Interciencia ISSN: 0378-1844 [email protected] Asociación Interciencia Venezuela

Capello, Soledad; Franceschini, María Celeste Number and duration of the juvenil e instars of the neotropical grasshopper cornops aquaticum (: ) in Argentina Interciencia, vol. 39, núm. 4, abril, 2014, pp. 255-259 Asociación Interciencia Caracas, Venezuela

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How to cite Complete issue Scientific Information System More information about this article Network of Scientific Journals from Latin America, the Caribbean, Spain and Portugal Journal's homepage in redalyc.org Non-profit academic project, developed under the open access initiative Number and duration of the juvenile instars of the Neotropical grasshopper Cornops aquaticum (Acrididae: Leptysminae) IN ARGENTINA

Soledad Capello and M. Celeste Franceschini

SUMMARY

Cornops aquaticum is a semiaquatic grasshopper that lives in ger nymphal development in females was determined by females close association with Pontederiaceae, especially the Eichhor- with six juvenile instars. Our results, as those from other South nia genus. In its native distribution range, information is only American locations about the life cycle of C. aquaticum, show available about life history traits of populations from Brazil that when the instar number and nymphal development are the and Uruguay. The aim of this work was to assess the number life history traits to be studied, rearing should be carried out of juvenile instars and duration of nymphal development of C. at several sites within its distribution, and in the periods when aquaticum in summer and spring nymphs from Argentinean newly hatched nymphs have peaks of abundance in wild popula- populations. Wild newly emerged nymphs were individually tions. Different juvenile instars in native populations respect to reared in Corrientes and Santa Fe cities under local climate, quarantine populations of C. aquaticum show the importance to without direct sun exposure. In both sites, nymphs showed consider life history traits throughout the native range and the males with five instars and females with five or six instars. Fe- effect of environmental factors as useful information for its pos- males with six instars were more frequent in summer nymphs. sible use as a biological control agent of Eichhornia crassipes The highest mortality occurred in the first two instars. Nymphal in non-native areas. development was longer in nymphs from the summer peak. Lon-

Introduction al., 2007; Franceschini, 2008; tant variations in its life history In this context, temperature and Capello, 2010). traits, mainly with respect to insolation have been mentioned Cornops aquaticum Bruner C. aquaticum seems to have the number of juvenile instars as key factors determining the (1906) is a semi-aquatic grass- originated in the Amazonian and duration of nymphal devel- number of juvenile instars and hopper that lives in close asso- floodplain (Amedegnato, 1977) opment (Zolessi, 1956; Silveira duration of nymphal develop- ciation with Pontederiaceae and currently is widely distrib- Guido and Perkins, 1975; Me- ment (Lhano, 2002; Adis et al., floating meadows, especially of uted in Latin America, from deiros, 1984; Hill and Oberhol- 2004; Capello et al., 2007). the genus Eichhornia (Center et Southeastern Mexico to central zer, 2000; Lhano, 2002; Vieira The length of the hind femur al., 2002). Nymphs and adults Argentina and Uruguay (Rob- and Santos, 2003; Adis and is the most constant and precise of this grasshopper feed on the erts and Carbonell, 1979; Lha- Junk, 2003; Adis et al., 2004; morphometric trait to distin- leaves of these macrophytes, no, 2006; Adis et al., 2007). Brede et al., 2007; Capello et guish each juvenile instar of C. and eggs are laid in an ootheca Climatic variables such as inso- al., 2007). Several scenarios aquaticum, because it is subject within the leaf petiole (Zolessi, lation, temperature and precipi- have been proposed to explain to little measuring bias and it is 1956). Currently, several aspects tation have been shown to vary the causes of these variations easy to measure when working of the life history of C. aquati- greatly over its area of distribu- (Adis et al., 2004). However, with a high number of individu- cum are intensely studied for its tion (Adis et al., 2004). the most plausible explanation als (Franceschini et al., 2005). possible release in non-native Recent studies carried out in seems to be that the variation is In Argentina, the reproduction ecosystems as a biological con- natural populations of C. aquat- a result of the phenotypic plas- of this grasshopper takes place trol agent of the water hyacinth icum from South America and ticity in response to different during spring and summer, Eichhornia crassipes (Lhano, quarantine populations reared in climatic factors of the respec- when abundance peaks of instar 2002; Adis et al., 2004; France- South Africa have confirmed tive environments where popu- I nymphs occur in populations schini et al., 2005; Capello et that this species shows impor- lations live (Brede et al., 2007). living on E. azurea (France-

Keywords / Eichhornia / Leptysminae / Life Cycle / Semiaquatic Grasshopper / Received: 07/13/2012. Accepted: 03/20/2014.

