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Published in International Journal of Pest Management Original Paper Targeted elimination of the exotic termite, Nasutitermes corniger (Isoptera: Termitidae: Nasutitermitinae), from infested tracts in southeastern Florida Rudolf H. Scheffrahn1, Hartwig H. Hochmair1, William H. Kern Jr.1, John Warner1, Jan Krecek1, Boudanath Maharajh1*, Brian J. Cabrera2, Robert Hickman3, and Steven Dwinell4 (1) University of Florida, Fort Lauderdale Research and Education Center 3205 College Avenue, Davie, Florida, 33314 (2) Agricultural Commissioner’s Office, Santa Barbara County, 263 Camino del Remedio, Santa Barbara, CA 93110-1335 (3) BASF Pest Control Solutions, 26 Davis Drive, Research Triangle Park, NC 27709 (4) Florida Department of Agriculture and Consumer Services, 3125 Conner Boulevard Suite F, Tallahassee, FL 32399-1650 *Deceased Rudolf H. Scheffrahn (Corresponding author) Email: rhsc@ufl.edu, Telephone: 954-577-6312, FAX: 954-475-4125 Received: Accepted: Published online: ABSTRACT. Populations of the exotic arboreal termite, Nasutitermes corniger, discovered in Dania Beach, Florida, in 2001, were surveyed and treated for eradication beginning in 2003. Most N. corniger nest locations were cryptic and some nests exceeded one million in population. Field efficacy of fipronil (Termidor® SC) and imidacloprid (Premise® 2) were compared in initial treatments and fipronil was selected as the lone residual spray treatment for termite-active sites through 2010. In 2006, expanded survey work was discontinued due to budget cuts. By 2010 all sites within the original survey area were termite-free. In 2011, a new population of N. corniger was discovered beyond the original 2003 infestated tracts. A newly funded 2011-2012 survey revealed a total of four areas outside the original tracts that were infested with N. corniger. No tract treated with fipronil between 2003 and 2010 yielded renewed termite activity in 2011-2012 thus confirming the efficacy of original targeted residual treatments. New procedures using nest-only and fipronil dust treatments were instituted by Florida state officials in May 2012. Many of these newly treated locations still contained live mature nests as of December 2012. Continued spread of N. corniger in

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tropical urban Florida will have serious economic consequences as a result of nuisance and structural damage. KEY WORDS: Arboreal invasive termite; eradication; termiticide application. INTRODUCTION In May 2001, a robust infestation of the exotic arboreal termite, Nasutitermes corniger (Motschulsky), was discovered in Dania Beach, Broward County, Florida (Scheffrahn et al. 2002). Although ship-borne infestations of Nasutitermes had previously been observed in Florida (Scheffrahn and Crowe 2011), this was the first land-based population of a Nasutitermes sp. in the continental U.S. and, at the time, was the first record of any higher termite (family Termitidae) established on a non-endemic land mass. As with Coptotermes spp. (Hochmair and Scheffrahn 2010), this infestation was likely the result of a dispersal fight originating from an infested boat docked in the area. Nasutitermes corniger is an important structural pest over much of Central and South America (Constantino 2002, Torales 2002) and the West Indies including Puerto Rico and the U.S. Virgin Islands (Scheffrahn et al. 2003). This termite has also invaded nonendemic locations in New Guinea (Scheffrahn et al. 2005) and the Bahamas (Scheffrahn et al. 2006). An expanding invasion of N. corniger poses an increasing and permanent economic impact on residents of tropical south Florida and may also have negative ecological consequences. In Puerto Rico, N. corniger is a more damaging and aggressive structural pest than the common subterranean termite, Heterotermes convexinotatus (Snyder) (ME Holston, pers. comm.). Nasutitermes corniger feeds on a broad range of structural lumber and other wood products. As an arboreal species, i.e., one that nests and forages on or above ground level, N. corniger would fill a niche not occupied by the existing subterranean and drywood termites of Florida (Scheffrahn & Su 1994). Because of its environmental visibility, multiyear colony maturation age, once-a-year dispersal flight season, and short flight distance, eradication of N. corniger was deemed feasible. However, initial requests for assistance from the United States Department of Agriculture and the Florida Department of Agriculture and Consumer Services (FLDACS) did not render support for a survey, treatment, or eradication program, apparently because N. corniger is not an agricultural pest. Herein we recount how an eradication effort was initiated and how a series of surveys and chemical treatments resulted in extirpation of massive populations of N. corniger from targeted tracts inside a 170-hectare area of mixed landscape between 2003 and 2010. Furthermore, we report the discovery of additional populations of N. corniger outside the original tracts during 2011 and 2012. We also argue that modification of the original eradication procedures will slow and complicate further eradication attempts. MATERIALS AND METHODS

