Natural ocurrence of hymenopterous parasitoids associated with Anastrepha fraterculus (Diptera: Tephritidae) in Myrtaceae species in Entre Rios, Northeastern Argentina.
Parasitoids of Anastrepha fraterculus (Wiedemann) were monitored on ripe fruit of 3 native and 1 exotic, wild Myrtaceae species in the Province of Entre Rios, NE Argentina, between Jan and Mar 1993 and 1994 with the aim of identifying indigenous parasitoid par·a·sit·oid
Any of various insects, such as the ichneumon fly, whose larvae are parasites that eventually kill their hosts.
Of or relating to a parasitic insect of this kind. species and determining natural parasitization rates and fruit infestation infestation /in·fes·ta·tion/ (-fes-ta´shun) parasitic attack or subsistence on the skin and/or its appendages, as by insects, mites, or ticks; sometimes used to denote parasitic invasion of the organs and tissues, as by helminths. levels. The fruit species surveyed were Psidium guajava L. (common guava guava (gwä`və), small evergreen tree or shrub of the genus Psidium of the family Myrtaceae (myrtle family), native to tropical America and grown elsewhere for its ornamental flowers and edible fruit. ), Feijoa fei·jo·a
An evergreen shrub (Acca sellowiana syn. Feijoa sellowiana) native to South America and cultivated as an ornamental and for its sweet, tart fruit. Also called pineapple guava. sellowiana (O. Berg) O. Berg (feijoa), Eugenia uniflora Noun 1. Eugenia uniflora - Brazilian tree with spicy red fruit; often cultivated in California and Florida
pitanga, Surinam cherry
Eugenia, genus Eugenia - tropical trees and shrubs with aromatic leaves and often valuable hard wood L. (Surinam cherry), and Myrcianthes pungens Myrcianthes pungens is a species of plant in the Myrtaceae family. It is endemic to Brazil. Source
Larvae: see lemures. per kg of fruit, respectively. Highest levels of parasitism parasitism: see parasite.
Relationship between two species in which one benefits at the expense of the other. Ectoparasites live on the body surface of the host; endoparasites live in their hosts' organs, tissues, or cells and often rely were recorded in P. guajava, whereas M. pungens had the lowest parasitization rates. However, overall mean parasitism levels (i.e., considering all parasitoid species) did not appear to have great differences when comparing Myrtaceae species, collection sites, and years. The relative abundance and parasitization rates data of the recovered parasitoids in the 4 Myrtaceae species suggest some degree of host plant preference by U. anastrephae and D. brasiliensis.
Key Words: fruit flies, parasitoids, Braconidae, Figitidae, Argentina, Myrtaceae
Con el proposito de identificar especies de parasitoides asociadas con Anastrepha fraterculus (Wiedemann) y determinar niveles de parasitismo natural y de infestacion en fruta en la provincia de Entre Rios, Noreste de Argentina, se colectaron tres especies nativas y una exotica ex·ot·i·ca
Things that are curiously unusual or excitingly strange: such gustatory exotica as killer bee honey and fresh catnip sauce. de Myrtaceae entre Enero y Marzo de 1993 y de 1994. Las especies frutales colectadas fueron Psidium guajava L. (guayaba comun), Feijoa sellowiana (O. Berg) O. Berg (feijoa), Eugenia uniflora L. (arrayan), y Myrcianthes pungens (Berg) Legrand (mato). Se obtuvo un total de 2.186 puparios, de los cuales el 95% correspondio a A. fraterculus y el 5% restante a Ceratitis capitata (Wiedemann). De los 1.667 insectos adultos que emergieron de estos puparios, 1.378 fueron A. fraterculus, 89 C. capitata y 200 fueron parasitoides larvo-pupales, representando cuatro cuat·ro
n. pl. cuat·ros
A small guitarlike instrument of Latin America, usually having four or five pairs of strings.