Soledad Capello. Biology Profes- Professor, Universidad de Con- María Celeste Franceschini: Biol- Universidad Nacional del sor and Specialist in Biodiver- cepción del Uruguay, Argentina. ogy Professor and Specialist in Nordeste, Argentina. e-mail: sity, Universidad Nacional del Address: Instituto Nacional de Entomology, Universidad Nacio- [email protected] Litoral, Argentina. Doctor in Limnología (INALI-CONICET- nal del Nordeste, Argentina. Natural Sciences, Universidad UNL). Ciudad Universitaria. Pa- Doctor in Natural Sciences, Uni- Nacional de La Plata, Argenti- raje El Pozo, (3000) Santa Fe, versidad Nacional de La Plata, na. CONICET Postdoctoral Fel- Argentina. e-mail: solecapello1@ Argentina. CONICET Research- low, Instituto Nacional de Lim- yahoo.com.ar er, Centro de Ecología Aplicada nología, Santa Fe, Argentina. del Litoral, Argentina. Professor,

APR 2014, VOL. 39 Nº 4 0378-1844/14/04/255-05 $ 3.00/0 255 Número y duración de los estadios ninfales de la tucura neotropical Cornops aquaticum (Acrididae: Leptysminae) en Argentina Soledad Capello y M. Celeste Franceschini RESUMEN Cornops aquaticum es un acridio semiacuático que vive en ninfas se produjo en los dos primeros estadios. La mayor du- estrecha asociación con Pontederiaceas, especialmente del gé- ración del desarrollo ninfal en las hembras estuvo determinada nero Eichhornia. En su área de distribución nativa, la infor- por aquellas de seis estadios. Estos resultados, y los de otros mación existente respecto al ciclo de vida esta referida a po- sitios de Sudamérica sobre el ciclo de vida de C. aquaticum, blaciones de Brasil y Uruguay. En este trabajo se determinó el muestran que al analizar el número de estadios y el desarrollo número de estadios ninfales y la duración del desarrollo ninfal ninfal de un acridio conviene realizar crías en distintos lugares de C. aquaticum en Argentina, en ninfas originadas en verano y y en periodos cuando las ninfas de estadio I presentan mayor primavera. Las crías se realizaron en dos localidades: Corrien- abundancia en las poblaciones naturales. Las diferencias en el tes y Santa Fe, utilizando ninfas recién eclosionadas en campo, ciclo de vida de C. aquaticum en ninfas nativas y en cuaren- criadas individualmente en recintos cerrados bajo influencia del tena muestran la importancia de considerar las particularida- clima local y sin exposición solar directa. En ambas localida- des del desarrollo ninfal en el área de distribución nativa y los des, los machos presentaron cinco estadios ninfales; las hem- factores climáticos implicados, como información valiosa ante bras cinco o seis. Las hembras con seis estadios fueron más su posible utilización en el control biológico de E. crassipes en frecuentes en las ninfas de verano. La mayor mortalidad de áreas no nativas.