Area-wide survey

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After observing the severity of the N. corniger infestation in 2002, FLDACS officials agreed to facilitate the organization of a “Tree Termite Task Force” to plan, implement, and verify an eradication program for this pest under Florida statutes. The task force consisted of personnel from State (University of Florida Fort Lauderdale Research and Education Center, FLREC), FLDACS, private (pesticide manufacturers and pest management professionals), and non-profit (pest control associations) entities. Private sector participation was voluntary and extramural funding by FLDACS was limited to field surveys and treatments by FLREC employees. The city of Dania Beach and Broward County officials gave their consent, support, and access to public lands for the eradication effort. Citizens in the eradication zone were informed of the planned treatments and owners of infested properties signed treatment authorization forms. Public support for this project was overwhelmingly positive. In January 2003, an area-wide visual survey was conducted for nests, foraging tubes, foraging sites, and debris harboring live N. corniger populations. Longitude/latitude waypoints for active sites were recorded using a WAAS enabled Global Positioning System (GPS) receiver. A photograph and termite voucher sample was taken at each site. New sites were marked at minimum 7-meter intervals until the entire infestation zone was surveyed. These same survey techniques were carried out through 2012. Chemical treatments Because N. corniger is such a visible species due to its above ground nests and dark foraging tubes, it was decided that eradication could be accomplished most efficiently by direct application of liquid termiticides onto active sites. Fumigation was used to augment outdoor treatments around heavily infested structures. Biological control was not considered because of historical failures against termite field populations (Chouvenc et al. 2011). On 23 April 2003, field treatments were conducted by the Tree Termite Taskforce in which over 70 volunteers and paid employees took part. Infested areas were divided into 12 treatment sectors (Fig. 1). Five sectors were assigned for treatment with Premise 2 (1,000 ppm imidacloprid) and seven with Termidor SC (1,200 ppm fipronil) both of which, at the time, were USEPA registered “new generation” termiticides. The application of Termidor and Premise away from structures was allowed under State of Florida experimental use permit no. FL 03-EUP-X. The sectors assigned to Premise® treatment contained 103 active sites, while those for Termidor® contained 101 sites. A treatment team, each consisting of a state-licensed applicator and two termite spotters holding aerial maps for reference, was assigned to each sector. Nasutitermes corniger activity, located by spotters, was sprayed with either a truck-mounted or backpack applicator. Active areas were defined as nests, foraging tubes, and active feeding sites such as dead tree stumps, branches, cellulosic debris piles, wood implements, structural lumber, wooden pallets, etc. Areas without termite activity were not sprayed. Tracker blue dye was added to all spray solutions to mark sprayed areas and to prevent duplicate spraying of the same area. Two boats in dry storage and two office mobile homes containing aerial colonies of N. corniger were fumigated with Vikane (sulfuryl fluoride) gas fumigant at the non-monitored drywood termite rate. After April 2003, all termiticide applications were limited to Termidor.

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Colony census In 2004, we collected and censused three nests of differing size, and presumably age, in order to estimate colony populations and roughly establish the overall population of N. corniger within the survey area. The smallest colony was collected inside an arm-sized piece of branch wood lying on the ground. The second was a basketball-sized nest collected at the base of a shrub at a different location. The third, and largest nest, was collected at the base of a Pinus tree at a third location. The smallest colony was censused by direct count. The two larger colonies were censused by extracting termites from the nest and suspending them in ethanol. Ten 10ml aliquots were removed from each suspension and counted. Total population was extrapolated by multiplying the mean aliquot count by the total number of 10ml aliquots contained in the whole colony suspension. The suspension was searched visually for kings and queens. RESULTS AND DISCUSSION Area-wide survey

