıveis de umidaderdquo Anais da SociedadeEntomologica do Brasil vol 16 pp 341ndash350 1988

[12] S L LapointeG Sotelo andG LArango ldquoImproved techniquefor rearing spittlebugs (Homoptera Cercopi-dae)rdquo Journal ofEconomic Entomology vol 82 pp 1764ndash1766 1989

[13] D C Peck ldquoNatural history of the spittlebug Prosapia bicincta(Homoptera Cercopidae) in association with dairy pastures ofCosta RicardquoAnnals of the Entomological Society of America vol91 no 4 pp 435ndash444 1998

[14] F Lopez C Cardona J W Miles G Sotelo and J Mon-toya ldquoScreening for resistance to adult spittlebugs (HemipteraCercopidae) in Brachiaria spp methods and categories ofresistancerdquo Journal of Economic Entomology vol 102 no 3 pp1309ndash1316 2009

[15] F Holmann and D C Peck ldquoEconomic damage caused by spit-tlebugs (Homoptera Cercopidae) in Colombia a first approxi-mation of impact on animal production in Brachiaria decum-bens pasturesrdquo Neotropical Entomology vol 31 no 2 pp 275ndash284 2002

[16] H F Kain and D S Atkison ldquoProblems of insect pest assess-ment in pasturesrdquo New Zealand Entomologist vol 6 no 1 pp9ndash13 1975

[17] J Diaz-Montano J C Reese W T Schapaugh and L R Camp-bell ldquoChlorophyll loss caused by soybean aphid (HemipteraAphididae) feeding on soybeanrdquo Journal of Economic Entomol-ogy vol 100 no 5 pp 1657ndash1662 2007

[18] C Cardona JWMiles andG Sotelo ldquoAn improvedmethodol-ogy formassive screening of Brachiaria spp genotypes for resis-tance toAeneolamia varia (Homoptera Cercopidae)rdquo Journal ofEconomic Entomology vol 92 no 2 pp 490ndash496 1999

[19] A Pabon C Cardona J W Miles and G Sotelo ldquoResponseof resistant and susceptible Brachiaria spp genotypes tosimultaneous infestation with multiple species of spittlebugs(Hemiptera Cercopidae)rdquo Journal of Economic Entomology vol100 no 6 pp 1896ndash1903 2007

[20] J Morgan G Wilde and D Johnson ldquoGreenbug resistance incommercial sorghum hybrids in the seedling stagerdquo Journal ofEconomic Entomology vol 73 pp 510ndash514 1980

[21] N Panda andEAHeinrichs ldquoLevels of tolerance and antibiosisin rice varieties having moderate resistance to the brown plan-thopper Nilaparvata lugens (Stal) (Hemiptera Delphacidae)rdquoEnvironmental Entomology vol 12 pp 1204ndash1214 1983

[22] C M Smith Z R Khan and E M D Pathak Techniques forEvaluating Insect Resistance in Crop Plants CRC Boca RatonFla USA 1994

[23] SISVAR Versao 53 (Biud 75) Sistemas de Analises de VarianciaPara Dados Balanceados Programa de Analises Estatısticase Planejamento de Experimentos Lavras MG UniversidadeFederal de Lavras 2010

[24] BIOESTAT Sistema Para Analises Estatısticas Versao 50 ParaUniversidade Federal do Para Belem Brazil 2007

[25] X Ni J P Wilson and G D Buntin ldquoDifferential responsesof forage pearl millet genotypes to chinch bug (HeteropteraBlissidae) feedingrdquo Journal of Economic Entomology vol 102no 5 pp 1960ndash1969 2009

[26] T Wang S S Quisenberry N I Xinzhi and V TolmayldquoAphid (Hemiptera Aphididae) resistance in wheat near-isogenic linesrdquo Journal of Economic Entomology vol 97 no 2pp 646ndash653 2004

[27] G BMartha Junior L G Barioni L Vilela and A O BarcellosldquoArea do piquete e taxa de lotacao no pastejo rotacionadordquoComunicado Tecnico 101 2003