Número E duraÇÃO dos estÁGIOS ninfaIs dO GAFANHOTO neotropical Cornops aquaticum (Acrididae: Leptysminae) NA Argentina Soledad Capello e M. Celeste Franceschini RESUMO Cornops aquaticum é um acrídeo semiaquático que vive em de ninfas se produziu nos dois primeiros estágios. A maior du- estreita associação com Pontederiáceas, especialmente do gê- ração do desenvolvimento ninfal nas fêmeas esteve determinada nero Eichhornia. Nos seus limites de distribuição nativa, a por aquelas de seis estágios. Estes resultados, e os de outros informação existente relativa ao ciclo de vida esta referida a lugares da América do sul sobre o ciclo de vida de C. aqua- populações de Brasil e Uruguai. Neste trabalho se determinou ticum, mostram que ao analisar o número de estágios e o de- o número de estágios ninfais e a duração do desenvolvimento senvolvimento ninfal de um acrídeo convêm realizar crias em ninfal de C. aquaticum na Argentina, em ninfas originadas no distintos lugares e em períodos quando as ninfas de estágios I verão e primavera. As crias se realizaram em duas localida- apresentam maior abundancia nas populações naturais. As di- des: Corrientes e Santa Fe, utilizando ninfas recém-eclodidas ferenças no ciclo de vida de C. aquaticum em ninfas nativas e em campo, criadas individualmente em recintos fechados sob em quarentena mostram a importância de considerar as parti- a influência do clima local e sem exposição solar direta. Em cularidades do desenvolvimento ninfal nos limites de distribui- ambas as localidades, os machos apresentaram cinco estágios ção nativa e os fatores climáticos implicados, como informação ninfais; as fêmeas cinco ou seis. Fêmeas com seis estágios fo- valiosa diante de sua possível utilização no controle biológico ram mais frequentes nas ninfas de verão. A maior mortalidade de E. crassipes em áreas não nativas.

schini et al., 2007) and E. enclosure under influence of in order to calculate the duration days/year in Corrientes and 1 crassipes floating meadows (Ca- the local climate, without di- of each nymphal instar and the day/year in Santa Fe (Bruniard, pello et al., 2004; Franceschini rect sun exposure. Rearings nymphal development. Experi- 1981). In both sites, the mean et al., 2008). were carried out in Corrientes ments were started with wild temperature during rearing var- In the native range of distri- City, in the northeast of Argen- newly emerged nymphs to in- ied from 13.9°C to 27.3°C. bution of C. aquaticum, infor- tina (27º28’14”S; 58º50’24”W), clude a high number of individu- Monthly mean of maximum mation is only available about and in Santa Fe City, 500km als in the data analysis; thus, the temperature varied from 30.3 to life history traits of populations further south (31°40’14”S; duration of first-instar nymphs 35.2°C in summer and from from Brazil and Uruguay. Thus, 60°34’44”W). was determined from nymphs 27.3 to 32.8°C in spring. the aim of this work was to as- Experiments started in Febru- that hatched from ovipositions Monthly mean of minimum sess the number of juvenile in- ary 2004 (Corrientes N= 29; contained in Eichhornia plants temperature varied from 16.8 to stars and duration of nymphal Santa Fe N= 29) and November in the summer of 2006. 22.2°C in summer and from development of C. aquaticum in 2004 (Corrientes N= 28; Santa 17.2 to 20.7°C in spring (Figure summer and spring nymphs Fe N= 26), in order to obtain Study sites 1). High levels of insolation from Argentinean populations. nymphs of C. aquaticum popula- were recorded during summer tions from the summer and The climate of the study area 2004 (241 to 351h of sun/ Material and Methods spring peaks, respectively (Ca- is classified as subtropical; sum- month); during spring, insola- pello et al., 2004; Franceschini mers are warm and prolonged, tion varied from 228 to 304h of Wild newly emerged nymphs et al., 2007, 2008). Observations while winters are shorter and sun/month. Total hours of sun were collected and individually were made every two days, re- mild (Bruniard, 1996), with a during 2004 were 2732 in Cor- reared in a screened outdoor cording the date of each moult frequency of frost days of 0.25 rientes and 2679 in Santa Fe.