A total of 204 active sites was recorded and mapped onto a geo-referenced aerial photograph of the surveyed area (Fig. 1). Of the ca. 170 hectares surveyed in early 2003, about 20% of the area contained active infestations on commercial, residential, and undeveloped properties (Fig. 1). We estimate that several hundred mature (alate-producing) colonies were in the area at that time. Maximum east-west and north-south separation of active sites was 1,000 and 1,150 meters, respectively. Eighteen structures, including mobile homes, single-family houses (Fig. 2 a-c), commercial buildings, sheds, and boats were infested. Damage to four mobile homes was severe. Many wood items (e.g. plywood and pressboard sheets, a ladder, fencing, decorative pieces, a patio bar, wooden shelves of a portable barbeque, etc.) were also damaged or destroyed. We suspect that N. corniger activity may have even vectored a pathogen that caused the death, within 2 years, of at least six native live oak (Quercus virginiana) trees on one heavily infested yard. After repeated field site visits in Dania Beach, it was determined that surveys were best suited for the beginning and end of the rainy season during May-June and November-December, respectively, because accelerated foraging and nest enlargement by N. corniger during these periods allowed for optimum visual detection of termite activity. In Dania Beach, N. corniger nests were almost exclusively epigeal and situated at the base of trees, palms (especially Sabel palmetto where the termites feed on dried leaf axils), or on other standing objects and structures. Nasutitermes corniger nesting sites vary in response to local precipitation and humidity (see climate classifications in Peel et al. 2007). In tropical high-moisture climates lacking extended dry seasons e.g., central Panama (Thorne & Haverty 2000), Amazonia (Bahder et al. 2009), eastern Nicaragua (Scheffrahn et al. 2005), Papua New Guinea (Roisin & Pasteels 1985), Trinidad (Merritt & Starr 2010), and southern Belize (Scheffrahn unpubl.), nests tend not to be in contact

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with the soil surface, but attached to tree trunks and branches up to tens of meters above the ground. As in Dania Beach, tropical climates with pronounced dry seasons (e.g. Providenciales, Turks and Caicos Is. (Scheffrahn et al. 1990), Pedernales, Dominican Republic, and Barquisimeto, Venezuela (Scheffrahn unpubl.) promote epigeal nest construction by N. corniger, possibly to sequester soil moisture. Ground nests are more difficult to locate because they are often covered with leaf litter and debris (Fig. 2 d,e) and require careful field observation to be found. Nests hidden in grass tussocks such as Tripsacum dactyloides (Fig. 2g), also occur in drier climates and are very difficult to detect. Even during prolonged dry seasons, N. corniger colonies do not require permanent sources of free water. Many of the colonies in Dania Beach were beyond the foraging range of waterways and lakes that are typical of the area. On Providenciales, N. corniger colonies were common at the base of Coccothrinax palms at least 0.5 km from the nearest body of water. In 2006, funding from FLDACS was eliminated due to the Florida’s economic downturn. At that point, surveys of the original 170-hectare area were discontinued because of the intense labor involved. It was decided, however, to use limited discretionary research funding held by RHS to complete the survey and treatment cycles of known tracts of activity until termites were no longer detected within them. Chemical treatments On April 23, 2003, a total of 1,272 liters (1.26 kg AI) of Premise 2 and 1,504 liters of Termidor SC (1.88 kg AI), were applied to outdoor infestations of N. corniger assigned to each termiticide. Eighteen kg of Vikane (sulfuryl fluoride, DowAgroSciences) was used to fumigate the four structural infestations of N. corniger. Two months after termiticide application, 71 active sites were located during the first post-treatment survey of the entire area. No live termites were detected in the four fumigated structures. Sixty-four active sites were recorded in Premise-treated sectors, while seven active sites were found in sectors treated with Termidor . All five sectors treated with Premise had at least three active sites while only two out of seven sectors treated with Termidor contained live termites (