[Spanish, from Latin quattuor, four; see quatrain.] especies de dos familias de Hymenoptera: Doryctobracon areolatus (Szepligeti), D. brasiliensis (Szepligeti), Utetes anastrephae (Viereck) (todos Braconidae, Opiinae), yAganaspis pelleranoi (Brethes) (Figitidae, Eucoilinae). Todas estas especies de parasitoides son nuevas citas para la provincia de Entre Rios. Ademas, estos registros representan el rango natural de distribucion mas austral en el continente americano para estas cuatro especies de parasitoides. Doryctobracon areolatus y A. pelleranoi fueron recuperados de todas las especies de Myrtaceae colectadas, y fueron las especies de parasitoides mas abundantes. Los niveles de infestacion por A. fraterculus en mato, arrayan, guayaba, y feijoa variaron de 15.2 a 41.8, 21.3 - 49.4, 34.1-109.2, y 78.9 -140.6 larvas por kg de fruta, respectivamente. Sin embargo, los valores medios totales de parasitismo (incluyendo todas las especies de parasitoides) no presentaron grandes diferencias cuando se consideraron las distintas especies de Myrtaceae, lugares de colecta, y anos de muestreo. Los datos sobre abundancia relativa de parasitoides y las tasas de parasitismo en las cuatro especies de Myrtaceae sugieren cierto grado de preferencia por la planta hospedera por parte de los parasitoides U. anastrephae y D. brasiliensis.
Translation provided by the authors.
The South American fruit fly, Anastrepha fraterculus (Wiedemann), and the Mediterranean fruit fly Mediterranean fruit fly: see fruit fly.
Mediterranean fruit fly
or Med fly
Fruit fly (Ceratitis capitata) proven to be particularly destructive to citrus crops, at great economic cost. , Ceratitis capitata (Wiedemann), are among the most significant insect pests of edible fruit in Argentina (Spinetta 2004) and in the remaining South American countries (Malavasi et al. 2000; Aluja et al. 2003a). The native A. fraterculus is basically restricted to NE and NW Argentina, where the climate mainly is warm and humid, whereas the exotic C. capitata is currently distributed throughout all fruit-growing regions of the country (Ovruski et al. 2003; Segura et al. 2006). Although A. fraterculus represents a cryptic species ensemble (Steck 1991; Aluja et al. 2003a; Hernandez-Ortiz et al. 2004; Vera et al. 2006) distributed throughout continental America from Mexico to Argentina (Aluja 1999; Norrbom 2004), the Argentine populations of the complex belong to a single biological species (Alberti et al. 2002).
In the northeastern province of Entre Rios, where A. fraterculus and C. capitata coexist in large citrus crop areas (Segura et al. 2006), common guavas grown in thin patches of wild vegetation adjacent to commercial orchards support larger local A. fraterculus populations (Turica & Mallo 1961; Putruele 1996). Though A. fraterculus has one of the broadest host ranges of all known Anastrepha species (Norrbom 2004), fruit in the Myrtaceae family are among its favored host plant (Aluja 1999; Raga et al. 2005).
Poisoned bait sprays for A. fraterculus and C. capitata control were tried with some success as early as 1930 in Entre Rios (Ovruski & Fidalgo 1994). Some attempts to develop biological control programs were made in the 1940s and the 1960s. Thus, 3 exotic larval larval
1. pertaining to larvae.
see cutaneous and visceral larva migrans. parasitoid species (Tetrastichus giffardianus Silvestri, Aceratoneuromyia indica (Silvestri), and Diachasmimorpha longicaudata (Ashmead)) were introduced and released in limited numbers in citrus crop areas of Entre Rios (Ovruski et al. 1999). Tetrastichus giffardianus and A. indica are 2 gregarious gre·gar·i·ous
1. Seeking and enjoying the company of others; sociable. See Synonyms at social.
2. Tending to move in or form a group with others of the same kind: gregarious bird species. eulophid parasitoids originally collected in West Africa West Africa
A region of western Africa between the Sahara Desert and the Gulf of Guinea. It was largely controlled by colonial powers until the 20th century.
West African adj. & n. and Southeast Asia, respectively, whereas D. longicaudata is a solitary braconid brac·o·nid
Any of several ichneumon flies of the family Braconidae, the larvae of which are parasitic on other insects.
[From New Latin Braconidae, family name, possibly from Greek parasitoid originally from the Malaysia-Philippine region (Ovruski et al. 2000). Only A. indica and D. longicaudata were recovered (Turica 1968).
