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Bionomics of Orasema Simplex (Hymenoptera: Eucharitidae), a Parasitoid of Solenopsis fire Ants (Hymenoptera: Formicidae) in Argentina

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Bionomics of Orasema Simplex (Hymenoptera: Eucharitidae), a Parasitoid of Solenopsis fire Ants (Hymenoptera: Formicidae) in Argentina Biological Control 48 (2009) 204–209 Contents lists available at ScienceDirect Biological Control journal homepage: www.elsevier.com/locate/ybcon Bionomics of Orasema simplex (Hymenoptera: Eucharitidae), a parasitoid of Solenopsis fire ants (Hymenoptera: Formicidae) in Argentina L. Varone *, J. Briano USDA-ARS South American Biological Control Laboratory, Bolivar 1559 (1686) Hurlingham, Buenos Aires Province, Argentina article info abstract Article history: Biological characteristics of the parasitoid Orasema simplex Heraty (Hymenoptera: Eucharitidae), a poten- Received 17 June 2008 tial candidate for the biological control of fire ants in the United States were investigated. Female survi- Accepted 9 October 2008 vorship, fertility and oviposition preferences were studied in the laboratory. Naturally parasitized Available online 17 October 2008 colonies were examined to determine offspring sex ratio, development success and time, and to artifi- cially parasitize healthy ant colonies. In addition, field studies were carried out to establish natural ovi- Keywords: position substrates and adult activity patterns. Orasema simplex female survivorship was 3.6 ± 1.5 days. Orasema simplex Newly emerged females contained 613.5 ± 114.0 mature eggs. The adult development success in natural Solenopsis richteri parasitized colonies was 22.2% with a female-biased sex ratio (4:1). The time required from planidia to Solenopsis invicta Fire ants adult was 29.5 ± 5.4 days. In the field, adults were mostly found around the ant nests at midday. A broad Biological control range of plant species was observed as oviposition substrates. The transfer of planidia to healthy ant col- onies was achieved but the development success was very low. Orasema simplex appears to have a limited potential as a fire ant biocontrol agent because of cosmetic damage to a wide variety of plants used for oviposition. However, further studies are necessary to evaluate the real damage exerted by oviposition punctures. Published by Elsevier Inc. 1. Introduction Initially, the planidia burrows under the host cuticle and swells slightly indicating limited feeding by the parasitoid. Upon pupa- The species of Eucharitidae (Hymenoptera) are parasitoid wasps tion, first instar larva becomes external (Heraty, 1994a, 2000). of ants (Das, 1963; Johnson et al., 1986; Williams and Whitcomb, The parasitoid completes development when the host enters the 1974). Almost all species of Orasema are parasitoids of ants in pupal stage (Johnson, 1988; Heraty et al., 1993). Orasema imma- the genera Solenopsis, Pheidole, Tetramorium and Wasmannia tures are treated by host workers as ant brood, since they acquire (Heraty, 1994b). The first studies on the biology and habits of the host colony odors that allows acceptance inside the colony (Vander Eucharitidae were carried out by Wheeler (1907), who found sev- Meer et al., 1989). When development is complete, adults emerge eral species of Orasema associated with ants of the genera Pheidole within the ant nest (Heraty, 1985). and Solenopsis in Texas and Colorado. Eucharitids have an unusual In Argentina and Brazil, unidentified Orasema spp. have been re- life cycle. Females lay their eggs in plant tissue near ant nests ported ovipositing in several plants (Parker, 1942; Tocchetto, (Wheeler, 1907; Johnson et al., 1986). The site of oviposition and 1942). They were first reported parasitizing fire ants of the Solenop- the range of plants used by the different eucharitid species are sis saevissima complex in 1964 in Uruguay (Silveira-Guido et al., uncertain. Therefore, the presumed specificity for oviposition 1964) and later in Brazil (Williams and Whitcomb, 1974). The im- plants may be a consequence of the scarce data available (Johnson, ported fire ants, Solenopsis invicta Buren and S. richteri Forel are two 1988). In general, the part of the plant the eucharitids utilized for species of this complex that were accidentally introduced in the oviposition is highly variable, including overwintering buds, open- United States from South America and became serious medical ing flower buds, stems of blossom clusters, seed pods and leaves and economic pests (Adams, 1986; Logfren 1986a,b; Drees et al., (Clausen, 1940). The emerged planidia attach themselves to a for- 1992; Lard et al., 2001; Pereira et al., 2002). Since then, several nat- aging worker ant or intermediate host and are carried to the nest ural enemies have been under study: two species of microsporidi- phoretically. Once there, they are transferred to an ant larva. an pathogens, several species of phorid flies, a congeneric parasitic ant and more recently the mermithid nematode Allomermis sole- nopsii (Briano et al., 1995a,b, 1997, 2002a; Calcaterra et al., 1999; * Corresponding author. Fax: +54 11 4452 1882x104. Oi et al., 2005; Orr et al., 1995; Pesquero et al., 1995; Porter, E-mail address: [email protected] (L. Varone). 