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Table 1). A chi-square test of independence was performed to examine the relation between treatment efficacy and chemical used. The relation between these two variables was significant, ƛ2(1, N=275)=34.7, p<.0001. Because N.corniger was less likely to be found in areas treated with Termidor than those with Premise, Termidor was selected for use in all further eradication treatments. The results of April 2003 treatment and the June 2003 survey are shown in Fig. 3. Blue cones indicate active termite sites from the original survey (Fig. 1) before the April 2003 treatments, whereas red cones indicate live termite locations found during the June 2003 survey. Most yellow sectors (treated with Termidor) are free of red cones, indicating the efficacy of this treatment. Contour lines are used in addition to point locations to visualize areas of effective treatment. For the contour lines, two Kernel Density Estimation (KDE) maps were created based on termite locations of pre- and post treatment (i.e., blue and red cones). In a second step, the KDE values from the second layer (post-treatment) were subtracted from those of the first layer (pre-treatment). Thus, areas with higher contour values indicate areas of efficient chemical treatment, i.e., a high density of initial termite sightings, and a small number of follow-up sightings after the treatment. In general, areas treated with Termidor showed higher contour values than the nearby areas treated with Premise. One exception is sector 4 (see Fig. 1), where new colonies were detected after treatment, possibly due to undetected young colonies while doing the treatment and inadequate Termidor application (26.5 l). Sector 12, where all termites were eliminated through Termidor® as well, is not shown in Fig. 3 to keep a closer zoom on the remaining sectors. Because the majority of active sites (59) in mid-2003 occurred on a heavily wooded 4-hectare lot (Fig. 4), it was decided to first implement another more localized retreatment effort there. Before treating the wooded lot and using previous methods, 167 liters of Termidor® was applied on 12-15 September 2003 to 12 active sites outside of the wooded lot (yellow triangles in Fig. 4). A November-December 2003 survey added 18 additional live N. corniger sites to now total 77 sites on or within the fence line of the wooded lot. The lot was divided into six treatment sectors (yellow rectangles in Fig. 4) and on 18 December 2003, six teams applied 2,233 liters of Termidor to active sites (Fig. 4, blue circles). Figure 5 depicts the active locations of N. corniger from 2004 to 2010. In May 2010, the last treatment was applied for live termites within the constraints of reduced survey efforts. In November 2010 and May 2011, no more live termites were found in the original active tracts (Fig. 1) and the program was ended with the cautious hope that N. corniger was eradicated from south Florida. Although the decline in the number of yearly live sites is evident (Fig. 7), one may argue that the decline in active termite sites is partially caused by land cover change, i.e., transformation of some undeveloped lots into newly constructed buildings and parking areas. However, a comparison of historic aerial satellite images reveals that land cover change in the study area has been generally small over the past nine years. There are two areas with noticeable changes over time, however. The first area is a former container lot shown in an aerial image taken in early 2003 (Fig. 8a). Live termite sites for this area were observed in 2003 (green circles) and 2004 (yellow triangles), but not thereafter. An

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image from November 2005 (Fig. 8b) shows that most containers are still on the lot, whereas a month later, in December 2005, most containers were removed for subsequent development (Fig. 8c). The 2005 survey was conducted in June, when the containers were still in place. However, no termite infestation was observed in that survey, which indicates that the disappearance of termites can be attributed to the chemical treatment in years 2003 and 2004, respectively, and not to clearance of the lot. A similar situation can be observed in a previously wooded lot (Fig.8 d-f). Fig. 8d shows the corresponding satellite image from January 2003, where treatment locations are shown for 2003, 2004, and 2006. No infestations were identified thereafter, i.e., 2007 and later. As shown in Fig. 8e, the lot canopy is unchanged as of November 2007, whereas the area was cleared sometime within the following year (Fig. 8f). This, again, suggests that the eradication of termites occurred because of chemical treatment rather than through clearance of the lot. The total amount of Termidor® applied between April 2003 and May 2010 was 5,103 liters (6.33 kg AI) in addition to the Premise® applied in April 2003. Based on label directions, the amount of Termidor® applied as a preventative termite treatment for a 23 m2 structure built on a concrete slab would equal a minimum of 1,722 liters or 2.14 kg AI using a 0.125% suspension. Therefore, the entire area-wide elimination of N. corniger used the approximate Termidor® AI equivalent required to preventatively treat three 23 m2 buildings. Colony census Census results of the three N. corniger nests collected in 2004 are given in Table 2. The smallest nest contained 5,239 individuals including the primary king and queen. This colony had no exposed exterior fecal carton and was completely hidden within the termite-hollowed branch. This is precisely how Thorne & Haverty (2000) describe an incipient colony of an N. corniger colony in Panama: “the royal pair remains sequestered in their original copularium embedded in wood for several years as their incipient colony grows. Workers and soldiers in the young colony also remain within the wood, first eating local areas of the host tree and eventually traveling through the tree core, or perhaps building external galleries, to other food sources. Colonies also may be initiated in stumps or logs. When the colony population size approaches 10,000 termites, and when the first rains of the wet season create the right conditions, the colony blossoms into a rapidly constructed arboreal nest”. The incipient colony is the most problematic stage of nest development for field surveys because of its cryptic existence. The second nest contained ca. 76,460 individuals not counting the foraging population outside the nest but no kings or queens were found suggesting that this was a satellite nest of a larger primary colony as has been noted by others for N. corniger (e.g. Thorne 1982, Atkinson and Adams 1997). The largest nest (65 liters, 16.2 kg) contained over one million individuals, again not counting the foraging population outside the nest at time of collection. This nest contained two primary kings and 12 primary queens exemplifying the polygyny that is typical of a mature primary N. corniger nest (Thorne 1982). We suspect that before treatments began, several hundred million termites were established in the 2003 survey area.

