Aris: diversity as affected by J. D. P. Aris flower species 3965 NW 27th Lane Gainesville, FL 32606 Journal of Economic Phone: 352 359-1846 Email: [email protected]

Aris: A Comparison of Native Wasp Diversity and

Abundance as Affected by Different Species of

Native Annual and Perennial Flowers

J. D. P. Aris

University of Florida, Department of Entomology and Nematology

970 Natural Area Drive, Gainesville, FL 32611 1 Abstract 2 3 pollination services are essential to the world’s agricultural production.

4 Until recently, most of the world and the United States relied on the services of the

5 managed European (Apis mellifera) for these pollination services. However, A.

6 mellifera has recently experienced a dramatic population decline, prompting research into

7 the use of alternative to achieve adequate crop pollination. One such option is

8 to use native pollinators such as native bees and . In this project, I established two

9 sites wherein I planted 5 plots of different assortments of easily grown, native Florida

10 wildflower species, as well as a control plot with no flowers planted. Flower plots were

11 grouped by their flower type (annual or perennial) and the number of species included in

12 the mixture (basic = 5-6 species and diverse = 9-11 species), with one plot being a

13 combination of the two flower types. Wasps were collected using an insect net from these

14 plots once every 3 weeks over a 12 week period between May and July of 2010. The

15 wasps were identified to family. The plot on which they were caught was recorded.

16 Statistical software was used to analyze the data for significant differences in wasp

17 abundance (number of individuals caught/minute) and diversity (number of families

18 caught/minute). The data indicated that flower plot affected wasp abundance, but not

19 wasp diversity. Additionally, certain wasp families were more prevalent than others.

20 21 22 Key Words: Pollination biology, wasps, native pollinators 23 24

25 26 Pollination services provided by are very important to human beings.

27 Thirty-five percent of the world’s crop production relies on insect pollination, as well as

28 60-80% of wild plant species. An even larger number than this benefit from animal

29 pollinators (Kremen et. al. 2007). At the moment, the U.S. relies mostly on the services

30 of managed honey bees (Apis mellifera) to achieve adequate crop pollintation. About

31 90% of commercially grown field crops, vegetables, and nuts grown in the U.S. rely on

32 these honey bees (Goldman 2010). Soon after 2000, honey bee populations started

33 declining due to a variety of factors. Much of the decline can be attributed to Varroa

34 destructor, a disease-carrying species of mite (Kremen et. al. 2007). Colony Collapse

35 Disorder (CCD) has also been responsible for large declines in honey bee populations

36 (Goldman 2010).

37 Such declines in honey bee populations are financially threatening since honey

38 bees are the most valuable worldwide (Goldman 2010). A possible remedy to

39 this problem is to investigate the use of native pollinators to pollinate U.S. crops. Native

40 pollinators are already responsible for about $3.07 billion of the fruits and vegetables the

41 United States produces (Losey & Vaughan 2006). If the U.S. could make use of these

42 pollinators, it would no longer have to rely on honey bees imported from elsewhere. In

43 addition to reducing our reliance on the threatened population of honey bees, this may

44 also reduce costs for farmers (Goldman 2010). Wild native pollinators can be just as

45 efficient as managed pollinators (Rickets et. al. 2004).

46 Most native pollinators are bees (Losey and Vaughan 2006) which do not fly

47 great distances (they generally forage between 100 and 200 meters from their hive, and

48 only very rarely go over 400 meters away), and rely on native habitats (Goldman 2010; 49 Henry et. al. 2012). This suggests that one must set aside some patches of land to be

50 maintained as native habitat in order to augment native bee populations, or at least

51 provide nearby structures within which native pollinators can nest (Gruber et. al. 2011).

52 By preserving or even increasing the abundance of native plants, areas could augment the

53 population of native pollinators and attract them to places where they would be

54 agriculturally useful. This process is called pollinator spillover, and there is some

55 evidence that this occurs, although research has focused on pollinators spilling over from

56 crops to native plants. (Hanley et. al. 2011).

57 While it has been shown that native bees are often the most active pollinators of

58 the native pollinators (Losey and Vaughan 2006), and a lot of research has focused on

59 them (Tonietto et. al. 2011, Jha & Dick 2010, Dohzono & Yokoyama 2010), other insects

60 are important as well. Other Hymenopterans, as well as Lepidopterans and a variety of

61 Dipterans and Coleopterans can act as pollinators. Among the hymenopterans, wasps

62 have also been observed as pollinators (Shuttleworth and Johnson 2010, Peakall et. al.

