Subterranean termites of the Oklahoma tallgrass prairie preserve cross timbers.
|Abstract:||A study was conducted to characterize termite colonies on the Nature Conservancy's Tallgrass Prairie Preserve Cross Timbers habitat in northeastern Oklahoma. The two test sites were established on a prescribed-burn area and no-burn area of the Cross Timbers habitat. Termites were identified through both morphological and molecular analyses. Foraging areas of five colonies were delineated. Numbers of termites in foraging groups, estimated using the 'weighted mean model', ranged from 103,093 ([+ or -] 7081) to 422,780 ([+ or -] 19,297) for Reticulitermes flavipes within the prescribed-burn area, and 44,179 ([+ or -] 4879) to 207,141 ([+ or -] 9190) for R. hageni within the no-burn area. Soldier percentages were determined for each foraging group. Estimates of foraging areas and populations are compared with those from previous studies in dissimilar tallgrass prairie habitats. Improved understanding of termite colony densities in various natural habitats provides an increased understanding of termite input in rural areas and could aid in the development of management strategies.|
(Protection and preservation)
Timber (Protection and preservation)
Smith, Matthew P.
Smith, Anita L.
Broussard, Greg H.
|Publication:||Name: The American Midland Naturalist Publisher: University of Notre Dame, Department of Biological Sciences Audience: Academic Format: Magazine/Journal Subject: Biological sciences; Earth sciences Copyright: COPYRIGHT 2012 University of Notre Dame, Department of Biological Sciences ISSN: 0003-0031|
|Issue:||Date: Jan, 2012 Source Volume: 167 Source Issue: 1|
|Organization:||Organization: Oklahoma State University|
|Geographic:||Geographic Scope: Oklahoma Geographic Code: 1U7OK Oklahoma|
Subterranean termites are valuable components of their ecosystems, recycling cellulosic materials and influencing soil fertility, composition and structure. The majority of manmade structures in the United States of America (USA) contain wooden components that are susceptible to termite attack. In the USA alone, termites are responsible for $1 billion to $11 billion in annual expenditures for preventive treatments and repair of structural damage (Su and Scheffrahn, 1990; Kard, 2003).
A better understanding of how termites affect natural and urban environments requires greater knowledge of the biology and behavior of Reticulitermes sp. (Isoptera: Rhinotermitidae). Termites responsible for the majority of damage to wooden structures in the USA are subterranean in nature, making it difficult to gather data on the size and distribution of their colonies. However, several investigators have studied termite behavior in a variety of habitats. In a Canadian study, foraging areas, maximum linear foraging distances, and foraging population estimates for R. flavipes colonies ranged from 266 to 1091 [m.sup.2], 48 to 79 m, and 2.1 to 3.2 million individuals, respectively (Grace et al., 1989). In Florida, Su et al. (1993) estimated R. flavipes foraging areas, maximum linear foraging distances, and foraging populations of 18 to 2361 [m.sup.2], 7.3 to 70.6 m, and 171,000 to 5.01 million individuals, respectively. Maximum linear foraging distances, cumulative foraging distances among monitoring stations, and foraging population estimates for Reticulitermes sp. colonies in California, were recorded as 25.3 m, 291.6 m, and 4500 to 492,000 individuals, respectively (Haverty et al., 2000). Additionally, Reticulitermes sp. foraging populations in Georgia ranged from 106 to 3.5 million individuals (Forschler and Townsend, 1996).
Historically, pest management professionals have relied on persistent, broad spectrum, repellant compounds to prevent and treat termite infestations. These termiticides are applied to the soil around and beneath a structure, creating a continuous chemical barrier. More recent advancements have included the use of non-repellents, targeted applications, and baits. These newer, more directed methodologies make understanding termite foraging density and behavior imperative.
Within Oklahoma, the population density of subterranean termites ranges from relatively moderate to high, except for counties located in the subtropical, extreme southeast where densities are considered 'very heavy' (Suiter et al., 2002). To date, four studies documenting distribution and behavior of termites in limited geographic areas of Oklahoma have been completed (Brown et al., 2004; Brown, 2005; Brown et al., 2008; Smith et al., 2010). The current study was conducted on the Nature Conservancy's Tallgrass Prairie Preserve (TGPP), 89 km northwest of Tulsa and 16 km north of Pawhuska, Oklahoma, in Osage County (36[degrees]50'N, 96[degrees]24'W). Two study sites established in the current study were located on Cross Timbers areas, one where prescribed burns were conducted and one where burning was excluded.