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2011 Reemergence On July 11, 2011, RHS received a call from a pest control technician who reported that N. corniger was invading the building housing the International Game Fish Association (IGFA) museum in Dania Beach, Florida. An immediate inspection of the IGFA property revealed very dense infestations of mature N. corniger colonies throughout a ca. 1-hectare wetlands exhibit north of the building (yellow triangles in southeast corner of Fig. 9). This newly discovered infestation was reported to a local politician who contacted FLDACS, and who, in return, contacted RHS. Soon thereafter, FLDACS awarded a grant to FLREC to reinstitute the area-wide surveys that were discontinued in 2006. FLDACS also directed some field personnel to assist FLREC employees in survey efforts. These surveys revisited the original survey and treatment areas (Fig. 1) and areas up to 1 km away, primarily east and south of the original 2003 survey area border. By March 2012, an extensive new survey was completed and 74 active N. corniger sites were located for treatment. These sites were distributed among four zones encompassing the IGFA wetlands exhibit, an undeveloped wooded lot and its adjoining residential area, and two additional residential neighborhoods (Fig. 9). Although one of the residential sites adjoined an area that was treated in 2006 (Figs. 5, 6), none of the post-2010 sites were within the pre-2010 treated zones with one exception. A wood-framed couch, discarded in the wooden lot treated in December of 2003, was being attacked by a colony foraging from the newly discovered zone to the east of the lot. The lack of overlap between locations treated between 2003-2010 and the new active sites found in 2011-2012 validated the long-term efficacy of the original Termidor® treatments. In April 2012, FLDACS declared that the N. corniger infestations would be conducted deploying an “Incident Command System Response Team” from FLDACS DAES to conduct treatment efforts. The pest control industry and termiticide manufacturers were not allowed to participate in the FLDACS program. FLDACS was encouraged by FLREC to implement chemical treatments as soon as possible because alates had already been observed in some N. corniger nests as early as March 2012 (Fig. 2f). It was known in Florida and elsewhere that N. corniger synchronizes dispersal flights with the first drenching rains of the wet season which typically begins in May (Scheffrahn et al. 2002). On May 7, 2012, the response team was activated at a command post in Ft. Lauderdale, FL. The team was primarily staffed by DAES with assistance from the Division of Plant Industry (DPI). FLREC personnel were asked to assist the teamassisted in locating termite nests before treatments. At this meeting, DAES annouced a change in original termiticide application procedures used during 2003-2010. New application procedures were mandated because of FLDACS concern about potential public exposure to treated areas as more private parcels were involved in 2012 than in 2003. The application of Termidor® SC 0.125% was now to be limited only to nest injections accomplished by creating one or more holes in the nest carton in which the Termidor® liquid would be injected. A new formulation of fipronil, Termidor Dry® (0.5% AI), was additionally enlisted to dust the interior of N. corniger foraging tubes away from nests using the