63 2010). What is not known is if native wasp populations can be augmented in the same

64 way as those of native bees, i.e. by providing floral resources to which they or their prey

65 are attracted. The purpose of this project is to investigate the diversity and abundance of

66 native wasps as affected by the availability of native flowering plants. The data from this

67 project also will allow one to identify patterns and trends in wasp behavior based on the

68 temporal availability of various wildflower species.

69 70 71 72 73 74 75 Materials and Methods 76 77 Preparation: Planting and Maintenance 78 79 Two acceptable sites were found (named 1 and 2) for planting flower strips. These

80 sites were 22 × 75 m (l × w). Six, 3 × 15 m wildflower plots were planted at each site.

81 Ten meters of mowed grass separated each plot (Figures 1 & 2). The plots at both sites

82 were treated with glyphosate (TouchDownTM) at 2.84 liters per acre and a nonionic

83 surfactant at 0.95 liters per acre. The grass at these sites was mowed with Hiniker 5700

84 Flail-chopper, and the thatch was harvested.

85 Seven weeks after the initial herbicide treatment, all of the plots were retreated as

86 before. All of the plants not killed by the herbicide were removed from the plots. The

87 remaining thatch was burned from the plots. Mats of dead Bahia grass roots remained in

88 some of the plots. These were broken up with a Lilly Rod Weeder. Each plot was then

89 rolled with a 16 ft. roller, raked a second time, and then rolled a second time. Once the

90 plots were prepared, they were planted using a Scotts Professional Drop spreader and

91 rolled a final time to push the seeds just into the surface of the ground. The 6 plots were

92 established per the seed mixes reported in Table 1.

93 An irrigation schedule was established such that the plots were irrigated every

94 week that rainfall did not reach a 0.635 cm. The plants were given 1.27 to 2.54 cm of

95 water during each irrigation. All undesirable weeds were pulled.

96 The six plots planted at each site were a composition of different flowers and a

97 control, in which nothing was planted but native vegetation was allowed to grow (Table

98 1). The plants listed in the table were selected because they are native to Florida and 99 easily grown and maintained. These are ideal characteristics if the plants are to be used

100 by farmers to attract native pollinators.

101 Collection and Identification 102 103 Each flower patch was divided into two half sections longitudinally before data

104 collection. Both halves were sampled for an equal amount of time. I sampled each half

105 using an insect net, trying to catch every wasp seen. I also randomly followed a sweep-

106 net protocol by swinging my net through the flower vegetation to catch cryptic wasps.

107 Wasps were collected once every 3 weeks for 12 weeks between the months of May and

108 July of 2010.

109 Wasps caught in the net were killed in the field using a cyanide-based kill jar.

110 Each wasp was placed into a test tube labeled with the date, site and plot on which it was

111 captured. The euthanized wasps were pinned and arranged in insect storage boxes

112 according to where and when they had been captured. Wasps were identified to family

113 and labeled appropriately. The sites were sampled for one day each on consecutive days.

114 The combined catches for the two sites were considered one collection.

115 116 Analysis 117 118 The number of individual wasps that were caught per minute was analyzed by

119 treatment (flower strip and collection number) with a one way ANOVA. The number of

120 families that were caught per minute was also analyzed by treatment (flower strip and

121 collection number) with a one way ANOVA. Furthermore, I used a two way ANOVA to

122 test the effects of plot, wasp family, and the interaction of treatment plot × family on the

123 number of wasps that were caught. Where necessary, I used Student’s T-tests to compare

124 the square roots of the means for treatments with less than 6 replicates, accepting 125 differences at P<0.05. I used Tukey tests to compare means for treatments with 6 or more

126 replicates.

127 128 129 130 Results 131 132 Flower strip did not affect the diversity of wasp families present though it did

133 affect wasp abundance (Table 2). The Annual Diverse plot had more wasps visiting the

134 sites than did all of the other plots except for the Perennial plot. A greater diversity of

135 wasp families (as measured by the number of families caught per minute) was present

136 during the third collection than the first or fourth, while the total number of wasps

137 collected was higher during the second collection than the first (Table 3). There were

138 significantly more individuals caught from certain wasp families (Table 4). More

139 individuals were caught from Crabronidae, Sphecidae and Scoliidae than from any other

140 family. Family Crabronidae was more populous than Sphecidae, and Sphecidae was more

141 common than Scoliidae.