Studies on native parasitoids attacking tephritid pests in northeastern Argentina have been largely neglected. Examples of fruit fly parasitoid surveys are only those developed in Misiones (Ogloblin 1937) and Corrientes (Turica & Mallo 1961; Ovruski & Schliserman 2003a). These studies were focused largely on commercial fruit in citrus growing areas. Some parasitoid species also were reported from fruit samples sporadically collected in Entre Rios (Blanchard 1947) and Corrientes (Brethes 1924; Blanchard 1966). Most of the available new information on native fruit fly parasitoids of Argentina is based on the surveys carried out in NW Argentina (Turica & Mallo 1961; Nasca 1973; Fernandez de Araoz & Nasca 1984; Ovruski 1995; Wharton et al. 1998; Ovruski & Schliserman, 2003b; Schliserman et al. 2004; Schliserman & Ovruski, 2004; Ovruski et al. 2004, 2005, 2006; Orono et al. 2005). As many as 11 native parasitoid species associated with A. fraterculus on native and exotic host fruit species have been so far recorded in Argentina (Ovruski et al. 2005). Most of them also have been found parasitizing diverse Anastrepha species in several Latin American countries, including Brazil (Leonel et al. 1995; Aguiar-Menezes & Menezes 1997; Canal & Zucchi 2000; Guimaraes et al. 2000; Carvalho 2001; Aguiar-Menezes et al. 2001; Uchoa-Fernandes et al. 2003), Venezuela (Katiyar et al. 1995), Colombia (Yepes and Velez 1989; Carrejo & Gonzalez 1999), Costa Rica (Jiron & Mexzon 1989; Wharton et al. 1981), Guatemala (Eskafi 1990), and Mexico (Aluja et al. 1998, 2003b; Lopez et al. 1999; Sivinski et al. 1997, 2000, 2001; Guillen et al. 2002; Hernandez-Ortiz et al. 1994, 2006).
The specific aims of this study were to survey selected wild Myrtaceae species commonly infested in·fest
tr.v. in·fest·ed, in·fest·ing, in·fests
1. To inhabit or overrun in numbers or quantities large enough to be harmful, threatening, or obnoxious: by A. fraterculus in order to provide information on infestation levels in the fruit sampled, degree of larval parasitization, and the parasitoid fauna attacking this fruit fly pest in Entre Rios, NE Argentina, as well as to document natural distribution ranges of these parasitoids along a latitudinal gradient. This article complements our previous study on the diversity and distribution of native fruit fly parasitoid in the extremely endangered subtropical sub·trop·i·cal
Of, relating to, or being the geographic areas adjacent to the Tropics.
of the region lying between the tropics and temperate lands
rainforests in north Argentina (Ovruski et al. 2004, 2005).
MATERIALS AND METHODS
Collections were made during Jan-Mar 1993 and 1994 in the following localities of Entre Rios, NE Argentina: Parana (31[degrees]64'S, 60[degrees]32'W elevation 90 m), La Paz La Paz, city, Bolivia
La Paz (lä päs), city (1992 pop. 713,378), W Bolivia, administrative capital (since 1898) and largest city of Bolivia. The legal capital is Sucre. (30[degrees]44'S, 59[degrees]38'W 75 m), and Concordia (31[degrees]24'S, 58'01'W 48 m). The climate is characterized as temperate-humid with long and warm summers, the temperature of the hottest month being >22[degrees]C and the temperature of the coldest month being <18[degrees]C; it rains all year long, but the proportion of winter rainfall is <5% of the total, the rainfall being between 933 mm (Cakf (w)) (Anonymous 1992).
Fruit samples included 3 native and 1 exotic, wild species of Myrtaceae: Psidium guajaua L. (common guava) (exotic species), Feijoa sellowiana (O. Berg) O. Berg (feijoa),Eugenia unifora L. (Surinam cherry), and Myrcianthes pungens (Berg) Legrand (mato) (all native species). Fruit were collected in backyard gardens in suburban areas and patches covered with wild native vegetation adjacent to small citrus orchards (sour and sweet oranges, tangerines, lime, sweet lemon, and grapefruit). All these fruit sampling sites were located in a linear corridor along the banks of Parana River (West sector of the NE Argentina region), except for the study site in Concordia, which was located close to the banks of Uruguay River Uruguay River
River, southeastern South America. Rising in southern Brazil, it forms the border between Argentina and Brazil and between Argentina and Uraguay. Above Buenos Aires, it combines with the Paraná River to form the great estuary of the Río de la Plata. (East sector of the NE Argentina region). The original native vegetation was subtropical rainforest locally known as "Selva en Galeria", which continued along the banks of southern and northern Parana and Uruguay River (Cabrera 1976).