256 APR 2014, VOL. 39 Nº 4 Statistical analysis est mortality of nymphs occurred mainly in the The frequency of occurrence first two instars (40-50%). of the different juvenile instars Development of was assessed by means of a nymphs from the summer contingence table. The duration peak took place more of nymphal development was slowly, from February to assessed using ANOVA with the end of June (summer post hoc Tukey tests, after hav- to beginning of winter), ing log10 transformed this vari- whereas development of able to normalize the distribu- nymphs from the spring tion and stabilize the variance. peak occurred from No- Values of p<0.05 were consid- vember to the end of De- ered significant. Statistical cember (spring to begin- analyses were performed using ning of summer). The InfoStat (2002). duration of each instar varied from 6 to 19 days Results in the instars I to IV, whereas in the instars V In the summer nymphs of to VI varied from 9.6 to Figure 1. Insolation (total hours of sun per month) and temperatures during the Corrientes, 72.2% of individuals 48 days. In the summer rearing of Cornops aquaticum nymphs in Corrientes and Santa Fe, Argentina. were males and 27.8% females; nymphs, the mean of Autumn is from Mar 21 to Jun 21 and spring is from Sept 21 to Dec 21. males required only five instars nymphal development to complete nymphal develop- reached 52 days in males ment, whereas females required and 81 days in females. In the comparing males and females show six or seven juvenile in- six. In the spring nymphs, spring nymphs, the mean of that developed with five juve- stars (Hill and Oberholzer, 52.9% of individuals were fe- nymphal development reached nile instars (ANOVA p=0.332). 2000). males and 47.1% males; all 49.6 days in males and 51 days High frequency of C. aquati- males developed with five in- in females (Tables I and II). Discussion cum nymphs with six instars in stars, whereas females devel- Nymphs from the spring peaks Argentina is coincident with oped with five or six instars had significantly shorter devel- All males of C. aquaticum high values of insolation record- (66.67% and 33.33 %, respec- opment times than nymphs reared in Argentina under the ed during summer 2004. This tively). from the summer peak (ANO- influence of the local climate result agrees with those found In the summer nymphs of VA p<0.0001). Nymphal devel- show five juvenile instars, by other authors (Lhano, 2002; Santa Fe, 31.25% of individuals opment was significantly longer whereas females develop with Adis and Junk, 2003; Vieira and were males and 68.75% fe- in females than in males five or six juvenile instars. Santos, 2003; Capello et al., males; males had five instars, (ANOVA p<0.0001). Compar- Nymphs of this species having 2007). In laboratory rearing, a while females went through five ing females that developed between five and six instars is a high frequency of females and or six instars (54.54% and with five and those that had six finding in agreement with the males with six instars has been 45.45%, respectively). In the instars, nymphal development results obtained in other sites of observed in C. aquaticum spring nymphs, 50% of indi- was significantly longer in fe- South America (Zolessi, 1954; nymphs reared in constant (24h) viduals were males and 50% males with six juvenile instars Medeiros, 1984; Adis and Junk, light at 27°C (Capello et al., females; all males had five in- (ANOVA p<0.0014). No sig- 2003; Vieira and Santos, 2003). 2007). In Manaus, where the stars, whereas females had five nificant differences in nymphal In non-native areas, quarantine insolation per year is lower and or six instars (60% and 40%, development were obtained populations of this grasshopper the monthly temperature is high- respectively). Frequency of females that TABLE I developed with six instars was Duration of the juvenile instars (from stage I to VI) and nymphal not significantly different be- development of Cornops aquaticum nymphs reared from the summer peak, tween sites (X2= 0.002; df= 1; in Corrientes and Santa Fe * p=0.96). In the summer Nymphal nymphs, females with six in- Males I II III IV V development stars were significantly more Corrientes 6.00 11.67 ±1.92 10.54 ±2.29 10.07 ±1.32 14.69 ±3.35 52.08 ±4.07 frequent than females with five Range - 9-15 7-15 7-13 10-22 41-57 instars. Conversely, in the Santa Fe 6.00 7.60 ±0.55 10.20 ±1.67 16.60 ±2.28 16.60 ±2.19 51 ±3.16 spring nymphs, females with Range - 7-8 8-12 7-19 14-20 48-56 five instars were significantly more frequent than females Females I II III IV V VI Nymphal with six instars (X2= 4.690; df= development 1; p=0.03). Corrientes 6.00 9.80 ±2.38 13.00 ±3.39 10.40 ±4.03 17.60 ±17.00 24.20 ±10.89 81 ±28.74 The survival rate of nymphs Range - 7-11 9-18 4-15 9-48 13-42 62-132 in Corrientes and Santa Fe was Santa Fe 6.00 9.91 ±1.04 11.27 ±2.00 11.55 ±1.81 17.45 ±5.84 22.71 ±5.15 70.64 ±10.24 higher in the summer (62.1 and Range - 9-12 8-15 9-14 12-33 16-30 55-82 55.17%, respectively) than in the spring nymphs (60.7 and Values are average ± standard deviation. Range: lowest and highest values recorded in the nymphal instars and 38.46%, respectively). The high- nymphal development.