142 143 Discussion 144 145 The data indicate that plot composition affected wasp abundance. This could be

146 explained by a variety of possible factors. Some of the wasps are attracted to flowers as

147 they search for nectar to supplement their diet (Kugimiya et. al. 2010). Others likely were

148 searching for prey or hosts (Fernandes et. al. 2010). Wasps visiting flowers to acquire

149 nectar may have preferred some flowers over others. It is not uncommon for wasps to

150 show preferences for some flowers (Wilmer 2011). Alternatively, flower diversity may

151 influence wasp abundance because wasp prey may prefer certain flowers. Wasps are 152 predators or parasites of various Lepidopterans and Dipterans, and even other

153 Hymenopterans (Woelke 2010, Hummel et. al. 2010, Rortais et. al. 2010). These species

154 are often present in flower patches in high densities (Bruinsma et. al. 2008). All of these

155 insect families have members that are known to have certain flower preferences and

156 specializations. Wasps seeking nectar likely had preferences, and wasps seeking a

157 host/prey item may have had “indirect” preferences because their host/prey item had

158 certain preferences.

159 While it is true that the main difference between annuals and perennials is their

160 life cycle, and not the flowers themselves, it may still be the case that different species

161 differ in attractiveness to different native insects (Albani and Coupland, 2010). Some of

162 the plant species in certain plots may be more attractive to more of the native insects. The

163 similarity in attractiveness shown by many of the plots may have been an artifact of a few

164 particularly attractive flowers being present in multiple plots, so investigation into the

165 attractiveness of individual plant species would be a worthy future enterprise. Knowing

166 which individual flowers were considered most attractive by wasps would be valuable

167 because these could be used to attract the greatest abundance of wasps, increasing the

168 chance of pollinator spillover.

169 This study does not conclude that wasps prefer a greater diversity of flowers over

170 a lesser diversity, although the data suggests that this may be the case. More extensive

171 research would be required to confirm this pattern. The density of flowers in each plot

172 was similar, so observed differences in wasp density in the various plots could indicate a

173 qualitative difference between the flowers, not a quantitative one. This indicates that the

174 presence/absence or abundance of certain individual plant species may have influenced 175 wasp preferences. This study revealed no data regarding which plant species may have

176 had such an influence.

177 The control plots grew a variety of native wildflowers that we did not plant, and

178 so presented their own separate array of flowers to attract insects that may be interested

179 in nectar. What one may conclude from these results and observations is that many plants

180 present in the control plots are as attractive to wasps as many of the plants included in the

181 other plots.

182 It is important to note that this project took place over a single year, so wasp

183 behavior may change in subsequent years based on the regrowth patterns of the flower

184 species. Furthermore, the data suggest there is no difference in the diversity of wasps

185 attracted to different flower types, though trends were present. As such, increasing the

186 sample size may have resulted in a clearer picture of wasp attraction to flowers.

187 The diversity of wasps found on each plot was measured by the number of

188 different families caught on each plot per minute sampled. This does not take into

189 account certain factors such as the number of individual wasps caught from each family.

190 So my diversity measure does not account for the possibility that there may have been

191 more individuals from some families than from others. Future investigation into the

192 diversity of wasps being found on these plots should include a diversity index that takes

193 the abundance of each family in comparison to the other families into account.

194 There are a number of possible reasons that certain wasp families were present in

195 greater numbers than were other families. For example, there could simply be more

196 individuals in some wasp families present in the native ecosystem than other wasp

197 families. It could also be the case that certain wasp families are attracted to flowers 198 and/or open fields in greater numbers than other wasp families. In this case, even a wasp

199 family that may be lower in abundance overall may have been more abundant on the

200 plots. Further research should be done on this topic. If certain wasp families are

201 differentially attracted to flowers and open fields, it would be to the advantage of farmers

202 to have this information as they could plant flowers to attract important predator species.

203 There were a high number of individuals from Crabronidae, Sphecidae, and

204 Scoliidae. Individuals from all three of these families have been observed behaving as

205 pollinators (Wilmer 2011, Patiny 2012). Sphecids (as will as Pompilids and Tiphids)

206 frequently gather nectar as fuel between longer, prey-gathering forage trips. While some

207 Sphecids, Pompilids, and Tiphids feed solely on insects, many frequently feed on liquid

208 flower products. Some even seem to require the sugary flower products at the beginning

209 of their adult life, while nest building, and hence are common on flowers during spring

210 time. They prefer open flowers with fully exposed nectar, but are also found on bowl-

211 shaped flowers with partially concealed nectar. A few species even visit tubular flowers

212 or composite inflorescences with multiple corolla tubes. Scoliids tend to be parasitic on

213 the larvae of other Hymenopterans, and often the females are wingless (making them

214 poor pollinators), but most of the winged forms visit shallow open flowers occasionally

215 to supplement their diet (Wilmer 2011). So all three of the most commonly found wasp

216 families on my plots are known pollinators, and hence are potentially valuable as native

217 pollinator alternatives to A. mellifera.

