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Foraging Populations and Distances of the Desert Subterranean Termite, Heterotermes Aureus (Isoptera: Rhinotermitidae), Associated with Structures in Southern Arizona

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Foraging Populations and Distances of the Desert Subterranean Termite, Heterotermes Aureus (Isoptera: Rhinotermitidae), Associated with Structures in Southern Arizona HOUSEHOLD AND STRUCTURAL INSECTS Foraging Populations and Distances of the Desert Subterranean Termite, Heterotermes aureus (Isoptera: Rhinotermitidae), Associated with Structures in Southern Arizona 1 2 PAUL B. BAKER AND MICHAEL I. HAVERTY J. Econ. Entomol. 100(4): 1381Ð1390 (2007) ABSTRACT MarkÐreleaseÐrecapture studies were conducted on foraging populations of Hetero- termes aureus (Snyder) (Isoptera: Rhinotermitidae) associated with three structures in Tucson, AZ. Foraging population estimates ranged from 64,913 to 307,284 termites by using the Lincoln Index and from 75,501 to 313,251 termites using the weighted mean model. The maximum distance between monitors ranged from 26 to 65 m, with minimum total foraging distance ranging between 297 and 2,427 m. Characterizations of the cuticular hydrocarbons of foraging groups were qualitatively identical. Quantitative similarities within sites and differences among sites suggested that each site was occupied by a single colony during the sampling period. The colony at each site had a proportion of soldiers (0.135, 0.069, and 0.040) that was signiÞcantly different from the colonies at each of the other sites. From this study, we question the assumption of equal mixing of marked H. aureus foragers throughout the occupied collars around structures. KEY WORDS colony density, colony size, cuticular hydrocarbons, markÐreleaseÐrecapture, soldier proportions Subterranean termites have signiÞcant economic im- tion of foraging populations requires knowledge of pact worldwide. In the United States, subterranean foraging biology. However, our ability to understand termites cost consumers at least US$1.5 billion (Su and the population and foraging dynamics of H. aureus is Scheffrahn 1990). Species of Reticulitermes, Copto- restricted by its cryptic nature. Direct and indirect termes formosanus Shiraki, and Heterotermes aureus methods of sampling H. aureus in undeveloped, native (Snyder), are among the most economically impor- environments have produced asymmetrical results tant pests of structures in the mainland United States ranging from 23,000 to 300,000 individuals per colony (Su and Scheffrahn 1990, Baker and Bellamy 2006), yet on the same site near Tucson, AZ (Haverty et al. 1975, there are relatively few studies of colony demograph- Haverty and Nutting 1975, Jones 1990b). ics or foraging characteristics. With the development We report here estimates of the foraging popula- of long-lasting dyes for marking foragers (Su et al. tions and foraging distances, as well as soldier propor- 1991) or ßuorescent paint markers (Forschler 1994), tions, of colonies of H. aureus associated with struc- markÐrecapture and markÐreleaseÐrecapture meth- tures in three different locations in Tucson, AZ. We ods have been used extensively to study and monitor used a markÐreleaseÐrecapture protocol similar to that subterranean termites worldwide, but not without used to assist the development of baits for control of controversy (Thorne et al. 1996, Evans et al. 1998). subterranean termites (Su and Scheffrahn 1994, Getty However, despite drawbacks, including wide varia- et al. 2000) and characterizations of cuticular hydro- tions in the estimated number of termites, Baroni- carbons (Haverty et al. 1996) and soldier proportions Urbani et al. (1978) suggested this technique repre- to associate foraging groups within a colony. An un- sents at least a practical approach for estimating social derstanding of the magnitude of the foraging distances insect population sizes. and populations is key to effective deployment of baits In Arizona, H. aureus is the most economically im- for protection of structures in urban settings, as well portant termite pest of structures. Control by destruc- as understanding the role H. aureus plays in natural systems. 1 Corresponding author: Department of Entomology, University of Arizona, Tucson, AZ 85720 (e-mail: [email protected]). Materials and Methods 2 Chemical Ecology of Forest Insects, PaciÞc Southwest Research Station, U.S. Department of AgricultureÐForest Service, P.O. Box 245, Sites. Three structures located in Tucson, AZ, with Berkeley, CA 94701; and Division of Organisms and the Environment, active infestations of H. aureus were chosen for this Department of Environmental Science, Policy, and Management, College of Natural Resources, 137 Mulford Hall 3114, University of study. None of the structures had received a termiti- California, Berkeley, CA 94720. cide application in the previous 5 yr. Site 1 is the 0022-0493/07/1381Ð1390$04.00/0 ᭧ 2007 Entomological Society of America 1382 JOURNAL OF ECONOMIC ENTOMOLOGY Vol. 100, no. 4 Fig. 1. Diagram of collar deployment at site 1, the University of ArizonaÕs Environmental Research Laboratory (Tucson, AZ). University of ArizonaÕs Environmental Research Lab- paver was placed on top. Approximately 5 mo after the oratory located near the