1998, 2000; Williams et al., 1999; Porter and Varone unpublished). 1049-9644/$ - see front matter Published by Elsevier Inc. doi:10.1016/j.biocontrol.2008.10.003 L. Varone, J. Briano / Biological Control 48 (2009) 204–209 205 The flies Pseudacteon tricuspis Borgmeier, Pseudacteon curvatus 2.2. Field observations Borgmeier, Pseudacteon litoralis Borgmeier, and Pseudacteon obtusus Borgmeier have been released (Porter et al., 2004; Vazquez et al., 2.2.1. Parasitism rate 2006; Graham et al., 2003; Gilbert et al., 2008) and the microspo- Individuals of Orasema simplex were obtained from parasit- ridian Vairimorpha invictae Jouvenaz and Ellis is still in quarantine. ized colonies collected from February 2005 to May 2007 at the Orasema was also listed as a potential candidate for the biological 24 positive sites found in Argentina (Varone and Heraty, unpub- control of Solenopsis and investigations on this organisms started at lished). The colonies were excavated, put in 10 L dusted buckets the South American Biological Control Laboratory (SABCL) in 2005 and transported to the laboratory for later flotation and separa- as part of the biological control program against fire ants. tion from the soil (Banks et al., 1981). The ant brood was sepa- In our recent surveys in Argentina, O. simplex Heraty was the rated from the colony using sorting sheets (Banks et al., 1981) eucharitid species most commonly found parasitizing fire ants, and then observed under the dissecting scope for the presence with a wide distribution but with low abundance and persistence of parasitoids. in the field (Varone and Heraty, unpublished). The objective of this Once detected, the Orasema larvae and pupae were returned to study was to determine the bionomics of O. simplex in the labora- their original host colonies that were previously reduced to 100– tory and to conduct field trials to detect natural oviposition sub- 200 workers and a small amount of Orasema-free brood. These strates and adult activity patterns. Such information is essential small fragments of colonies were kept in rearing chambers at to assess the real potential of O. simplex as a biological control 30 ± 2 °C in plastic vented containers with food and humidity agent of imported fire ants in the United States. sources, and checked daily for the emergence of the adult parasit- oids. The development time from planidia to adult was estimated 2. Materials and methods and the sex ratio of emerged adults recorded. The CBS was selected to conduct ecological studies because of 2.1. Laboratory studies the high occurrence and persistence of O. simplex (Varone and Heraty, unpublished). The site chosen was the entrance of the sta- 2.1.1. Survivorship and fertility tion, a 250 m long–20 m wide dirt road surrounded by a xerophytic Orasema simplex female survivorship was estimated with 30 forest. newly emerged females (obtained from field collected colonies) ex- posed to different plant species as oviposition substrates. The spe- 2.2.2. Adult activity cies tested were: (1) Eupatorium aff. laevigatum L. (Asteraceae), (2) Adult presence was monitored observing 36 fire ant mounds in Vinca rosae L. sp. (Apocynaceae), (3) Impatiens sp. (Balsaminaceae), the morning (8–10 h; 19.2 ± 1.7 °C), at noon (11–13 h; 24.8 ± and (4) Viola sp. (Violaceae). No plants were available in the control 0.2 °C) and in the afternoon (15–17 h; 27.6 ± 1.6 °C) for three treatment; 4–9 replicates were considered for each treatment. consecutive days. Each unmated female was placed in a plastic vented bottle at- Each mound and the surrounding area (1 m diameter) were ob- tached to a stem of the potted test plant or, in the controls, placed served during 2–3 min; individuals found were captured with an in a plastic container with humid tissue paper. All females fed hon- aspirator, marked with latex paint and released. The volume of ey–water solution and were checked daily. Potted plants were each mound was estimated as a hemisphere with the formula 3 placed in a rearing chamber at 25 ± 2 °C and 14–10 h L–D. Fertility V = (2/3)pR (V, volume; R, radius) to evaluate a size preference was estimated by dissecting other 10 females immediately after of O. simplex. Sixteen of the 36 mounds were excavated and trans- emergence and by counting the mature oöcytes in the ovarioles. ported to the laboratory in dusted buckets for later brood examination. 2.1.2. Transfer of planidia Several tests with different approaches were conducted to arti- 2.2.3. Oviposition substrates ficially transfer newly emerged planidia from the Corrientes Bio- Natural oviposition substrates were recorded in three different logical Station (CBS, S 27° 33,1730;W58° 40,7710) to receptor habitats with high occurrence of O. simplex: (1) the entrance of the laboratory S. invicta colonies that were free of the parasitoid. The CBS, (2) a pasture in Colonia Hughes, Entre Ríos (S 32° 22,7850;W approaches were: (1) plants with oviposition marks and/or plani- 58° 16,6780) and (3) abandoned land in Concepción del Uruguay, dia were potted in the field, transported to the laboratory and ex- Entre Ríos (S 32° 27,7610;W58° 14,1960).
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