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Spatio-Temporal Population Sampling of a Fire Ant Parasitoid Donald C

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Spatio-Temporal Population Sampling of a Fire Ant Parasitoid Donald C DOI: 10.1111/j.1570-7458.2008.00763.x Blackwell Publishing Ltd Spatio-temporal population sampling of a fire ant parasitoid Donald C. Henne*, William S. Hilbun & Seth J. Johnson Department of Entomology, Louisiana State University Agricultural Center, 404 Life Sciences Building, Baton Rouge, LA 70803, USA Accepted: 18 June 2008 Key words: Pseudacteon tricuspis, Solenopsis invicta, Thelohania solenopsae, SADIE, red imported fire ant, decapitating fly, Phoridae, Diptera, Hymenoptera, Formicidae, Microsporidia, Thelohaniidae Abstract The spatial and temporal population structure of Pseudacteon tricuspis Borgmeier (Diptera: Phori- dae), an introduced parasitoid of the invasive red imported fire ant, Solenopsis invicta Buren (Hymenoptera: Formicidae), was investigated at three locations in Louisiana, USA, during the fall of 2005. Georeferenced disturbed fire ant mounds that attracted adult P. tricuspis were analyzed using Spatial Analysis by Distance IndicEs to model the population structure of this parasitoid on a local spatial scale, and relate spatial abundance to host colony social form and infection with the fire ant pathogen, Thelohania solenopsae Knell, Allen and Hazard (Microsporidia: Thelohaniidae). Spatial associations between P. tricuspis counts and host S. invicta colonies infected with T. solenopsae were not detected. Overall, spatio-temporal count patterns of P. tricuspis were largely random. However, high P. tricuspis counts were significantly associated with areas of polygyne host colonies. Occasional aggregations of P. tricuspis populations into discrete spatial patches and gaps were also found. The implications of this study are that population surveys of P. tricuspis should be conducted in several areas at a location to account for local patchiness in fly populations and/or varying densities and social form of host ant colonies. Analyzing counts of organisms that are based on tradi- Introduction tional methods of sample variance–mean relationships It is a trivial fact that populations of organisms occupy and derivatives thereof (see Taylor, 1984) only provide certain areas at certain times, and some species are more information on the numeric properties of the underlying mobile than others. Understanding the processes that frequency distribution, and therefore have limited affect the spatial distribution of plant and animal popu- capability to describe the essential spatial information of lations is an important focus in ecology (Tuda, 2007). counts (Perry & Hewitt, 1991; Perry et al., 1999). For example, For mobile animal species, spatial information is often a highly skewed series of counts (e.g., negative binomial) restricted to trap counts at specific locations (Perry, 1995). with one or more very large values relative to the others These count locations may be spatially arranged in regular, may nevertheless be completely random in space (Perry & random, or clustered patterns without regard to the properties Dixon, 2002). Current analyses of insect spatial and tem- of the underlying count frequency distribution (Ferguson poral distributions utilize location information of sample et al., 2000). Given that population sampling relies on units to detect and measure degree of non-randomness spatial dispersion patterns (Southwood, 1978), determining in the spatial pattern in two-dimensional space (Perry, 1998a) the spatial and temporal population structure of organisms and spatial association between data sets collected on is an important objective toward understanding their different occasions (Winder et al., 2001). This methodology population dynamics. However, the difficulty of directly has been applied to the study of several host–parasitoid studying movement of individual animals, particularly systems (e.g., Ferguson et al., 2000, 2006; Weaver et al., small and numerous species such as insects, presents 2005). Therefore, key insights into spatial dynamics of special problems for ecologists (Perry, 1998a). host–parasitoid systems, such as spatial aggregations of parasitoids and their hosts, may be achieved by incorpo- *Correspondence: E-mail: [email protected] rating spatial information of count data into analyses. © 2008 The Authors Entomologia Experimentalis et Applicata 129: 132–141, 2008 132 Journal compilation © 2008 The Netherlands Entomological Society Fire ant parasitoid dynamics 133 The red imported fire ant, Solenopsis invicta Buren P. tricuspis to host social form and presence/absence of (Hymenoptera: Formicidae), is an exotic invasive insect in T. solenopsae. the USA and is regarded as a significant economic pest (Lofgren, 1986; Porter et al., 1992). Management of Materials and methods S. invicta in