APR 2014, VOL. 39 Nº 4 257 Table II cies of Cornops (Turk, 1984; Duration of the juvenile instars (from stage I to VI) and nymphal Turk and Aquino, 1996) and development of Cornops aquaticum nymphs reared from the spring peak, Leptysminae (Aquino and Turk, in Corrientes and Santa Fe * 1997; Nunes et al., 1992; Amor- Nymphal im and Adis, 1994, 1995). Males I II III IV V development Higher mortality of C. aquat- icum nymphs in the instar I and Corrientes 6 7.20 ± 0.82 7.00 ±1.30 7.75 ±1.04 12.38 ±2.50 40.33 ±2.40 II is also mentioned by Lhano Range - 7-9 4-8 7-10 9-16 36-76 (2002). Santa Fe 6 9.80 ± 2.49 10.00 ±2.24 10.40 ±0.89 11.80 ±1.48 49.60 ±6.02 Nymphal development of 81 Range - 8-14 7-13 9-11 10-14 44-59 days found in females from Nymphal the summer peak in Argentina Females I II III IV V VI development is a high mean value in com- Corrientes 6 8.75 ±2.27 7.56 ±3.28 6.89 ±3.26 11.67 ±3.57 13.00 ±3.05 45.24 ±6.07 parison with those mentioned Range - 4-12 2-14 3-14 7-16 10-16 36-55 for C. aquaticum nymphs from native population (Zo- Santa Fe 6 8.20 ±1.30 10.60 ±1.52 11.00 ±2.00 9.60 ±1.95 14.00 ±1.41 51 ±7.91 lessi, 1954; Silveira Guido and Range - 6-9 9-13 9-14 8-13 13-15 43-60 Perkins, 1975; Medeiros, 1984; Values are average ± standard deviation. Range: lowest and highest values recorded in the nymphal instars and Lhano, 2002; Adis and Junk, nymphal development. 2003; Vieira and Santos, 2003). A similar duration of nymphal development was ob- er than Corrientes and Santa Fe, reared under local photoperiod demonstrated that both climatic tained in males and females nymphs of C. aquaticum reared and temperatures from 22 to factors have a synergic effect. reared under constant (24h) under the influence of local cli- 30°C go through six or seven The number of juvenile instars dark at 25°C (Lhano, 2002). mate go through five instars juvenile instars (Hill and Ober- of this grasshopper has been In quarantine populations in during the rainy period (low in- holzer, 2000) (Figure 2). primarily related to the flood South Africa, nymphal devel- solation) and through six instars Adis et al. (2004) pointed pulse (Adis and Junk, 2003), opment takes 50 days with during the dry period (high in- that the number of juvenile in- quantity of food (Hill and temperature between 22 and solation) (Adis and Junk, 2003; stars and duration of nymphal Oberholzer, 2000) and rearing 30°C (Hill and Oberholzer, Vieira and Santos, 2003). development in C. aquaticum conditions (Medeiros, 1984). 2000). In laboratory rearing, In Curitiba, where the insola- mirrors the temperature and The variation in the number Capello et al.. (2007) find that tion per year is lower than in photoperiod of different climatic of juvenile instars, with addi- the duration of nymphal devel- Corrientes and Santa Fe, C. conditions in the respective re- tional instars in females, has opment in C. aquaticum under aquaticum nymphs reared under gions, and Capello et al. (2007) also been reported in other spe- constant long days with 24h of the local photoperiod at light is 33 days at 25°C have five instars 27°C and 75 days at in males, whereas fe- 36°C. The duration of males have predomi- nymphal development nately five instars and is shorter under con- only 5% of them have stant long days with six juvenile instars (Me- 24h of light in C. deiros, 1984). In Cuiabá aquaticum from the (Pantanal of Mato Gros- Pantanal of Mato Gros- so), where insolation is so (Lhano, 2002) and lower and temperature fissicauda is higher than in Corri- fissicauda from Ama- entes and Santa Fe, zonian (Amorim and males and females Adis, 1995). reared under the influ- Higher duration of the ence of the local cli- instars V and VI in mate have five and six comparison with the in- instars (Lhano, 2002; stars I to IV found in Adis et al., 2004). Al- this study is also men- though temperatures tioned to C. aquaticum and insolation in nymphs from the Ama- Carrasco, Uruguay, are zonian floodplain (Adis lower than in Corrientes and Junk, 2003). The and Santa Fe, the Figure 2. Summary of the variations in the number of juvenile instars and nymphal devel- duration of nymphal de- nymphs of C. aquati- opment of Cornops aquaticum in native and quarantine populations. Circles indicate an- velopment in summer cum develop with six nual average insolation; the darker circles indicate higher number of hours of sun per year. nymphs from the center juvenile instars (Zolessi, A: Manaus, Brazil (Adis and Junk, 2003; Vieira and Santos, 2003); B: Cuiabá, Brazil of Argentina (Silveira (Lhano, 2002; Adis et al., 2004); C: Curitiba, Brazil (Medeiros, 1984); D: Carrasco, Uru- 1956). In quarantine guay (Zolessi, 1954); E and F: data from Corrientes and Santa Fe, Argentina; G: Pretoria, Guido and Perkins, populations in Pretoria, South Africa (Hill and Oberholzer, 2000). Values in parenthesis indicate minor frequency 1975) is similar to South Africa, nymphs of occurrence in number of juvenile instars; Nd indicates number of days to complete nymphal development of of this grasshopper nymphal development. the spring nymphs from