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applicator included with the product. A label rate of 0.1g of Termidor Dry® (0.5 mg AI) was to be applied to foraging tubes every three m or “as practical”. On 8 May 2012, treatments were applied to the infested sites in Dania Beach (Fig. 9, yellow triangles). Treatments were halted early in the afternoon because of heavy thunderstorms. During twilight that evening, massive flights of N. corniger alates were observed emanating from the IGFA wetlands exhibit (Fig. 2g). The occurrence of a dispersal flight is significant because of the multi-year time lag before a colony constructs a carton nest and becomes detectable (Thorne and Haverty 2000). The new DAES-mandated application procedures jeopardize treatment efficacy because they eliminate the residual circum-foraging deposits of Termidor® applied in 2003-2010. Treating only nests and foraging tubes will intoxicate targeted colonies but will have little or no impact on nests (e.g. nest 1 above, Table 1) and active sites missed by visual inspection. With the original treatment strategy, overlooked populations of N. corniger would be drawn into the same preferred and treated foraging niches (e.g. abandoned foraging tubes and active feeding sites such as dead tree stumps, branches, cellulosic debris piles, wood implements, structural lumber, wooden pallets, etc.) used by the previously targeted population. Even partial contact with residual Termidor® deposits would result in mortality due to its extreme toxicity to termites (Remmen and Su 2005). Contact toxicity against termites has been demonstrated with deposits of as little as 10 ppb fipronil (Bagnères et al. 2009). Other active ingredients such as chlorantraniliprole (Scheffrahn and Scherer 2012) may prove to be options for residual applications in the future. All of the active tracts discovered in 2011-12 zones still contained live nests as of December 2012. The N. corniger experience in southeastern Florida brings to light a lack of a public policy for reacting to such inevitable urban pest invasions in the United States. With the exception of the New Orleans Mosquito and Termite Control Board (Guillot et al. 2010), no local, state, or federal agency has a clear mandate to react to an outbreak of non-agricultural or non-medical pests. A policy to deal with exotic structural pests might prevent or, at least, decelerate widespread economic losses in any such future event. ACKNOWLEDGEMENTS

The following Tree Termite Taskforce members were essential to field operations during this project: Mike Beckers and Norm Smith (Certified Pest Control Operators of Florida), Florida Pest Management Association, Fumigation Advisory Council (FAC), Russ Barnes, Ron Box (Hulett Environmental Services), Bart Bruni (Bestec Exterminators), Jon Depriest, Jeff Edwards (Dead Bug Edwards), Al Hoffer (Al Hoffer’s Pest Protection), Gil Livingston (PQ Termite Services), Lance McDonald (Terminix), Steve Niedzwiedzki (Univar USA), Gary Spero (Patriot Pest Control) Hugh Turner (Hugh Turner Pest Control), and Brian Van Dam (Fulton Pest Solutions); Chip Anderson, Eric Lenz, & Jorge Moreno (Bayer Environmental Sciences); Chris Key and Joe Hughes (BASF); Ellen Thoms and Rudolfo Subieta (Dow AgroSciences); Bayardo Herrera (Lesco); David Obrien, Kevin Nitsch, Laura Remmen, Matt Remmen (FLREC); Jude Plummer

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(MetroDade); Mary Peters, Charles Livio, Mike Orfanedes (Broward County Cooperative Extension Service); Charlie Clark, Steve Dwinell, Eric Reiss, Gary Tucker, George Moore, Kathleen Osgood, Mary Cohen, Mike Petrozzino, Neil Richmond, Wayne Gale, Eric Reiss, Richard Lucas (FLDACS DAES); Brian McElroy, Connie Riherd, Dan Phelps, Don Harris, Ed Burns, Jim Dowling, Jose Diaz, Mark Fagan, Micaelle Elicier, Randall Dean,Terry Smith, Fabian Philippoussi (FLDACS Division of Plant Industry); Lori Parrish (Broward County Board of Commissioners); Curtis Johnson (Broward County Aviation Department), Kurt Cobrick & Bill Gilmartin (Broward County Parks & Recreation Department); Mike Sheridan (City of Dania Beach Public Works and Utilities), Ivan Pato (City of Dania Beach City Manager); Brian and Kate Gaskill, (Banyan Bay Marina), Fred Elsen, Wilson Atkinson, Sully Sullivan (propery owners); Rob Kramer, Mike Myatt, Bob Greshman (IGFA).

