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(Hymenoptera: Vespidae) in a Temperate Deciduous Forest

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(Hymenoptera: Vespidae) in a Temperate Deciduous Forest 1068 Florida Entomologist 94(4) December 2011 VERTICAL DISTRIBUTION AND SEASONALITY OF PREDATORY WASPS (HYMENOPTERA: VESPIDAE) IN A TEMPERATE DECIDUOUS FOREST MICHAEL D. ULYSHEN1*, VILLU SOON2 AND JAMES L. HANULA3 1USDA Forest Service, Southern Research Station, Starkville, MS, USA 2Department of Zoology, Institute of Ecology and Earth Sciences, University of Tartu, Estonia 3USDA Forest Service, Southern Research Station, Athens, GA, USA *Corresponding author; E-mail: [email protected] Efforts to investigate the vertical dimension of predators may influence voltinism (number of forests continue to refine our thinking on issues of generations) in some lepidopteran species. biodiversity and ecology. Arthropod communities Vespidae were sampled at two heights as part exhibit a high degree of vertical stratification in of a larger study on the vertical distribution of forests worldwide but the vertical distribution insects in a temperate deciduous forest. Detailed patterns of most taxa remain largely unexplored methodology and results for beetles and bees can or poorly understood. For example, only 2 studies be found in Ulyshen & Hanula (2007) and Uly- provide information on the vertical distribution shen et al. (2010), respectively. Briefly, 12 domi- patterns of predatory wasps in temperate decidu- nant overstory trees were selected in a mature ous forests. In Canada, Vance et al. (2007) found bottomland forest within the Oconee National Sphecidae to be more abundant in flight intercept Forest in Oglethorpe County, Georgia. Each tree traps suspended 18 m or more above the ground had 1 flight intercept trap suspended in the can- than in traps suspended near ground level. Simi- opy (≥15 m) and one suspended near ground larly, in Germany, Sobek et al. (2009) found Sphe- level (0.5 m). The traps, consisting of intersect- cidae and Vespidae (Eumeninae) to produce more ing clear plastic vanes attached to white buck- brood cells in artificial cavities placed in tree ets, were operated continuously for most of the crowns than in those placed near the forest floor. growing season, from 5 Apr to 28 Jun and from Nothing is currently known about how social 12 Jul to 4 Oct 2005. Samples were collected ev- predatory wasps in the subfamilies Vespinae and ery 2 weeks. Polistinae are distributed within forest canopies. In total, 522 vespids were collected represent- This is unfortunate considering the ecological sig- ing 8 species (Table 1). Vespula maculifrons nificance of these large and highly mobile insects. (Buysson) dominated the samples (83%) followed For instance, social vespids represent one of the by Dolichovespula maculata (Linnaeus) (8%) and principal mortality factors for caterpillars in Vespula squamosa (Drury) (4%). Almost all speci- many habitats (Morris 1972; Steward et al. 1988; mens (99%) were captured in the upper traps. Lichtenberg & Lichtenberg 2003). In an Arkansas Vespid abundance increased dramatically in mid forest, for example, Steward et al. (1988) found season, peaking in late Jul and early Aug (Fig. 1). Vespula maculifrons (Buysson) and V. squamosa These results show that vespids, including the (Drury) to be the most important predators of He- ground-nesting Vespula, are much more active in liothis virescens (Fabricius) larvae, removing over the canopy than near the ground. These patterns 90% of those placed 1.8-2.5 m above the ground. likely reflect the greater availability of prey (espe- Similarly, Polistes fuscatus Fabricius and P. cially caterpillars) and carbohydrate sources such dominulus Christ removed 77-99% of Hemileuca as honeydew and sap in the canopy, although the lucina Hy. Edwards placed in a Massachusetts availability of plant fibers, water and nesting lo- garden (Stamp & Bowers 1988). This intense pre- cations may also be important (Richter 2000; dation pressure is known to have immediate ef- Vance et al. 2007; Sobek et al. 2009; Ulyshen fects on caterpillar behavior (Stamp & Bowers 2011). In addition, some vespids are known to cir- 1988) and may select for certain life history traits cle high above feeding sites when returning to as well. For example, by preferentially preying their nests, presumably for orientation purposes upon caterpillars with a wide host range (presum- (Richter 2000). ably because they are more palatable), vespids The striking mid-season increase in vespid may act to increase host-plant specificity over abundance (Fig. 1), an observation consistent evolutionary time (Richter 2000, and references with previous studies (Spradbery 1971; Mac- therein). Donald & Matthews 1981), is particularly inter- The objectives of this paper are to 1) present esting. Considering this seasonal pattern and