Based on fruit size (measured as individual fruit weight) of each plant species, approximately 10 fruit were randomly taken from each guava and feijoa tree, and 35 fruit from each Surinam cherry and mato tree. Guava and feijoa plants had medium size fruit (mean [+ or -] SD: 45.8 [+ or -] 4.9 g and 39.2 [+ or -] 5.8 g, n = 100, respectively) while Surinam cherry and mato plants had small size fruit (mean [+ or -] SD:6.8 [+ or -] 2.3g,and7.2 [+ or -] 5.8g,n=100,respectively). Fruit was picked both from the tree and from the ground under the tree canopy. Therefore, each sample included fruit collected from tree plus fruit of the same tree that had fallen on the ground. Only ripe fruit that was about to fall from the tree was harvested. This allowed fruit fly larvae to complete development, and gave fruit fly parasitoids the opportunity to parasitize par·a·sit·ize
To live on or in a host as a parasite.
to live on or within a host as a parasite. larvae throughout their development (Lopez et al. 1999). Fruit samples were placed in styrofoam boxes with sand in the bottom as pupation pu·pate
intr.v. pu·pat·ed, pu·pat·ing, pu·pates
1. To become a pupa.
2. To go through a pupal stage.
pu·pa medium for larvae, and they were taken to the laboratory. All material collected in the field was processed in the laboratory of the CIRPON institute (Centro de Investigaciones para la Regulacion de Poblaciones de Organismos Nocivos) in San Miguel de Tucuman San Mi·guel de Tu·cu·mán or Tucumán
A city of northern Argentina at the foot of an eastern range of the Andes north-northwest of Córdoba. The country's independence was proclaimed here in July 1816. Population: 833,000. (26[degrees]50'S, 65[degrees]13'W, elevation 426 m), Tucuman, Northwestern Argentina. Each styrofoam box contained only one fruit sample and all cages were kept inside a room at 27 [+ or -] 2[degrees]C and 60 [+ or -] 10% relative humidity relative humidity
The ratio of the amount of water vapor in the air at a specific temperature to the maximum amount that the air could hold at that temperature, expressed as a percentage. . Fruit fly puparia were recovered weekly during 1 month and then transferred to plastic trays containing sterilized humid sand, which was re-moistened every 3 d until all samples had been processed. Each tray was then placed inside a sealed wooden box. All wooden cages were kept inside a rearing room at 25 [+ or -] 1[degrees]C and 75 [+ or -] 5% relative humidity for 4 months. Puparia of C. capitata and Anastrepha Schiner were separated based on pupal pu·pa
n. pl. pu·pae or pu·pas
The nonfeeding stage between the larva and adult in the metamorphosis of holometabolous insects, during which the larva typically undergoes complete transformation within a protective cocoon or characters (White & Elson-Harris 1992). Emerged flies and/or parasitoids were identified, counted and sexed by sample and summarized for each locality. Afterward, unemerged puparia also were counted. Additional samples of parasitoid specimens collected in El Palmar El Palmar is the name of several geographical locations:
Parasitoid specimens were identified to species by S. Ovruski using Wharton and Marsh's (1978) key and the taxonomic tax·o·nom·ic also tax·o·nom·i·cal
Of or relating to taxonomy: a taxonomic designation.
tax descriptions by Wharton et al. (1998). Fruit flies were identified by S. Ovruski using Zucchi's (2000) taxonomic key. Nomenclature for the Opiinae follows Wharton (1997) and for the Eucoilinae Wharton et al. (1998). Parasitoid and fly specimens were placed in the entomological en·to·mol·o·gy
The scientific study of insects.
ento·mo·log collection of the Fundacion Miguel Lillo (FML Fml Formal
FML Football Manager Live (video game)
FML Field Manipulation Language
FML Forms Markup Language
FML Ferret Mailing List
FML Fibre Metal Laminate ) (San Miguel de Tucuman, Argentina). All plant samples were compared to herbarium herbarium, collection of dried and mounted plant specimens used in systematic botany. To preserve their form and color, plants collected in the field are spread flat in sheets of newsprint and dried, usually in a plant press, between blotters or absorbent paper. specimens at the FML, and identified by Alejandra Roldan (Facultad de Ciencias Naturales e Instituto Miguel Lillo, Universidad Nacional de Tucuman). Nomenclature employed for plant identification was based on Morales et al. (1995).
All parasitism values and fruit infestation levels reported here are based on the number of emerging adult flies and parasitoids (Lopez et al. 1999), and on the number of fruit fly larvae per kg of fruit (Aluja et al. 2000), respectively. Where appropriate, means and standard deviations (mean [+ or -] SD) were calculated as summary statistics for the parasitism percentage, fruit size (weight), and ovipositor ovipositor /ovi·pos·i·tor/ (o?vi-pos´it-er) a specialized organ by which many female insects deposit their eggs.
a specialized organ by which many female insects deposit their eggs. length of opiine parasitoid species.