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REFERENCES Atkinson L, Adams ES (1997) The origins and relatedness of multiple reproductives in colonies of the termite Nasutitermes corniger. Proc R Soc Lond (B) 264:1131–1136 Bagnères A-G, Pichon A, Hope J, Davis RW, Clément JL (2009) Contact versus feeding intoxication by fipronil in Reticulitermes termites (Isoptera: Rhinotermitidae): Laboratory evaluation of toxicity, uptake, clearance, and transfer among individuals. J Econ Entomol 102:347-356 Bahder BW, Scheffrahn RH, Krecek J, Keil C, Whitney-King S (2009). Termites (Isoptera: Kalotermitidae, Rhinotermitidae, Termitidae) of Ecuador. Ann de la Société entomologique de France 45: 529-536 Constantino, R (2002) The pest termites of South America: taxonomy, distribution and status. J Appl Entomol 126:355-365 Chouvenc T, Su N-Y, Grace JK (2011) Fifty years of attempted biological control of termites - Analysis of a failure. Biol Control 59:69-82 Guillot FS, Ring DR, Lax AR, Morgan A, Riegel C, Boykin D (2010) Area-wide management of the Formosan subterranean termite, Coptotermes formosanus Shiraki (Isoptera: Rhinotermitidae), in the New Orleans French Quarter. Sociobiology 55:311-338 Hochmair HH, Scheffrahn RH (2010) Spatial association of marine dockage with land-borne infestations of invasive termites (Isoptera: Rhinotermitidae: Coptotermes) in urban south Florida. J Econ Entomol 103:1338-1346 Merritt NRC, Starr CK (2010) Comparative nesting habits and colony composition of three arboreal termites (Isoptera: Termitidae) in Trinidad & Tobago, West Indies. Sociobiology 56:611-622 Peel MC, Finlayson BL, McMahon TA (2007) Updated world map of the Köppen-Geiger climate classification. Hydrology and Earth System Sciences Disc 4:439-473 Remmen LN, Su N-Y. (2005) Tunneling and mortality of eastern and Formosan subterranean termites (Isoptera: Rhinotermitidae) in sand treated with thiamethoxam or fipronil. J Econ Entomol 98:906-910 Roisin Yves, Pasteels JM (1985) A new Nasutitermes species from New Guinea (Isoptera: Termitidae). Indo-Malayan Zool 2:325-330 Scheffrahn RH, Crowe W (2011) Ship-borne termite (Isoptera) border interceptions in Australia and onboard infestations in Florida, 1986–2009. Florida Entomol 94:57-63

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Scheffrahn RH, Su N-Y (1994) Keys to soldier and winged adult termites (Isoptera) of Florida. Florida Entomol 77:460-474 Scheffrahn RH, Cabrera BJ, Kern Jr WH, Su N-Y (2002) Nasutitermes costalis (Isoptera: Termitidae) in Florida: first record of a non-endemic establishment by a higher termite. Florida Entomol 85:273-275 Scheffrahn RH, Jones SC, Krecek J, Chase JA, Mangold JR, Su N-Y (2003) Taxonomy, distribution, and notes on the termites (Isoptera: Kalotermitidae, Rhinotermitidae, and Termitidae) of Puerto Rico and the U.S. Virgin Islands. Ann Entomol Soc America 96:181-201 Scheffrahn RH, Krecek J, Maharajh B, Chase JA, Mangold JR, Moreno J, Bayardo H (2005) Survey of the termites (Isoptera: Kalotermitidae, Rhinotermitidae, Termitidae) of Nicaragua. Florida Entomologist 88:549-552 Scheffrahn RH, Krecek J, Chase JA, Maharajh B, Mangold JR (2006) Taxonomy, biogeography, and notes on the termites (Isoptera: Kalotermitidae, Rhinotermitidae, Termitidae) of the Bahamas and Turks and Caicos Islands. Ann Entomol Soc America 99:463-486 Scheffrahn RH, Scherer CW (2012) Efficacy of Altriset® on the tropical arboreal termite, Nasutitermes corniger (Isoptera: Termitidae: Nasutitermitinae). Florida Entomologist 96 (in press) Scheffrahn RH, Su N-Y, Diehl B. (1990). Native, introduced, and structure-infesting termites of the Turks and Caicos Islands, BWI (Isoptera: Kalotermitidae, Rhinotermitidae, Termitidae). Florida Entomologist 73:622-627 Scheffrahn RH, Krecek J, Szalanski AL, Austin JW (2005) Synonymy of the neotropical arboreal termites, Nasutitermes corniger and N. costalis (Isoptera: Termitidae), with evidence from morphology, genetics, and biogeography. Ann Entomol Soc America 98:273-281 Thorne BL, Haverty MI (2000) Nest growth and survivorship in three species of neotropical Nasutitermes (Isoptera: Termitidae). Environ Entomol 29:256-264 Thorne BL (1982) Polygyny in termites: Multiple primary queens in colonies of Nasutitermes corniger (Motschulsky) (Isoptera: Termitidae). Insectes Soc 29:102-107 Torales GJ (2002) Termites as structural pests in Argentina. Sociobiology 40:191-206