results on the vertical distribution and seasonal- the extremely high predation rates documented ity of Vespidae from a southeastern U.S. temper- elsewhere (Stewart et al. 1988; Stamp & Bowers ate deciduous forest and 2) propose that these 1988), we speculate that vespids may play a pre- TABLE 1. LIFE HISTORY INFORMATION AND MEAN ± SE (N = 12) ABUNDANCE OF VESPID SPECIES CAPTURED IN FLIGHT INTERCEPT TRAPS SUSPENDED IN THE CROWNS (≥15 M) OR NEAR THE BASES (0.5 M) OF TREES IN A TEMPERATE DECIDUOUS FOREST. Species Sociality Nest location Nest materials ≥15 m 0.5 m Dolichovespula maculata (Linnaeus) social vegetation pulp 3.58 ± 1.64 0.08 ± 0.08 Eumenes fraternus Say solitary vegetation mud 0.08 ± 0.08 0 Polistes annularis (L.) social vegetation pulp 0.83 ± 0.41 0 Polistes metricus Say social vegetation pulp 0.58 ± 0.29 0 Pseudodynerus quadrisectus (Say) solitary in wood mud 0.33 ± 0.19 0.08 ± 0.08 Vespula maculifrons (Buysson) social underground pulp 35.92 ± 11.35 0.25 ± 0.25 Vespula squamosa (Drury) social underground pulp 1.58 ± 0.61 0 Zethus spinipes Say solitary uncertain uncertain 0.17 ± 0.11 0 Total 43.08 ± 11.96 0.42 ± 0.26 Scientific Notes would beofconsiderableinterest. addressingthesequestions Future research 2010). et al. (Murphy spines or 1997) (Camara sequestrationofhostdefensecompounds 1988), etal. nighttimeforaging(Fitzgerald include This might species orearly-seasonconspecifics. thanunivoltine or morphologicalcharacteristics exhibit agreaterincidenceofdefensivebehaviors populationsmay to thesehighmid-seasonalwasp dation pressure. sponse tothecoincidentincreaseinvespidpre- itcouldalsorepresentare- however, species, some For 1991). (Schaefer host-plant quality in uted tothewell-documentedseasonaldecline This patternhastraditionallybeenattrib- liage. their larvalfeedingactivitiesonearlyspringfo- only asinglegenerationyearandfocusing producing are univoltine, deciduous forests ate Manylepidopteranspeciesintemper- doptera. thenumberofgenerations)inLepi- inism (i.e., viously unrecognizedroleindeterminingvolt- combined) in a temperate deciduous forest. combined) inatemperatedeciduousforest. captured attwelvelocations(canopyandgroundtraps ( cussed. in determining lepidopteranvoltinism is dis- The possibleimportance ofthesepredators Aug. andearly peakinginlate Jul ically inmidseason, abundanceincreaseddramat- Vespid upper traps. most allspecimens(99%)were capturedinthe (Linnaeus) and maculifrons The mostcommonspecieswere wasps. a temperatedeciduousforestyielded522vespid ≥ 15 m) and near the bases (0.5 m) of 12 trees in trees 12 of m) (0.5 bases near the and m) 15 We furtherspeculatethatcaterpillars exposed We Fig. 1. Mean±SE( 1. Fig. Flight intercepttrapssuspendedinthecrowns (Buysson), (Buysson), Vespula squamosa Vespula S n UMMARY =12)numberofvespidwasps Dolichovespula maculata Dolichovespula Al- (Drury). Vespula 1069 1070 Florida Entomologist 94(4) December 2011 REFERENCES CITED story: does tree diversity affect bee and wasp com- munities and their natural enemies across forest CAMARA, M. D. 1997. Predator responses to sequestered strata? Forest Ecol. Manag. 258: 609-615. plan toxins in buckeye caterpillars: Are tritrophic in- SPRADBERY, J. P. 1971. Seasonal changes in the popula- teractions locally variable? J. Chem. Ecol. 23: 2093- tion structure of wasp colonies (Hymenoptera: Vesp- 2106. idae). J. Animal Ecol. 40: 501-523. FITZGERALD, T. D., CASEY, T., AND JOOS, B. 1988. Daily STAMP, N. E., AND BOWERS, M. D. 1988. Direct and indi- foraging schedule of field colonies of the eastern tent rect effects of predatory wasps (Polistes sp.: Vespi- caterpillar Malacosoma americanum. Oecologia 76: dae) on gregarious caterpillars (Hemileuca lucina: 574-578. Saturniidae). Oecologia 75: 619-624. LICHTENBERG, J. S., AND LICHTENBERG, D. A. 2003. Pre- STEWARD, V. B., SMITH, K. G., AND STEPHEN, F. M. dation of caterpillars on understory saplings in an 1988. Predation by wasps on lepidopteran larvae Ozark forest. Southeastern Naturalist 2: 423-432. in an Ozark forest canopy. Ecological Entomol. 13: MACDONALD, J. F., AND MATTHEWS, R. W. 1981. Nest- 81-86. ing biology of the eastern yellowjacket, Vespula ma- ULYSHEN, M. D. 2011. Arthropod vertical stratification culifrons (Hymenoptera: Vespidae). J. Kansas Ento- in temperate deciduous forests: Implications for con- mol. Soc. 54: 433-457. servation-oriented management. Forest Ecol. Man- MORRIS, R. F. 1972. Predation by wasps, birds, and ag. 261: 1479-1489. mammals on Hyphantrea cunea. Can. Entomol. 104: ULYSHEN, M. D., V. SOON, AND HANULA, J. L. 2010. On 1581-1591. the Vertical Distribution of Bees in a Temperate De- MURPHY, S. M., LEAHY, S. M., WILLIAMS, L. S., AND ciduous Forest. Insect Conservation and Diversity 3: LILL, J. T. 2010. Stinging spines protect slug cater- 222-228. pillars (Limacodidae) from multiple generalist pred- ULYSHEN, M. D., AND HANULA, J. L.
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