The objectives of this study were to investigate foraging behavior and colony characteristics of Reticulitermes sp. within the Cross Timbers. Our studies provide additional information on termite biology and behavior on the TGPP within two different Cross Timbers habitats. This information combined with previous studies conducted on the tallgrass area (Brown et al., 2008), may lead to improved and more effective termite management strategies.
MATERIALS AND METHODS
Monitoring devices.--In-ground stations consisted of cylindrical 10.2-cm inside diameter (i.d.) polyvinyl chloride (PVC) pipe cut to 20.3-cm lengths. Each pipe had four equally spaced, parallel rows of twelve 3.2-mm-diameter drill holes running lengthwise. Drill holes began 1.3 cm from one end of the pipe and were spaced 1.3 cm apart. Each pipe was vertically inserted into an 18-cm-deep hole in the soil that was pre-drilled with a gas-powered auger equipped with a 10.2-cm-diameter bit. Each station was provisioned with a wooden 'sandwich' consisting of seven parallel, rectangular 17.8 x 6.4 x 0.6-cm pine sapwood slats, each separated by a flat wooden tongue depressor and bound together with nylon 'zip' ties. The resulting sandwich was wrapped with a rectangular 37.5 x 18.5-cm section of corrugated cardboard around the long axis and inserted into each PVC station (Figs. la, b). A standard, removable PVC cap was placed on top of each pipe to exclude sunlight, moisture and animals (Fig. 1c) (Brown et al., 2004).
Additionally, soil-surface rectangular ground-boards of fir/spruce/pine, each measuring 30.5 x 15.2 x 2.5 cm, were placed on bare soil (surface vegetation removed). A standard, solid building brick was placed on top of each ground-board to reduce displacement or loss due to severe weather or animal activity (Fig. ld). Ground-boards allow additional monitoring of termite foraging at the soil surface.
Study sites.--Site 1 was established on the prescribed-burn area and Site 2 on the no-burn area. Bison roam freely on the TGPP but are excluded from a 142 ha area that includes the habitats used for this study. The TGPP encompasses 15,659 ha of land consisting mainly of native tallgrass prairie with a north-to-south central swath of Cross Timbers. The Cross Timbers ecosystem consists of xeric oak woodlands interspersed with patches of savanna and prairie that encompasses 4.9 million ha in central southeastern Kansas, central Oklahoma, and northern Texas (Clark and Hallgren, 2003). The Cross Timbers contains prescribed-burn and no-burn sites that are managed by the Nature Conservancy in cooperation with range scientists from Oklahoma State University.
[FIGURE 1 OMITTED]
Prescribed-burn area.--The prescribed-burn area supports vegetation comprised primarily of native grasses and young, regenerating trees. The predominant plants are the legume, Tephrosia virginiana (L.) Pers. (goat's rue), and the grasses, Sorghastrum nutans (L.) Nash (indiangrass), Panicum virgatum L. (switchgrass) and Andropogon gerardii Vitman (big bluestem) (Palmer, 2007). Mature trees are not found on this area due to cyclic prescribed burns. Quercus stellata (Wangenh.) (post oak) and Q. marilandica (Muenchh.) (blackjack oak) regenerate on this area, but most are [approximately equal to] 1.5 m in height, clustered and shrub-like.
Twenty-five in-ground stations were initially installed as a 12.0 x 12.0-m square grid, configured in straight lines in a "checkerboard" arrangement with spacing of 3.0 m between stations (Fig. 2). Additionally, a 9.0 x 9.0-m square grid of 20 ground- boards was overlaid so that each ground-board was centered between four in-ground stations. The result was a total of 45 monitoring devices, each subtending an area of 4.5 [m.sup.2]. This original grid was expanded when stations [less than or equal to] 6 m from the perimeter became active with termites. In such cases, additional stations and ground-boards were added to encompass the colony's foraging area. The prescribed-burn grid was eventually expanded to 136 in-ground stations and 122 ground-boards covering a 31.5 x 39.0-m area (Fig. 2). Three colonies were delineated within this area, during the 2005, 2006, and 2007 growing seasons, and designated as prescribed-burn sites PB05 (2005), PB06 (2006), and PB07 (2007).