Tucson International Airport. establishment of the collars, a tripleÐmarkÐrecapture This 70 m2, split-level structure, with a half basement, program (Su et al. 1993) was used to delimit colonies is made of slump block walls (Fig. 1). Site 2 is called and estimate the foraging populations and distances. the Shrimp House, and it is located on the West Ag- In addition, Sentricon monitoring stations (Dow Agro- ricultural Campus of the University of Arizona (Tuc- sciences LLC, Indianapolis, IN) were placed equidis- son, AZ). This concrete block structure covers 139.4 tance or 3.3 m between collars, but they were used m2, and it was built 3.2 m below the soil surface with only for evaluating foraging distances when occupied Ϸ0.7 m above ground (Fig. 2). Site 3, the Olson res- by marked termites. Sentricon Termite Colony Elim- idence, is a 337-m2 home with a standard ßoating slab, ination System is a Dow AgoSciences product devel- and it is constructed of slump blocks (Fig. 3). oped for the commercial termite market for use by Colonies, Population Estimates, and Foraging Dis- pest management professionals. tances. Size and dispersion of foraging populations Termites were captured, marked, released, and re- were estimated with markÐreleaseÐrecapture studies. captured over a period of Þve consecutive weeks in Monitoring stations (collars) were placed at 3.3-m the late summerÐfall 2000. The number of workers and intervals around each structure. Collars consisted of soldiers was determined by direct counts of individ- 15.8-cm-long by 15.8-cm-diameter polyvinyl chloride uals. One collar with at least 200 termites was selected pipe forced 2 mm into the ground. Within each collar and force-fed Þlter paper (Whatman no. 1, Whatman, we placed a 10- by 120-cm section of rolled cardboard Maidstone, United Kingdom) with 0.1% (wt:wt) Nile (B ßute, SF Roll Corp., Tucson Container Corp., Tuc- Blue A (Sigma-Aldrich, St. Louis, MO) (Su et al. 1993, son, AZ). A piece of 2.5-by 1.5- by 10-cm ash (Fraxinus Forschler and Townsend 1996) in an 85-mm petri dish sp.) was positioned in the center, and cardboard was in complete darkness at 25ЊC for 7 d. After 7 d, blue wrapped around the wood and held in place with a termites were counted, and they were returned to the rubber band. Once the rolled cardboard was placed monitoring collar from which they had been collected inside the collar, a 16- by 16- by 1.5-cm concrete brick (Su et al. 1993). August 2007 BAKER AND HAVERTY:FORAGING POPULATIONS OF THE DESERT SUBTERRANEAN TERMITE 1383 Fig. 2. Diagram of collar deployment at site 2, the Shrimp House, West Agricultural Campus, University of Arizona (Tucson, AZ). Collections were made every 7 d after the release of tances between all possible pairs of collars occupied the marked termites, for 21 d. Termites were collected by the colony. These measurements assume foraging from all collars and brought back to the laboratory for galleries between collars for sites 1 and 2 were along sorting and counting. Termites found within a collar solid structure guidelines, such as block walls, whereas that contained marked termites at any time within the at site 3 it was calculated based on a straight line, and 21-d sampling period were considered nestmates. All thus is a conservative estimate. foraging nestmates within these collars were fed dyed Relationship of Foraging Groups by Quantification Þlter paper for another 7 d and released back into their of Cuticular Hydrocarbon Mixtures. Termites were corresponding location. The weighted mean model gathered from individual monitoring collars to char- (WMM) was used to estimate termite foraging pop- acterize the cuticular hydrocarbons of each foraging ulations by counting the numbers of marked and un- group. Our hypothesis was that foraging groups from marked termites collected and the number of marked the same colony would have cuticular hydrocarbon termites released at each cycle (Begon 1979, Haverty mixtures that were quantitatively similar and that et al. 2000). As a comparison, population estimates were quantitatively different from foraging groups based on the Lincoln Index (LI) (Bailey 1951) were that belonged to a different colony. To validate this made using the Þrst markÐreleaseÐrecapture cycle. methodology, we compared the cuticular hydrocar- For the purposes of this article, we considered a bon mixtures of foraging groups from the three sites, colony to be foraging groups of H. aureus sharing because we were certain that the sites shared no col- interconnected galleries (Su and Scheffrahn 1996). onies in common. Our deÞnition assumes that these foragers also are Termite samples were collected 2 or 6 mo after the associated with other conspeciÞcs involved in coop- markÐreleaseÐrecapture cycle, and they were brought erative rearing of offspring (Wilson 1971). The max- to the laboratory at the University of Arizona, where imum foraging distance between collars used by a the termites were separated from cardboard and any colony was determined by measuring (or calculating) other debris. Samples of 50Ð200 foragers (pseuder- the linear distance between the two connecting col- gates or workers) were placed in separate vials, frozen, lars that are furthest apart. The minimum total forag- and then dried (Haverty et al.
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