218 Future research on wasp diversity as affected by flower diversity should be

219 conducted beyond the family level. Many patterns may emerge if one investigates the

220 array of different genera or species present on the different flower plots. Such 221 investigations may also make it easier to investigate the relationships between the biology

222 of the wasps and their behavior in the field. In my experience, it was difficult to tell how

223 many wasps were present and acting as pollinators, and how many were there to hunt for

224 prey. If wasps are investigated at the genus or species level, it will be easier to research

225 the relative abundance of pollinators and parasites, and how their presence affects flower

226 pollination.

227 I believe that it would also be valuable to do research on individual flowers

228 instead of a multitude of flowers all at once. For example, the plots attracting wasps may

229 be more attractive because of only one or two of the flowers in the plot. If certain plant

230 species attract wasps in greater number, then it would be far more economical for farmers

231 to plant only these species.

232 In conclusion, it would be valuable to investigate the effect of the plots on wasp

233 abundance and diversity on nearby crops. It is possible that crops would benefit from

234 having more wasps in the area. Alternatively, the opposite may happen with wasps

235 moving from the crops to the flower strips instead. My research is a preliminary

236 investigation into the feasibility of trying to use native pollinators to pollinate crops by

237 causing pollinator spillover from nearby native vegetation. Future research will

238 hopefully reveal how this goal of using native pollinators to compensate for the declining

239 A. melifera population might optimally be accomplished.

240 My research found that both the flowers present and the time of year can affect

241 how many native wasps are present. These findings are valuable because they suggest

242 that we have the ability to manipulate the behavior of native wasps such that we might

243 use their pollination services on our crops as part of our effort to eliminate our 244 dependency on A. melifera. Additionally, we might try to time the flowering of our crops

245 to correspond the periods of highest native pollinator activity.

246 247 Acknowledgements 248 249 I thank Kevin Aris for his help with technical aspects of this project. Kaitlin Self

250 provided technical support and data entry/manipulation support.