[FIGURE 2 OMITTED]
[FIGURE 3 OMITTED]
No-burn area.--This area is populated by a mature stand of blackjack oak averaging 6-8 m in height. Fifty in-ground stations were installed on a 27.0 x 12.0-m rectangular grid with 3.0-m spacing between stations. Additionally, a 27.0 x 12.0-m rectangular grid of 50 ground-boards was overlaid resulting in a total of 100 monitoring devices. The overlay spacing was the same configuration as the prescribed-burn site and was expanded as needed to encompass all detectable termite activity. This grid was eventually expanded to 75 in-ground stations and 82 ground-boards (Fig. 3). Two colonies were delineated within this area, during the 2006 and 2007 growing seasons, and designated no-burn sites NB06 (2006) and NB07 (2007).
Morphological identification.--Termites collected from the study areas were preserved in 99.5 + % ethyl alcohol. Standard taxonomic dichotomous keys were used to identify the termites to species (Banks, 1946; Gleason and Koehler, 1980; Scheffrahn and Su, 1994; Brown et al., 2005).
Molecular techniques.--Termites were also identified using polymerase chain reaction (PCR) techniques. Two termites from each sample of preserved specimens were placed on filter paper and allowed to dry at room temperature. The DNA of whole workers from each sample group was extracted using a Qiagen DNeasy[R] Tissue Kit (Qiagen Sciences, Germantown, MD). The extract was quantified using a ND-1000 nanodrop spectrophotometer located in the Oklahoma State University (OSU) Biochemistry Microarray Core Facility. Extracts with a 260/280 ratio below 1.6 or with a mass <10 ng/[micro]l were discarded. Extracts were amplified using FastStart PCR Master[R] (Roche, Indianapolis, IN). The PCR primers LR-J-13017 (5'-TTACGCTGTTATCCCTAA-3') (Kambhampati and Smith, 1995) and LR-N- 13398 (5'-CGCCTGTTTATCAAAAACAT-3') (Austin et al., 2004) were used to amplify a [approximately equal to]428 bp region of the mtDNA 16S rRNA gene. The product was cleaned of excess dNTPs and primers using ExoSAP-IT[R] (USB, Cleveland, Ohio), and a combination of the hydrolytic enzymes Exonuclease I and Shrimp Alkaline Phosphatase. A sample of the resulting product was submitted to the OSU Recombinant DNA/Protein Resource Facility for sequencing. Sequencing was performed with the Applied Biosystems BigDye[R] Terminator Cycle Sequencing Kit version 1.1 using standard protocols and analyzed with an Applied Biosystems Model 3730 DNA Analyzer. The resulting sequence was submitted to the National Center for Biotechnology Information (NCBI) website and compared with known sequences using the Basic Local Alignment Search Tool nucleotide collection (BLASTn). Upon verification of correct morphological identification, a consensus sequence was identified using the ClustalW program at EMBL-EBI (European Bioinformatics Institute). The consensus sequences used for all subsequent molecular identifications were: AY257235.2 (Reticulitermes hageni) and DQ001971.1 (R. flavipes).
Delineation of foraging areas.--A triple-mark-release-recapture (TMRR) technique (Begon, 1979; Haverty et al., 2000) was used to delineate termite foraging areas. The cardboard-wrapped sandwich within each station that became active with termites was removed and placed individually into plastic containers for transportation to the laboratory. A new cardboard-wrapped sandwich was then placed into the station. Once at the laboratory, the termite-infested sandwich was placed on a plastic tray and dismantled to facilitate termite removal. A low-vacuum aspirator was used to aspirate the termites into a collection tube. Termites were then counted and sorted according to caste (worker, soldier, or nymph).