A total of 1,532 fruit, representing 33.1 kg was processed during this study. Out of these, 420 (3.1 kg) fruit were E. unifora, 310 (3.2 kg) M. pungens, 615 (23.8 kg) P. guajaua, and 107 (3.0 kg) F. sellowiana. In total, 2,186 tephritid puparia were obtained from these fruit, 95% of which were A. fraterculus and 5% of which were C. capitata. Of 1,667 adult insects that emerged from these puparia, 1,378 were A. fraterculus, 89 C. capitata, and 200 parasitoids, representing 4 species of 2 Hymenoptera families. No parasitoids were recovered from C. capitata puparia.
Only E. unifora and P. guajaua were infested by C. capitata, and infestation rates ranged from 0.1 to 1.9 and from 4.3 to 15.21arvae/kg of fruit, respectively. 0.2 Approximately 17% and 33% of all Surinam cherry and guava samples were simultaneously infested by both C. capitata and A. fraterculus. Infestation patterns by A. fraterculus in M. pungens, E. unif ora, P. guajaua, and F. sellowiana varied from 15.2 to 41.8, 21.3 to 49.4, 34.1 to 109.2, and 78.9 to 140.6 larvae/kg of fruit, respectively.
Table 1 summarizes A. fraterculus and parasitoid species abundance, and parasitization rates based on fruit species collected per study site and year. Three species of opiine Braconidae parasitoids and 1 species of eucoiline Figitidae parasitoid were identified: Doryctobracon areolatus (Szepligeti), D. brasiliensis (Szepligeti), Utetes anastrephae (Viereck) (Braconidae), and Aganaspis pelleranoi (Brethes) (Figitidae). These species constituted 45.4, 14.0, 12.5, and 28.1% of the total recovered parasitoids, respectively. Doryctobracon areolatus was the most abundant parasitoid species in 71% of total fruit samples and it was recovered from all of the Myrtaceae species sampled. Aganaspis pelleranoi was also found in association with A. fraterculus in E. unifora, F. sellowiana, M. pungens and P. guajaua. This eucoiline parasitoid was recovered from 52% of the total fruit samples. Doryctobracon brasiliensis was recovered only from 33 and 50% of the total guava and feijoa samples, respectively. Approximately 76% of all U. anastrephae specimens was recovered from both M. pungens and E. unifora. This opiine species was obtained from 8, 20, 67, and 100% of all guava, feijoa, mato, and Surinam cherry samples, respectively.
Highest levels of parasitism were recorded in P. guajaua, whereas M. pungens had the lowest parasitization rates (Table 1). However, overall mean parasitism levels (i.e., considering all parasitoid species) did not appear to have great differences when comparing Myrtaceae species, collection sites, and year (Table 1). Parasitization by A. pelleranoi has accounted for only [less than or equal to] 10% of the total parasitization of A. fraterculus on F. sellowiana, M. pungens and E. unif ora. In contrast, in P. guajava the degree of larval parasitization by eucoiline species ranged from 24 to 73% of the total parasitization.
From the additional parasitoid specimens collected by colleagues in Entre Rios, 9 A. pelleranoi and 8 D. areolatus were identified. Six D. areolatus were recovered from guavas collected in El Palmar, whereas the remaining 2 D. areolatus and 9 A. pelleranoi were obtained from guavas in Concordia.