251 252

253 References Cited 254 255 Albani, Maria C. and George Coupland. 2010. Comparative Analysis of Flowering in 256 Annual and Perennial Plants. Current Topics in Developmental Biology 91: 323-348. 257 Bruinsma, Maaike, H. Ijdema, J. J A. van Loon, M. Dicke. 2008. Differential effects 258 of jasmonic acid treatment of Brassica nigra on the attraction of pollinators, 259 parasitoids, and butterflies. Entomologia Experimentalis et Applicata 128(1): 109-116. 260 Dohzono, Ikumi, and Jun Yokoyama. "Impacts of Alien Bees on Native Plant- 261 pollinator Relationships: A Review with Special Emphasis on Plant 262 Reproduction." Applied Entomology and Zoology 45.1 (2010): 37-47. Print. 263 Fernandes, F. L., M. E. D. Fernandes, M. C. Picanco, G. C. Geraldo, A. J. Demuner, 264 R.S. da Silva. 2010. Coffee Volatiles and Predatory Wasps (Hymenoptera: Vespidae) 265 of the Coffee Leaf Miner Leucoptera coffeella. Sociobiology 56(2): 455-464. 266 Goldman, R. L. 2010. Ecosystem Services. Environment Magazine. 267 Gruber, Bernd, Katharina Eckel, Jeroen Everaars, and Carsten F. Dormann. "On 268 Managing the Red Mason Bee (Osmia Bicornis) in Apple Orchards." Apidologie 42.5 269 (2011): 564-76. Print. 270 Hanley, M.e., Franco, M., Dean, C.E., Franklin, E.L., Harris, H.R., Haynes, A. G., 271 Rapson, S.R., Rowse, G., Thomas, K. C., Waterhouse, B. R., and Knight, M. E. 272 (2011) “Increased bumblebee abundance along the margins of a mass flowering crop: 273 evidence for pollinator spillover. Oikos 120: 1618-1624. doi: 10.1111/j.1600- 274 0706.2011.19233.x 275 Henry, Mickaël, Marie Fröchen, Julie Maillet-Mezeray, Elisabeth Breyne, Fabrice 276 Allier, Ean-Francois Odoux, and Axel Decourtye. "Spatial Autocorrelation in 277 Honeybee Foraging Activity Reveals Optimal Focus Scale for Predicting Agro- 278 environmental Scheme Efficiency." Ecological Modeling 225 (2012): 103-14. Print. 279 Hummel, Jeremy, L. M. Clayton, W. George, K. N. Harker, J. T. O’Donovan. 2010. 280 Responses of the parasitoids of Delia radicum (Diptera: Anthomyiidae) to the 281 vegetational diversity of intercrops. Biological Control 55(3): 151-158. 282 Jha, S., and C. W. Dick. "Native Bees Mediate Long-distance Pollen Dispersal in a 283 Shade Coffee Landscape Mosaic." Proceedings of the National Academy of Sciences 284 107.31 (2010): 13760-3764. Print. 285 Kugimiya, Soichi, M. Uefune, T. Shimoda, J. Takabayashi. 2010. Orientation of the 286 parasitic wasp, Cotesia vestalis (Haliday) (Hymenoptera: Braconidae), to visual and 287 olfactory cues of field mustard flowers, Brassica rapa L. (Brassicaceae), to exploit 288 food sources. Applied Entomology and Zoology 45(3): 369-375. 289 Kremen, Claire, N. M. Williams, M. A. Aizen, B. Gremmill-Herren, G. LeBuhn, R. 290 McKinley, L. Packer, S. G. Potts, T. Roulston, I. Steffan-Dewenter, D. P. 291 Vasquez, R. Winfree, L. Adams, E. E. Crone, S. S. Greenleaf, T. H. Keitt, A. M. 292 Klein, J. Regetz, T. H. Rickets. 2007. Pollination and other ecosystem services 293 produced by mobile organisms: a conceptual framework for the effects of land-use 294 change. Ecology Letters 10: 299-314. 295 Losey, J. E. and Mace Vaughan. 2006. The Economic Value of Ecological Services 296 Provided by Insects. BioScience 56(4): 311-323. 297 Patiny, Sebastien. "Paleoecology of Bees and Plants." Evolution of Plant-pollinator 298 Relationships. Cambridge: Cambridge UP, 2012. 138. Print. 299 Peakall, Rod, D. Ebert, J. Poldy, R. A. Barrow, W. Francke, C. C. Bower, F. P. 300 Schiesti. 2010. Pollinator specificity, floral odour chemistry and the phylogeny of 301 Australian sexuall deceptive Chiloglottis orchids: implications for pollinator-driven 302 speciation. New Phytologist 188(2): 437-450. 303 Rortais, Agnes, C. Villemant, O Gargominy, Q. Rome, J. Haxaire, A. 304 Papachristoforou, G. Arnold. 2010. A new enemy of honeybees in Europe: the 305 Asian hornet, Vespa velutina. Atlas of biodiversity risk 181 306 Shuttleworth, A. and Steven D. Johnson. 2010. The missing stink: sulfur compounds 307 can mediate a shift between fly and wasp pollination systems. Proceedings of the 308 Royal Society B-Biological Sciences 277(1695): 2811-2819. 309 Tonietto, Rebecca, Jeremie Fant, John Ascher, Katherine Ellis, and Daniel 310 Larkin. "A Comparison of Bee Communities of Chicago Green Roofs, Parks and 311 Prairies." Landscape and Urban Planning 103.1 (2011): 102-08. Print. 312 Willmer, Pat. "Generalist Flowers and Generalist Visitors: Hymenoptera." 313 Pollination and Floral Ecology. Princeton, NJ: Princeton UP, 2011. 295-98. Print. 314 Woelke, Joop. 2010. Parasitoids of the large cabbage white (Pieris brassicae). Vlinders 315 25(3): 10-13. 316 317 Table 1. Flower species present in each plot. 318 Annual Basic Annual Diverse Perennial Basic Perennial Diverse Combo Blanketflower( Blanketflower Blanketflower Dense Gayfeather Dense Gayfeather Gaillardia (gaillardia (gaillardia (Liatris spicata) (Liatris spicata) pulchella) pulchella) pulchella) Dye Flower Dye Flower Dye Flower Black-eyed Susan Black-eyed Susan (Coreopsis (Coreopsis (Coreopsis (Rudbeckia hirta) (Rudbeckia hirta) basilis) basilis) basilis) Laevenworth's Laevenworth's White Wild Dotted Horsemint Partridge Pea Coreopsis Coreopsis Indigo (Baptisia (Monarda (Chamaechrista (Coreopsis (Coreopsis alba) puncata) fasciculata) laevenworthii) laevenworthii) Narrowleaf Partridge Pea Partridge Pea Osceola Clover Ironweed Sunflower (Chamaechrist (Chamaechrista (Trifolium (Vernonia (Helianthus a fasciculata) fasciculata) repens) gigantea) angustifolius) Crimson Pinebarren Golden Tickseed Ironweed Clover Goldenrod Tropical Sage (Coreopsis (Vernonia (Trifolium (Solidago (Salvia coccinea) tinctoria) gigantea) incarnatum) fistulosa) Pinebarren Ironweed Tropical Sage Goldenrod Dense Gayfeather (Vernonia (Salvia coccinea) (Solidago (Liatris spicata) gigantea) fistulosa) Standing White Wild Cypress Black-eyed Susan Indigo (Baptisia (Ipomoposis (Rudbeckia hirta) alba) rubra) Lanceleaf Drummond White Wild Tickseed Phlox (Phlox Indigo (Baptisia (Coreopsis drummondii) alba) lanceolata) Button Pinebarren Crimson Clover Rattlesnakemaster Goldenrod (Trifolium (Enungium (Solidago incarnatum) yiccifoliumi) fistulosa) Butterfly Weed Osceola Clover (Asceipias (Trifolium tuberosa) repens) Osceola Clover (Trifolium repens) 319 320 321 322 323 324 Table 2. Number of individuals and families caught per minute325 on plot. Data are mean + standard error individuals/families326 caught per minute, N. Data in columns followed by the same327 letter are not different at P < 0.05. Student T tests were used to compare means. Flower Strip # individuals/min # families/min Annual Diverse 0.64 + 0.18, 4a 0.14 + 0.04, 4a Perennial Diverse 0.58 + 0.27, 4ab 0.12 + 0.05, 4a Combo 0.53 + 0.17, 4bc 0.15 + 0.04, 4a Annual Basic 0.53 + 0.13, 4bc 0.18 + 0.05, 4a Control 0.44 + 0.21, 4bc 0.07 + 0.01, 4a Perennial Basic 0.42 + 0.14, 4c 0.10 + 0.03, 4a