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Paris, USA. pp 41-64 (Bruner, 1908) (Orthoptera: study in a given species, rearing France. 385 pp. Franceschini MC, Capello S, Lhano Acrididae) em um lago de should be carried out at several Amorim MA, Adis J (1994) Con- M, Adis J, de Wysiecki ML várzea da Amazônia Central, geographical sites of its distribu- sumo de alimento por um ga- (2005) Morfometría de los esta- Brasil. Bol. Mus. Para. Emílio tion, and in the periods when fanhoto neotropical, Stenacris dios ninfales Cornops aquati- Goeldi sér. Zool. 8: 349-374. new hatched nymphs have the fissicauda fissicauda (Bruner, cum Bruner (1906) (Acrididae: Roberts HR, Carbonell CS (1979) A 1908) Orthoptera: Acrididae) da peaks of highest abundance in Leptysminae) en Argentina. revision of the genera Stenopo- várzea Amazônica. Acta Amaz. Amazoniana 18 (3-4): 373-386. la and Cornops (Orthoptera: the wild populations. Different 24: 289-302. Acrididae: Leptysminae). Proc. juvenile instars in native popula- Franceschini MC, Adis J, Poi de Amorim MA, Adis J (1995) Desen- Neiff A, de Wysiecki ML Acad. Nat. Sci. Philadelphia. tions respect to quarantine popu- volvimento ninfal do gafanhoto (2007) Fenología de Cornops 131: 104-130. lations of C. aquaticum show Neotropical semi-aquático, aquaticum Bruner (Orthoptera: Silveira Guido A, Perkins BD the importance to consider life Stenacris fissicauda fissicauda Acrididae: Leptysminae), en un (1975) Biological and host (Bruner, 1908) (Orthoptera: camalotal de Eichhornia azurea specificity of Cornops aquati- history traits throughout the na- Acrididae) em condições con- tive range and the effect of envi- (Pontederiaceae) en Argentina. cum (Bruner) (Orthoptera: Ac- troladas. Acta Amaz. 25: 73-92. Amazoniana 19: 149-158. rididae), a potential biological ronmental factors as useful in- Aquino AL, Turk SZ (1997) Ciclo Franceschini MC, Adis J, Poi de control agent for waterhyacinth. formation for the possible use of vital de Leptysma argentina Neiff A (2008) Phenology of Environ. Entomol. 4: 400-404. this grasshopper as a biological Bruner 1906 (Acrididae: Lep- Cornops aquaticum Br uner Turk SZ (1984) Acrididos del NOA control agent of Eichhornia tysminae: Leptysmini). Variabi- (Orthoptera: Acrididae) on VI: El ciclo de vida de Cornops crassipes in non-native areas. lidad en el esquema pre-repro- Eichhornia crassipes floating frenatum cannae Roberts & ductivo y reproducción. Acta meadows in a wetlands of the Carbonell (Acrididae: Leptys- Entomol. Chil. 21: 93-99. Paraná River Floodplain (Ar- minae) con