Two pieces of Whatman[R] #1 90-mm-diameter filter paper, that were previously impregnated with 0.1% (wt/wt) Nile Blue A dye (Aldrich, Milwaukee, WI), were moistened using reverse-osmosis water, pressed between two paper towels to remove excess water and placed into a 10.2-cm-diameter x 6.4-cm-tall plastic container (Haagsma and Rust, 1993). Approximately 500 termites were placed into the container and maintained under dark conditions at 22 C with [greater than or equal to] 95% humidity and allowed to feed on the filter paper for 14 d. Termites were recounted to determine the number of dyed and non-dyed termites to be returned to the field. Termites were returned to their original station fifteen days after collection. Released termites were allowed to disperse throughout the colony for a period of 2 wk. Following that period, the contents from active monitoring stations surrounding the original station were collected, all termites were counted, and the number of dyed and non-dyed termites determined. The TMRR process was repeated twice. During the second and third TMRR collections, termites were counted according to caste and color: non- dyed worker, soldier and nymph, and dyed worker, soldier and nymph. Subsequent TMRR foragers collected from different monitoring devices were fed filter paper impregnated with 0.5% (wt/wt) Neutral Red dye (Aldrich, Milwaukee, WI) (Haagsma and Rust, 1993). Using different dyes ensured colonies were readily delineated.
Foraging population estimates.--Both Lincoln Index and weighted mean model calculations were used to estimate the number of foraging termites for each delineated colony (Lincoln, 1930; Bailey, 1951; Begon, 1979; Grace et al., 1989; Su, 1993; Haverty et al., 2000; Brown et al., 2008). For the Lincoln Index, P = Mn/m and SE = [square root of [M.sup.2]n(n - m)/[m.sup.3], where P = total foraging population, M = number of termites marked and released, n = total number of termites recaptured in the second sample and m = number of marked termites recaptured in the second sample. For the weighted mean model, P = ([summation] [M.sub.i][n.sub.i]/[([summation] [m.sub.i]) + 1] and SE = [square root of [1 / ([summation] [m.sub.i] + 1)] + [2/[([summation] [m.sub.i] + 1).sup.2]] + [6/[([summation] [m.sub.i] + 1).sup.3]]]] where P = number of foraging termites, M = total number of marked termites released, n = total number of termites recaptured and m = number of marked termites recaptured. Soldier percentages within the foraging populations were calculated as (total # soldiers/ (total # soldiers and workers)) x 100.
Foraging population comparison.--Estimates of foraging populations from the prescribed-burn sites were compared with those from the no-burn sites using contrast analysis (Steel and Torrie, 1980). These analyses compared foraging populations of two different termite species located in two different habitats. Our null hypothesis was foraging populations within prescribed-burn sites are equal to foraging populations within no-burn sites.
RESULTS AND DISCUSSION
Species identification.--Reticulitermes flavipes and R. hageni were found within the study sites. Morphological identifications of R. hageni soldiers were consistent with molecular identifications. Due to overlap of morphological measurements between soldiers of R. flavipes and R. virginicus, both found in Oklahoma, R. flavipes species designations based on soldier morphology were confirmed molecularly. During this study, termites within the prescribed-burn area were exclusively R. flavipes, whereas R. hageni was the only species found in the no-burn area. Discovery and identification of R. hageni was notable because all termites previously identified on the TGPP were R. flavipes. Subsequent sampling and identifications conducted after the completion of this study found that R. hageni also inhabited the prescribed-burn area, and R. flavipes was found within the no-burn area, thus both species inhabit these different vegetative regimes.
Several hundred additional termites collected from one in-ground station on the prescribed-burn area were identified as Reticulitermes tibialis. Reticulitermes virginicus has never been collected on the TGPP. This species may be present and simply not encountered as R. virginicus is rarely collected from in-ground stations (Haverty et al., 1999). A similar result was seen in a recent survey of Oklahoma termites as R. virginicus were not collected from inground stations or ground boards located across southern counties (Smith et al., 2010).
The presence of two Reticulitermes hageni colonies within the no-burn area was notable. This area has a dense shade canopy of scrub blackjack oak. A sample from a third R. hageni colony located on the prescribed-burn area, collected after the completion of this study, came from an in-ground monitoring station located in full shade. This finding corroborates results of the survey by Smith et al. (2010), in which all R. hageni samples were collected from shaded areas. These studies suggest that in Oklahoma, R. hageni preferentially colonizes shaded areas possibly due to cooler temperatures, wetter soil, and increased woody debris usually associated with full-shade areas. These observations contrast with those of Houseman (1999), who showed R. hageni prefer warm, dry conditions in Texas. The colonization preference of R. hageni in varying habitats merits further study.