the USA includes biological control through importation of natural enemies from the indigenous range Study locations of S. invicta in South America, particularly parasitic flies The P. tricuspis populations sampled in this study of the genus Pseudacteon Coquillet (Diptera: Phoridae). originated from a release conducted 17 km northeast of These flies are solitary endoparasitoids of Solenopsis fire Covington, St. Tammany Parish, LA, USA (30°36′N, ants (Morrison & Porter, 2005) and adults of one intro- 90°01′W) from 8–13 September 1999 (2 165 flies released) duced Pseudacteon species, Pseudacteon tricuspis Borgmeier, (see Henne et al., 2007). Populations of P. tricuspis were are attracted to alarm pheromones emitted by S. invicta sampled weekly at three widely separated locations in (Morrison & King, 2004). Pseudacteon females insert a Washington Parish, LA, USA, between 21 September and single egg into the host ants’ thorax, and the maggot 19 October 2005. The GPS coordinates for each study migrates to the hosts’ head capsule, consumes the contents location were recorded with a MagellanTM GPS 315/320 of the head over a 2-week period, and eventually decapitates (Magellan, Santa Clara, CA, USA). The first study location the host (Porter, 1998). was 8 km south of Bogalusa (30°41′N, 89°53′W; hereafter Phorid fly population dynamics are not well-understood called Farm 2), and was approximately 70 × 70 m in an (Morrison, 2000). In particular, almost no information is unmaintained cattle pasture. The second study location available concerning the spatial and temporal dynamics of was 14 km southwest of Franklinton (30°48′N, 90°18′W; various Phoridae (Disney, 1994). Populations of Pseu- hereafter called Farm 4), and was approximately 40 × 60 m dacteon parasitoids of the tropical fire ant, Solenopsis geminata in a mowed cattle pasture. The third study location was (Fabricius), in central Texas were characterized as having 16 km west of Bogalusa (30°46′N, 90°01′W; hereafter significant variations in abundance, both spatially and called Farm 7), and was approximately 170 × 100 m in an temporally (Morrison et al., 1999). Morrison & King unmaintained cattle pasture. (2004) evaluated phorid abundance at disturbed S. invicta At each study site, all active S. invicta colonies (usually mounds at multiple sites in North Central Florida over 15–25) were located in an arbitrarily defined area and time and determined that abundance deviated signifi- permanently marked with a barcode to enable future cantly from a regular distribution. However, neither of identification. The barcode consisted of a high durability these studies evaluated phorid abundance at the same weather-resistant polyester barcode label adhered to a individual mounds over time nor modeled abundance in 10 × 5 cm piece of rigid plastic sheeting that was anchored a spatio-temporal context. Additionally, no studies have flush with the ground with 10-cm long wire staple. The evaluated potential spatial patterns of phorid flies in barcode label assigned a unique number to each S. invicta relation to host colony social forms (monogyne vs. polygyne) mound that was retained throughout the study phase. and/or infection with microsporidian disease. Two social A Symbol® MC 3000 batch mobile computer (Motorola, forms of S. invicta, monogyne (single-queen) and polygyne Holtsville, NY, USA) was used to scan and record each (multiple-queens) occur in the USA (Glancey et al., 1973). barcode’s unique value into a database. The two forms are regulated by a single gene that is Decimal-degree locations of S. invicta mounds were homozygous (Gp-9B) in monogyne queens and hetero- taken with a GPS to obtain x, y-coordinates of sampling zygous (Gp-9B and Gp-9b) in polygyne queens (Ross & locations. Generally, at least 10 m separated S. invicta Keller, 1998; Krieger & Ross, 2002). mounds. At each location, weekly mean soil moisture The microsporidian entomopathogen, Thelohania levels were obtained using a Lincoln soil moisture meter solenopsae Knell, Allen and Hazard (Microsporidia: (Forestry Suppliers Part No. 3052; Lincoln Irrigation, Thelohaniidae), has been isolated from polygynous Lincoln, NE, USA), driven into the soil to a depth of 10 cm. S. invicta colonies in both North and South America, Ten measurements at 5 m intervals were made along a causing significant mortality in infected colonies (Oi & transect through each study area. Williams, 2002). However, it is unknown if S. invicta social form or infection with T. solenopsae influence P. tricuspis Sampling Pseudacteon tricuspis spatial distributions. Therefore, the objectives of this study Sampling Pseudacteon parasitoids of Solenopsis involved were to (1) characterize the number of attracted P. tricuspis simply disturbing host nests and awaiting arrival of flies. to disturbed S. invicta mounds in a spatio-temporal way Two observers disturbed
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