New fruit fly parasitoid species records for Entre Rios are D. areolatus, D. brasiliensis, U. anastrephae and A. pelleranoi, all native of the Neotropical region. Prior to this report, only the eucoiline Rhoptromeris haywardi (Blanchard) had been recorded from both C. capitata and A. fraterculus in Entre Rios (Blanchard 1947). All of these species belong to the fruit fly parasitoid guild number 2 described by Ovruski et al. (2000), which is characterized by solitary, koinobiont larval-pupal endoparasitoids ofAnastrepha spp. The braconids D. areolatus and U. anastrephae, and the figitid A. pelleranoi are widely distributed Adj. 1. widely distributed - growing or occurring in many parts of the world; "a cosmopolitan herb"; "cosmopolitan in distribution"
bionomics, environmental science, ecology - the branch of biology concerned with the relations between organisms in Latin American, while D. brasiliensis is known to occur in southern Brazil and northern Argentina (Ovruski et al. 2005). The discovery of these 4 native A. fraterculus-parasitoid species in Entre Rios represents their southernmost natural distribution range in the Americas. During this study, D. brasiliensis and U. anastrephae were found at 31[degrees]64'S latitude (Parana), whereas both D. areolatus and A. pelleranoi were also collected at 32[degrees]16'S latitude (El Palmar). Previously, these parasitoid species had been recorded from A. fraterculus in the provinces of Corrientes (27[degrees]15'-30[degrees]43'S and 59[degrees]41'-56[degrees]12'W) (Ovruski & Schliserman 2003a), Misiones (25[degrees]30'-28[degrees]10'S and 53[degrees]38'-56[degrees]03'W) (Ogloblin 1937; Turica & Mallo 1961), Catamarca (25[degrees]09'-29[degrees]16'S and 65[degrees]25'-69[degrees]06'W) (Ovruski & Schliserman 2003b), Tucuman (26[degrees]05'-28[degrees]01'S and 64[degrees]28'-66[degrees]13'W) (Nasca 1973; Ferndndez de Araoz & Nasca 1984; Ovruski 1995; Schliserman et al. 2004; Ovruski et al. 2004), and Salta (22[degrees]02'-26[degrees]21'S and 62[degrees]21'-68[degrees]34'W) (Ovruski et al. 2005; Orono et al. 2005). Pupal parasitoids were not found because all of the A. fraterculus and C. capitata were collected as larvae, so they would not have been obtained even if present at the collection site.
Rhoptromeris haywardi was not found during this study. However, as discussed by Wharton et al. (1998), published records for R. haywardi attacking tephritids (Blanchard 1947; Turica & Mallo 1961; Nasca et al. 1980) are questionable and need verification. All records of this eucoiline species come from bulk samples of fruit, from which parasitoid species of both Drosophilidae and Tephritidae could emerge. Without isolation of tephritid puparia, correct host fly--parasitoid species association cannot be made (Wharton et al. 1998).
Neither of the 2 introduced and released exotic fruit fly parasitoid species were recovered from fruit samples collected during this study, even though both D. longicaudata and A. indica were reported by Turica (1968) as established on Anastrepha spp. As noted by Ovruski et al. (1999), both exotic parasitoid species were recovered immediately following release in Concordia in the 1960s, and their establishment was not verified later. Interestingly, D. longicaudata was recently recovered in the northeastern province of Misiones (Schliserman et al. 2003) and in the northwestern province of Salta (Orono & Ovruski 2007) approximately 40 years after its first release. Similarly, recent fruit fly parasitoid surveys made in the provinces of Jujuy (northwestern Argentina), Cordoba cor·do·ba
See Table at currency.
[American Spanish córdoba, after Francisco Fernández de Córdoba (1475?-1526?), Spanish explorer.]
Noun 1. (Central Argentina), and Misiones (northeastern Argentina) recorded the presence of A. indica (Ovruski et al. 2006).
Although no C. capitata parasitoids were recovered in this study, some species, like A. pelleranoi, were obtained from C. capitata pupae in Tucuman. For example, in a recent fruit fly parasitoid survey in Citrus crop areas of Tucuman, A. pelleranoi has been recovered from C. capitata pupae in Citrus aurantium L. (Rutaceae), an exotic fruit mainly infested by Medfly (Schliserman & Ovruski 2004). According to according to
1. As stated or indicated by; on the authority of: according to historians.
2. In keeping with: according to instructions.
3. Ovruski et al. (2004), C. capitata is not heavily parasitized by neotropical Anastrepha parasitoid species. With the exception of the figitids A. pelleranoi, A. nordlanderi Wharton, and Odontosema anastrephae Borgmeier (Wharton et al. 1998), braconid parasitoids appear to adapt poorly to the exotic C. capitata (Ovruski et al. 2004).