[28] C Cardona J W Miles E Zuniga and G Sotelo ldquoIndepen-dence of resistance in Brachiaria spp to nymphs or to adultspittlebugs (Hemiptera Cercopidae) implications for breedingfor resistancerdquo Journal of Economic Entomology vol 103 no 5pp 1860ndash1865 2010

[29] T T Resende A M Auad M G Fonseca T H Santos andT M Vieira ldquoImpact of the spittlebug Mahanarva spectabilison signal grassrdquoTheScientificWorldJournal vol 2012 Article ID926715 6 pages 2012

[30] R H Painter Insect Resistance in Crop Plants MacMillan NewYork NY USA 1951

[31] J J Soares R S Melo C A Almeida and A M C FerreiraldquoProgresso no manejo integrado de pragas do algodoeiro noOeste Baianordquo Comunicado Tecnico 194 Embrapa AlgodaoCampina Grande Brazil 2003

[32] C R Weaver and J W Hibbs ldquoEffects of spittlebug infestationon nutrition value of alfalfa and red cloverrdquo Journal of EconomicEntomology vol 45 pp 626ndash628 1952

[33] R BMarthur andR L Pienkowski ldquoInfluence of adultmeadowspittlebug feeding on forage qualityrdquo Journal of EconomicEntomology vol 60 pp 207ndash209 1967

6 The Scientific World Journal

[34] E B Fagan Bionomics and control of the two-lined spittlebugProsapia bicincta on Florida pastures and notes on Prosapiaplagiata in Costa Rica (Homoptera Cercopidae) [Dissertation]University of Florida 1969

[35] J R Valerio ldquoConsideracoes sobre a morte de pastagensde Brachiaria brizantha cv Marandu em alguns Estados doCentro e Norte do Brasil enfoque entomologicordquo ComunicadoTecnico EmbrapaGado de Corte CampoGrande Brazil 2006

Submit your manuscripts athttpwwwhindawicom

Nutrition and Metabolism

Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Food ScienceInternational Journal of

Agronomy

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Microbiology

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom

Applied ampEnvironmentalSoil Science

Volume 2014

AgricultureAdvances in

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

PsycheHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BiodiversityInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

ScientificaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Plant GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Biotechnology Research International

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Forestry ResearchInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Journal of BotanyHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

EcologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Veterinary Medicine International

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Cell BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Evolutionary BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

6 The Scientific World Journal

[34] E B Fagan Bionomics and control of the two-lined spittlebugProsapia bicincta on Florida pastures and notes on Prosapiaplagiata in Costa Rica (Homoptera Cercopidae) [Dissertation]University of Florida 1969

[35] J R Valerio ldquoConsideracoes sobre a morte de pastagensde Brachiaria brizantha cv Marandu em alguns Estados doCentro e Norte do Brasil enfoque entomologicordquo ComunicadoTecnico EmbrapaGado de Corte CampoGrande Brazil 2006

Submit your manuscripts athttpwwwhindawicom

Nutrition and Metabolism

Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Food ScienceInternational Journal of

Agronomy

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Microbiology

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom

Applied ampEnvironmentalSoil Science

Volume 2014

AgricultureAdvances in

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

PsycheHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BiodiversityInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

ScientificaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Plant GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Biotechnology Research International

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Forestry ResearchInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Journal of BotanyHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

EcologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Veterinary Medicine International

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Cell BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Evolutionary BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Submit your manuscripts athttpwwwhindawicom

Nutrition and Metabolism

Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Food ScienceInternational Journal of

Agronomy

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

International Journal of

Microbiology

The Scientific World JournalHindawi Publishing Corporation httpwwwhindawicom Volume 2014

Hindawi Publishing Corporationhttpwwwhindawicom

Applied ampEnvironmentalSoil Science

Volume 2014

AgricultureAdvances in

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

PsycheHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

BiodiversityInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

ScientificaHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Plant GenomicsInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Biotechnology Research International

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Forestry ResearchInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Journal of BotanyHindawi Publishing Corporationhttpwwwhindawicom Volume 2014

EcologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Veterinary Medicine International

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Cell BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014

Evolutionary BiologyInternational Journal of

Hindawi Publishing Corporationhttpwwwhindawicom Volume 2014