ANOVA --> F = 0.1893; df = F = 0.967; df = 5,23; P =0.01 5,23; P > 0.05 328 329 Table 3. Number of individuals and families caught per minute during each sampling period. Data are mean + standard error individuals/families caught per minute, N. Data in columns followed by the same letter are not different at P < 0.05. Tukey tests were used to compare means. Collection # # individuals/min # families/min 1 0.2472 + 0.04, 6b 0.0612 + 0.01, 6b 2 0.8223 + 0.14, 6a 0.1225 + 0.02, 6ab 3 0.5777 + 0.14, 6ab 0.2111 + 0.04, 6a 4 0.4458 + 0.13, 6ab 0.116 + 0.02, 6b

ANOVA --> F = 4.0547; df = F = 7.2479; df = 3,23; P = 0.0210 3,23; P = 0.0018 330 331 332 333 Table 4. Number of individuals caught per strip in each family. Data are mean + standard error number of individuals. Flower strip (plot) did not affect the number of individuals caught (F = 1.87; df = 5,287; P = 0.10); neither did the interaction between flower strip and family (F = 1.07; df = 55,287; P = 0.37). Data in columns followed by the same letter are not different at P < 0.05. Tukey tests were used to compare means. Family # Individuals Crabronidae 6.67 + 1.52, 24a Sphecidae 4.92 + 0.93, 24b Scoliidae 3.42 + 1.12, 24c Mutillidae 1.42 + 0.35, 24d Ichneumonidae 1.08 + 0.63, 24de Halictidae 1.00 + 0.38, 24de Vespidae 0.42 + 0.17, 24ef Pompiliidae 0.38 + 0.17, 24ef Tiphiidae 0.29 + 0.11, 24ef Chrysididae 0.25 + 0.15, 24ef Braconidae 0.13 + 0.09, 24f Chaclididae 0.08 + 0.06, 24f

ANOVA --> F = 11.0981; df = 11,287; P < 0.0001 334 335 336 Figure 1. Diagram of spatial configuration of Site 1 (Citra, Marion Co., Florida) 337

338 339 340 341 Figure 2. Diagram of spatial configuration of Site 2 (Citra, Marion Co., Florida). 342

343