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Table 1. Volume of termiticide applied on 23 April 2003 in Dania Beach and live N. corniger infestation sites before and after the 23 April 2003 treatment (see text for statistical data). Live sites Live sitesSector Chemical Liters Pretreat Posttreat

1 Premise 0.1% 163 17 42 Termidor 0.125% 378 24 03 Premise 0.1% 867 40 394 Termidor 0.125% 26 16 65 Premise 0.1% 38 15 156 Termidor 0.125% 859 31 17 Premise 0.1% 45 13 38 Premise 0.1% 159 18 39 Termidor 0.125% 220 19 0

10 Termidor 0.125% 5 5 011 Termidor 0.125% 8 1 012 Termidor 0.125% 8 5 0

Total Premise 0.1% 1,272 103 64Termidor 0.125% 1,504 101 7

Table 2. Populations of Nasutitermes corniger nests, Dania Beach, Florida.

No. by Caste3

E L W N A PS S K Q Total

Colony 1a 994 1559 2339 1 0 101 243 1 1 5239

Colony 2b 10 50 61230 870 0 1520 12780 0 0 76460

Colony 3c 156200 353200 521900 6800 0 19200 74700 2 16 1132000

a Total count; incipient colony between branch and soil; about 6cm and 60 cm in area. b Estimate from subsample count c E = eggs, L = larvae, W = workers, N = nymphs, A = alates, PS = presoldiers, S = soldiers, K = kings, Q = queens.

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Fig. 1 Results of N. corniger field survey from 7 April 2003, in Dania Beach, FL with 12 designated sectors for chemical treatments (background image from Google).

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Fig. 2 Nasutitermes corniger habitus, Dania Beach, Florida. a: Damage to a wooden door jamb. b: Foraging tubes on house. c: Foragers emerging from indoor molding. d: Nest under leaf litter against chain-link fence. d: Nest at base of tree. e: nest containing alates one month before flight. f: Alates emerging for dispersal flight from nest at base of grass tussock.

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Fig. 3 Visualization of N. corniger locations before (blue) and after (red) 23 April 2003 treatment, with contour lines indicating effectiveness of chemical treatment in different sectors. Yellow sectors were treated with Termidor and green sectors were treated with Premise.

Fig. 4 Nasutitermes corniger treatment sites on 12-15 Sep 2003 outside wooded lot (triangles), and on 18 Dec 2003 in wooded lot (circles) in Dania Beach, FL (background image from Google).

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Fig. 5 Annual N. corniger treatment sites for the years 2004-2010 in Dania Beach, FL (background image from Google).

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Fig. 6 Locations of known active N. corniger sites between April 2003 and June 2011 in Dania Beach, FL (background images from Google).

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Fig. 7 Number of active N. corniger sites by year in Dania Beach, FL

(a) 28 Jan 2003 (b) 11 Nov 2005 (c) 30 Dec 2005

(d) 28 Jan 2003 (e) 25 Nov 2007 (f) 30 Dec 2008 Fig. 8 Change of land cover at two active N. corniger study areas in Dania Beach, FL (background images from Google).

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2003 2004 2005 2006 2007 2008 2009 2010 2011

Treated Termite Sites

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Fig. 9 Original April 2003 170-hectare survey area in Dania Beach, FL. Area surveyed approximately biannually until June 2006. Red circles are N. corniger active sites detected from April 2003 to December 2010 that were treated. Yellow triangles are active sites detected from July 2011 to April 2012.