Delineation of foraging areas.--Three foraging populations, PB05, PB06, and PB07, were identified within Site 1 on the prescribed-burn area, and two foraging populations, NB06 and NB07, were identified within Site 2 on the no-burn area (Figs. 2, 3). Foraging areas ranged from 13.5 to 58.5 [m.sup.2] and 22.5 to 27.0 [m.sup.2] on the prescribed- burn and no-burn sites, respectively (Table 1). Estimated foraging areas for individual colonies on the nearby tallgrass area ranged from 9.0 to 92.3 [m.sup.2] (Brown et al., 2008). The mean foraging area on the Cross Timbers area was 28.8 [m.sup.2] compared with 42.0 [m.sup.2] for the tallgrass area. Linear foraging distances ranged from 8.5 to 17.0 m, and 6.7 to 10.5 m on the prescribed-burn and no-burn sites, respectively (Table 1). Linear foraging distances for the tallgrass area ranged from 3.0 to 19.0 m (Brown et al., 2008).
Foraging population estimates.--Foraging termite population estimates ([+ or - ]SE), using the Lincoln Index, ranged between 59,249 ([+ or -]17,732) to 138,641 ([+ or - ]23,378) within the prescribed-burn area, and between 27,715 ([+ or -]5831) to 127,743 ([+ or - ]7373) within the no-burn area. These same estimates, using the weighted mean model, ranged between 103,093 ([+ or -]7081) to 422,780 ([+ or -]19,297) termites within the prescribed-burn area, and 44,179 ([+ or -]4879) to 207,141 ([+ or -]9190) within the no-burn area (Table 2). For four of the five colonies evaluated, estimates of foraging populations using the Lincoln Index were smaller than estimates using the weighted mean model. These variations are consistent with previous studies (Haverty et al., 2000; Brown, 2005).
The number of marked termites recaptured in this study varied from 1 to 4% of the number of marked termites released. Although model assumptions attribute this variability to differences in population sizes, it is also possible that marked termites do not distribute evenly or randomly within a foraging group. Variability in the number of marked termites recaptured could also be affected by loss of coloration from dyed termites. Dyes may not persist for a similar period in all marked termites, thus some previously marked and recaptured individuals may not have been apparent.
It is difficult to verify the validity of estimates of termite foraging populations using either Lincoln Index or the weighted mean model. Although the disparity between such estimates is problematic, these two methods continue to be commonly used to estimate foraging populations when using non-destructive sampling methods.
Discrepancies between mark-recapture techniques were addressed in a study conducted on mound-building subterranean termites in Australia. The results indicate that inherent variation in population estimates is attributable, in part, to violation of the assumptions of mark-recapture protocols (Evans et al., 1998). Other studies concerning forager population estimates determined using marked-recapture techniques often indicate much larger numbers compared with actual counts of all colony members (Forschler and Townsend, 1996; Thorne et al., 1996; Evans et al., 1998). However, mark-recapture field studies continue to provide valuable information, and when coupled with population genetic studies may provide further insight into termite foraging behavior and foraging area.
Although NB06 had an estimated foraging population approximately 4.6 times greater than NB07, NB07 encompassed a larger foraging area. This difference proved true for both the Lincoln Index and the weighted mean model estimates, and may be attributable to the availability of suitable cellulose resources within each colony's foraging area. Colony NB06 was located in a dense thicket of oak trees, whereas the foraging area of NB07 encompassed an open area in the canopy. This open area was marked by the presence of fewer tree roots, increased sunlight penetration, and the proliferation of big bluestem. If Reticulitermes hageni uses tree roots as a primary nutrition source, relatively fewer roots within the area of NB07 could explain the need for the smaller population to forage over a larger area.
Foraging population comparisons.--Contrast analysis indicated a moderate difference between Lincoln Index estimates within the prescribed-burn and no-burn areas, (P = 0.054). This significant difference between areas was more marked, when weighted mean model estimates were evaluated (P = 0.0001). These results must be considered with care as the factors responsible for the observed differences in estimated foraging populations between the two areas are not clear. The most obvious factor other than the area management regime is that the termite activity in the two areas was attributable to two different species. The differences in foraging numbers may be due to the type of habitat within which the termites are found, to interspecific variation, or to other unknown factors. Additional studies evaluating how such factors influence foraging population size would be beneficial.