Even though the number and size of fruit samples collected during this study were relatively small, the results seemingly suggest that U. anastrephae and D. brasiliensis would have a certain degree of fruit preference when searching host larvae. For example, U. anastrephae with the shortest ovipositor of any of the opiine species obtained (0.7 [+ or -] 0.1 times as long as the metasoma, n = 12), was most commonly found attacking A. fraterculus larvae in M. pungens and E. uniflora, the smallest fruit species sampled (7 and 8 times smaller than the guava or the feijoa, respectively). This relationship between smaller host fruit species and U. anastrephae has been previously reported by Hernandez-Ortiz et al. (1994), Lopez et al. (1999), Sivinski et al. (1997, 2000, 2001), and Aluja et al. (2003b) collecting fruit species of Spondias (Anacardiaceae) in Mexico, and by Aguiar Menezes et al. (2001) collecting Eugenia species (Myrtaceae) in Brazil. On the other hand, D. brasiliensis, which has the longest ovipositor of the parasitoid species collected (3.3 [+ or -] 0.2 times longer than the metasoma, n = 12), was observed only in association with A. fraterculus in F. sellowiana and P. guajava, the largest fruit species sampled. This peculiarity has also been noted by Aguiar-Menezes & Menezes (1997) in Brazil, when collecting A. fraterculus larvae parasitized by D. brasiliensis exclusively from Prunus persica Prunus persica,
n See peach. (L.) Batsch (peach), and by Ovruski et al. (2004) in NW Argentina, where D. brasiliensis was mainly recovered from both peach and guava. Interestingly, wild fruits of Prunus persica, P. guajaua, and F. sellowiana have a similar size (Ovruski et al 2004; Ovruski & Schliserman 2003a). Doryctobracon areolatus, which has a relatively long ovipositor (2.3 [+ or -] 0.2 times longer than the metasoma, n = 12) but 1.4 times shorter than D. brasiliensis' ovipositor, may be occupying an intermediate position between U. anastrephae and D. brasiliensis in host fruit preference. As reported previously Sivinski et al. (1997), Lopez et al. (1999), and Aguiar Menezes et al. (2001), D. areolatus might not have any host fruit preference. This opiine species has the broadest host-fly and host-fruit range of any of the Mexican (Sivinski 1997, 2000, 2001; Aluja et al. 2003b) and Brazilian (Canal & Zucchi 2000) fruit fly parasitoid species. In NW Argentina, D. areolatus has been reported foraging on a wide range of A. fraterculus host plant species (Ovruski et al. 2004). Apparently, a short ovipositor would allow U. anastrephae to have access to fly larvae mainly in small fruit, whereas a long ovipositor would allow D. areolatus (or D. brasiliensis) to reach host larvae in both large and small fruit. While studying a Mexican Anastrepha-parasitoid guild of 5 opiine species, including U. anastrephae and D. areolatus, Lopez et al. (1999) and Sivinski et al. (1997, 2001) found a strong correlation between ovipositor length of parasitoids and the size of host-in-fested fruits. This relationship would imply that the ovipositor length is an important limitation on foraging (Sivinski et al. 1997). Furthermore, Sivinski et al. (2001) and Sivinski & Aluja (2003) pointed out that the ovipositors in the opiine species attacking Mexican fruit fly would have originally diverged due to the action of environmental factors such as temperature, humidity, seasonality, and/or host-fruit abundance and diversity.
Although data provided here are preliminary, parasitization rates appear to be too low to consider parasitoids as a significant natural mortality factor of A. fraterculus in the study area. However, reliable data on the impact of the parasitoids upon natural A. fraterculus populations can be obtained only by closely monitoring the host and parasitoid population fluctuations for many generations in exotic and native host plant species. Thus, further studies on the ecology and behavior of A. fraterculus and their natural enemies should be carried out in Entre Rios for implementing a feasible, but cautious, biological control strategy against this fruit fly pest. As suggested by Aluja (1996, 1999), native host plants could be managed to naturally augment parasitoid numbers and to sustain parasitoid populations in wild vegetation areas. Therefore, conservation of native A. fraterculus-parasitoids would be an attractive alternative to the indiscriminate introduction of exotic parasitoids (Sivinski & Aluja 2003).
We express our gratefulness to Alejandra Roldan for the identification of all plant samples and to Eduardo Frias, Carolina Colin, and Jose Saez for technical support. Special thanks to Teresa Vera, Fanny Manso, Diego Segura, Jorge Cladera, Ricardo Tomassi, Natalia Petit, Norma Vaccaro, and Graciela Putruelle for sending additional samples of parasitoid collected in Entre Rios. We thank Martin Aluja (INECOL, A.C., Xalapa, Veracruz, Mexico) for sharing with us his vast experience on ecology, biology, and ethology ethology, study of animal behavior based on the systematic observation, recording, and analysis of how animals function, with special attention to physiological, ecological, and evolutionary aspects. of fruit fly parasitoids. Financial support was provided by the Consejo Nacional de Investigaciones Cientificas y Tecnicas de la Republica Argentina (CONICET CONICET Consejo Nacional de Investigaciones Científicas Y Técnicas (National Council for Science and Technology, Argentina) ) (Grant PIP Nos. 4973/ 97, 0702/98, and 5129/05).