especial referencia a Acknowledgments Brede EG, Adis J, Schneider P gentina).: 8th INTECOL Inter- su oviposición endofítica. Rev. (2007) What is responsible for national Wetlands Confer- Soc. Entomol. Arg. 43: 91-100. This research is dedicated to the variance in life history ence. (July 2008). Cuiabá, Turk SZ, Aquino AL (1996) Aci- the late Joachim U. Adis (Trop- traits of South American semi- Brazil. p. 80. doideos del NOA VIII: Nuevo aquatic grasshopper (Cornops Franceschini MC (2008) Biología y aporte a la bioecología y distri- ical Ecology Group of the Max- aquaticum)? A test of three Ecología de la Tucura Semiac- bución del genero Cornops Planck Limnology Institute, possible hypotheses. Stud. uática Cornops aquaticum (Or- Stal: Cornops paraguayense Neotrop. Fauna Environ. 42: Plön, Germany), who was in- thoptera: Acrididae: Leptys- (Br.) (Acrididae: Leptysminae: 225-233. volved in coordinating and dis- minae) en Relación a Dos Mac- Tetrataeniini). Acta Zool. Lill. cussing the Cornops studies and Bruniard ED (1981) El Clima de las rófitas Dominantes en Hume- 42: 427-432. gave us many ideas for carrying Planicies del Norte Argentino. dales del Nordeste de Argenti- Vieira MF, Santos AC (2003) Dura- Vol. I-II. Resistencia, Argenti- na: Eichhornia crassipes y E. ção do ciclo de vida de Cor- out this study, which is includ- na. Universidad Nacional del ed in the International Project azurea (Pontederiaceae). Thesis nops aquaticum (BRUNER, Nordeste. 379 pp. Universidad Nacional de La 1906) (Othoptera: Acrididae: “Host- Co-evolution on Bruniard ED (1996) Geografía de Plata. Argentina. 222 pp. Leptysminae) e aspectos de seu Water Hyacinth” (HICWA) ini- los Climas y de las Formacio- Hill MP, Oberholzer IG (2000) Host comportamento alimentar na tiated by Dr. Adis. Field trips nes vegetales (Aportes para un specifity of the grasshopper, Amazonia central. Acta Amaz. and laboratory tests were fund- Modelo Fitoclimático Mundial). Cornops aquaticum a natural 33: 711-714. Las Zonas Térmicas y la Vege- enemy of water hyacinth. In Zolessi LC (1956) Observaciones ed by the HICWA Project of tación Natural. EUDENE. Re- the Max-Planck Limnology In- Neal RS (Ed.) Proc. X Int. sobre Cornops aquaticum Br. sistencia, Argentina. 382 pp. Symp. on Biological Control of (Acridoidea, Cyrtacanthacr.) en stitute, PICT 2160-2011 FON- Capello S, Adis J, Marchese M Weeds. (1999) Bozeman, Mon- el Uruguay (1). Rev. Soc. Urug. CYT, PIP 6316 of CONICET. (2004) Fenología de Cornops tana, USA. pp. 349-356. Enomol. 1: 3-28.

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