Soldier percentage determinations.--Soldier percentages of 0.53, 1.86, and 2.08% for Reticulitermes flavipes (Table 2) were lower than those of 2.69, 3.65, and 4.46% recorded for the open prairie area of the TGPP (Brown, 2005). Earlier reports of soldier percentages for mature R. flavipes colonies ranged from 8.4 to 14% (Banks and Snyder, 1920; Haverty, 1977). These numbers indicate soldier percentages may vary depending upon both species and habitat. Previous studies have shown that observed caste ratios may also be influenced by sampling location within the colony area (Brown, 2005). Soldier percentages may be greater in centralized areas of the foraging area, compared with those observed from the colony's periphery (Brown et al., 2008).
The two colonies of Reticulitermes hageni located within the no-burn area had mean soldier percentage estimates <1.00%. This relatively small percentage of soldiers may not simply reflect the fact that the number of soldiers per worker within R. hageni colonies is less than that of R. flavipes but may, in fact, indicate a difference in where soldiers are primarily allocated within the colony's structure.
Soldier ratios were variable, ranging from 0.53 to 1.86% for Reticulitermes flavipes and 0.76 to 0.96% for R. hageni. Soldier percentage data show that colonies of R. flavipes on the TGPP contain fewer soldiers compared with percentages in other published studies (Banks and Snyder, 1920; Haverty, 1977).
Because termites live within the soil, it is not practical to count the total number of termites in a subterranean colony. Therefore, the modification of established wildlife markrecapture techniques to address insect populations is useful but can provide widely disparate numbers (Evans et al., 1998). Both single-mark-recapture as well as multiple- mark-recapture techniques have been used with some success. However, several assumptions concerning dye markers and termite behavior, e.g., dye retention and non-lethality of dye, are required for these techniques to provide accurate estimates. These assumptions do not always hold true, thus foraging population estimates may be much greater than the actual number of termites (Forschler and Townsend, 1996; Thorne et al., 1996). However, these techniques provide useful insight into termite colony characteristics.
Different species of termites predominated within the two different study area habitats. Reticulitermes hageni, a relatively less common species than R. flavipes in Oklahoma, predominated in shady, wooded areas, whereas R. flavipes dominated the grassy, prescribed-burn area. Foraging areas, although variable, were generally similar in both habitats. Mean maximum linear foraging distances, foraging populations, and soldier percentages within the prescribed-burn area were greater than within the no-burn area.
The current study identifies multiple areas for future research that promises to provide a better understanding of the biology and ecology of native subterranean termites and their impacts on various ecosystems. Of particular interest are further analyses of nutritional resources utilized by Reticulitermes hageni within the no-burn area, as well as a comparison of foraging populations/areas between colonies located in shaded and open locations. Such studies should include evaluations of the influence of other abiotic factors such as soil moisture and temperature on termite foraging activity. Finally, further study of caste ratios for both Reticulitermes sp. would aid in better understanding of colony structure. Such information will serve as a base line for evaluating emerging management tools such as the use of insect growth regulators to disrupt normal soldier percentages.
Acknowledgments.--We thank Robert G. Hamilton of The Nature Conservancy's Tallgrass Prairie Preserve for providing field study sites. We thank Jesse Eiben, Doug Kuehl, Matthew and Ngan Rawlings, and Zachary Smith for their assistance in conducting field studies. Thanks also to the reviewers whose comments improved this manuscript. This work was approved for publication by the Director of the Oklahoma Agricultural Experiment Station and supported in part under Project H- 2480.