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SERGIO M. OVRUSKI, PABLO SCHLISERMAN, LUIS E. ORONO, SEGUNDO R. NUNEZ-CAMPERO,
PATRICIA PATRICIA Practical Algorithm To Retrieve Information Coded In Alphanumeric
PATRICIA Proving and Testability for Reliability Improvement of Complex Integrated Architectures
PATRICIA PApilloma TRIal Cervical cancer In young Adults ALBORNOZ-MEDINA, LAURA Laura, subject of the love poems of Petrarch. She is thought to be Laura de Noves (1308?–1348), wife of Hugo de Sade, but this has not been proved.
Petrarch’s perpetual, unattainable love. [Ital. Lit. P. BEZDJIAN AND GUIDO GUIDO Guidance Officer A. VAN NIEUWENHOVE
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TABLE 1. ABUNDANCE OF A. fraterculus PARASITOIDS COLLECTED FROM 4 MYRTACEAE SPECIES IN ENTRE TRIOS, NW ARGENTINA, BETWEEN JAN AND MAR 1993 AND 1994. Collection sites Host plant No. of No. and weight (kg) and years species samples of fruit sampled Concordia (1993) P. guajava 14 143 5.0 Concordia (1994) P. guajava 6 62 2.7 La Paz (1993) F. sellowiana 5 50 1.3 P. guajava 4 41 1.8 La Paz (1994) F. sellowiana 5 57 1.7 P. guajava 22 225 8.3 ParanA (1993) E. uniflora 6 211 1.6 M. pungens 3 104 1.1 P. guajava 8 83 3.9 ParanA (1994) E. uniflora 6 209 1.5 M. pungens 6 206 2.0 P. guajava 6 61 2.1 No. of recovered Collection sites Host plant A. fraterculus and years species pupae and adults Concordia (1993) P. guajava 277 170 Concordia (1994) P. guajava 207 191 La Paz (1993) F. sellowiana 111 69 P. guajava 109 71 La Paz (1994) F. sellowiana 204 164 P. guajava 559 327 ParanA (1993) E. uniflora 79 64 M. pungens 46 28 P. guajava 231 149 ParanA (1994) E. uniflora 77 51 M. pungens 81 39 P. guajava 94 55 No. of recovered parasitoid Collection sites Host plant adults (1) and years species D.a D.b Concordia (1993) P. guajava 2 1 Concordia (1994) P. guajava 17 5 La Paz (1993) F. sellowiana 6 5 P. guajava 5 3 La Paz (1994) F. sellowiana 17 0 P. guajava 12 2 ParanA (1993) E. uniflora 2 6 M. pungens 1 0 P. guajava 12 0 ParanA (1994) E. uniflora 3 6 M. pungens 1 0 P. guajava 3 0 No. of recovered parasitoid Collection sites Host plant adults (1) and years species U.a A.p Concordia (1993) P. guajava 0 8 Concordia (1994) P. guajava 0 7 La Paz (1993) F. sellowiana 0 1 P. guajava 0 11 La Paz (1994) F. sellowiana 1 5 P. guajava 4 13 ParanA (1993) E. uniflora 8 1 M. pungens 3 0 P. guajava 1 7 ParanA (1994) E. uniflora 4 0 M. pungens 4 1 P. guajava 0 2 Mean % Collection sites Host plant parasitism and years species ([+ or -] SD) Concordia (1993) P. guajava 11.4 [+ or -] 3.2 Concordia (1994) P. guajava 13.2 [+ or -] 4.7 La Paz (1993) F. sellowiana 12.7 [+ or -] 2.3 P. guajava 18.9 [+ or -] 3.1 La Paz (1994) F. sellowiana 12.7 [+ or -] 2.7 P. guajava 9.8 [+ or -] 6.3 ParanA (1993) E. uniflora 11.2 [+ or -] 6.9 M. pungens 4.9 [+ or -] 4.3 P. guajava 14.8 [+ or -] 7.4 ParanA (1994) E. uniflora 8.7 [+ or -] 9.2 M. pungens 5.8 [+ or -] 8.1 P. guajava 9.7 [+ or -] 3.3 (1) D.a, D. areolatus; D.b, D. brasiliensis; U.a., U. anastrephae; A.p, A. pelleranoi.