SUBMITTED 13 AUGUST 2010
ACCEPTED 13 JULY 2011
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MATTHEW P. SMITH, (1) ANITA L. SMITH, AND BRAD KARD,
Department of Entomology and Plant Pathology, Oklahoma State University,
KENNETH S. BROWN
BASF Pest Control Solutions, St. Louis, Missouri 63122
GREG H. BROUSSARD
Department of Math and Science, Connors State College, Warner, Oklahoma 74469
(1) Corresponding author: Telephone: (405) 744-8860; e-mail: firstname.lastname@example.org
TABLE 1.--Foraging areas and maximum linear foraging distance of five colonies of subterranean termite (Retirulitermes sp.) on the Nature Conservancy's Tallgrass Prairie Preserve Cross Timbers Number of Maximum linear active Foraging foraging monitoring territory distance, Colony devices [m.sup.2] [m.sup.2] PB05 * 13 58.5 17.0 PB06 6 22.5 8.5 PB07 3 13.5 9.5 NB06 ** 5 22.5 6.7 NB07 6 27.0 10.5 * PB = prescribed-burn area: 05, 06 and 07 are the year of study, e.g., 05 = 2005. Colonies are R. flavipes ** NB = no-burn area. Colonies are R. hageni TABLE 2.--Average soldier percentages and foraging population estimates (based upon Lincoln Index and weighted mean model calculations) of five colonies of Reticulilermes sp. on the Nature Conservancy's Tallgrass Prairie Preserve Cross Timbers Mark-release-recapture cycle (a) Colony Station MI n1 ml M2 n2 m2 M3 PB05 (b) 13 381 774 16 500 1170 54 949 14 0 2564 2 1947 0 0 0 36 0 2259 4 1569 527 31 446 37 0 2099 4 1661 0 0 0 38 0 1163 2 470 117 11 77 39 0 376 2 0 2273 93 499 H 0 1380 6 844 0 0 0 12 0 0 0 0 377 48 310 AH 0 0 0 0 1163 29 894 U 0 0 0 0 427 5 317 18 0 0 0 0 0 0 0 25 0 0 0 0 0 0 0 O 0 0 0 0 0 0 0 Total 381 10,612 36 6991 6054 271 3492 PB06 70 667 256 1 140 0 0 0 54 0 3728 14 2765 2150 99 1776 55 0 2153 10 1321 0 0 0 87 0 247 3 115 0 0 0 88 0 891 7 308 0 0 0 107 0 0 0 0 0 0 0 Total 667 7275 35 4649 2150 99 1776 PB07 77 876 0 0 0 0 0 0 60 0 655 5 584 2882 15 2309 118 0 89 6 26 153 2 153 Total 876 744 11 610 3035 17 2462 NB06 28E 10,285 2090 179 1868 38 10 5 27D 0 714 41 505 13 5 9 28 F 0 624 56 548 0 0 0 27 F 0 0 0 0 3915 181 3379 28D 0 0 0 0 0 0 0 Total 10,285 3428 276 2921 3966 196 3393 NB07 21D 725 45 5 42 0 0 0 20B 0 471 13 372 0 0 0 20D 0 325 4 296 0 0 0 20A 0 0 0 0 226 31 215 GB20D 0 0 0 0 35 3 24 GB35A 0 0 0 0 434 6 326 Total 725 341 22 710 695 40 565 Mark-release-recapture cycle (a) Weighted Lincoln mean Soldier Index model Colony Station n3 m3 percentages (SE) (SE) PB05 (b) 13 2113 29 2.21 112,342 422,780 14 0 0 1.72 (18,692) (19,297) 36 910 17 3.06 37 0 0 1.72 38 4200 47 1.95 39 2967 39 1.81 H 0 0 1.49 12 0 0 5.57 AH 0 0 1.12 U 0 0 1.81 18 1449 19 0.41 25 1414 12 1.06 O 1224 11 0.41 Total 14,277 174 1.86 (c) PB06 70 0 0 3.47 138,641 103,093 54 337 39 1.96 (23,378) (7081) 55 0 0 0.98 87 364 32 3.11 88 0 0 2.69 107 114 8 2.63 Total 815 79 2.08 PB07 77 431 4 0.36 59,249 212,224 60 328 6 0.44 (17,732) (34,409) 118 0 0 2.89 Total 759 10 0.53 (c) NB06 28E 0 0 0.86 127,743 207,141 27D 0 0 1.93 (7373) (9190) 28 F 460 19 1.11 27 F 39 4 1.14 28D 586 14 0.51 Total 1085 37 0.96 NB07 21D 0 0 0.23 27,715 44,179 20B 0 0 1.91 (5831) (4879) 20D 0 0 0.31 20A 1052 21 0.55 GB20D 0 0 0 GB35A 0 0 1.61 Total 1052 21 0.76 (a) Numbers (1-3) indicate mark-release-recapture-cycle. M indicates the number of marked termites released, n indicates the number of termites recaptured (marked plus unmarked), and m indicates the number of marked termites recaptured (b) PB =prescribed-burn area, R. flavipes, and NB = no-burn area, R. hageni. 05, 06 and 07 are the year of study, e.g., 05 = 2005 (c) Mean of values immediately above
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