A comparison between above-ground bait stations and experimental caulk baits for elimination of field colonies of the Asian subterranean termite, Coptotermes gestroi (Blattodea: Rhinotermitidae).

Survey stakes and in-ground (IG) stations have failed to intercept underground tunnels of the Asian subterranean termite, Coptotermes gestroi (Wasmann), in southeastern Florida, rendering nearly useless the IG bait station that is the main tool of commercial baiting systems. When placed over an active infestation, above-ground (AG) bait stations were readily fed on by C. gestroi, resulting in colony elimination. A new experimental type of AG bait application being studied is an injectable caulk bait. In this study, we compared the efficacy of AG bait stations and an experimental AG caulk bait against field colonies of C. gestroi. Following the applications of AG bait stations at 3 sites with infested houses and trees, C. gestroi colonies were eliminated 7.1-28.1 wk later. The experimental AG caulk bait was applied at 7 sites including 4 houses, 2 office buildings, and 1 docked sailboat. Colony elimination times at caulk bait sites ranged from 3.1 to 13.6 wk, but there was no statistical difference in mean colony elimination time between AG station bait sites (17.4 ± 10.5 wk) and AG caulk bait sites (8.4 ± 3.8 wk). This study showed that AG caulk bait applications were equally efficacious in eliminating existing C. gestroi in structures as AG bait stations and will offer another tool for termite control professionals to better manage subterranean termites including C. gestroi.

Elimination of structural and tree infestations of the Asian subterranean termite, Coptotermes gestroi (Wasmann) (Blattodea: Rhinotermitidae) with noviflumuron baits in above-ground stations.

The traditional stake survey and in-ground (IG) monitoring stations have been ineffective in aggregating the Asian subterranean termite, Coptotermes gestroi (Wasmann) in southeastern Florida. In this study, we used both IG and above-ground (AG) Sentricon stations to monitor and bait C. gestroi, and as expected, none of the 83 IG stations was intercepted. Despite this, AG bait stations with 0.5% noviflumuron were successfully used to eliminate C. gestroi colonies. From 2 field experiments, the mean colony elimination time (±SD) using AG baits were 6.4 ± 3.8 wk (n = 4) and 8.0 ± 2.1 wk (n = 12), respectively. Such results were compatible with baiting studies against field colonies of C. gestroi elsewhere, that is, 4-9 wk. The successful rates in monitoring and baiting of C. gestroi with IG stations in other regions also varied, which may be due to the variabilities in tunnel geometry of this species in different environments. In areas with established C. gestroi populations, routine inspection for signs of activity in structures and surrounding trees can be a critical component for pest control providers for early detection of infestation and colony elimination with AG bait stations.

Age-based spatial distribution of workers is resilient to worker loss in a subterranean termite.

Elaborate task allocation is key to the ecological success of eusocial insects. Termite colonies are known for exhibiting age polyethism, with older instars more likely to depart the reproductive center to access food. However, it remains unknown how termites retain this spatial structure against external disturbances. Here we show that a subterranean termite Coptotermes formosanus Shiraki combines age polyethism and behavioral flexibility to maintain a constant worker proportion at the food area. Since this termite inhabits multiple wood pieces by connecting them through underground tunnels, disastrous colony splitting events can result in the loss of colony members. We simulated this via weekly removal of all individuals at the food area. Our results showed that termites maintained a worker proportion of ~ 20% at the food area regardless of changes in total colony size and demographic composition, where younger workers replaced food acquisition functions to maintain a constant worker proportion at the food area. Food consumption analysis revealed that the per-capita food consumption rate decreased with younger workers, but the colony did not compensate for the deficiency by increasing the proportion of workers at the feeding site. These results suggest that termite colonies prioritize risk management of colony fragmentation while maintaining suitable food acquisition efficiency with the next available workers in the colony, highlighting the importance of task allocation for colony resiliency under fluctuating environments.

Soil organic matter is essential for colony growth in subterranean termites.

Intrinsic dinitrogen (N2) fixation by diazotrophic bacteria in termite hindguts has been considered an important pathway for nitrogen acquisition in termites. However, studies that supported this claim focused on measuring instant N2 fixation rates and failed to address their relationship with termite colony growth and reproduction over time. We here argue that not all wood-feeding termites rely on symbiotic diazotrophic bacteria for colony growth. The present study looks at dietary nitrogen acquisition in a subterranean termite (Rhinotermitidae, Coptotermes). Young termite colonies reared with wood and nitrogen-rich organic soil developed faster, compared to those reared on wood and inorganic sand. More critically, further colony development was arrested if access to organic soil was removed. In addition, no difference of relative nitrogenase expression rates was found when comparing the hindguts of termites reared between the two conditions. We therefore propose that subterranean termite (Rhinotermitidae) colony growth is no longer restricted to metabolically expensive intrinsic N2 fixation, as the relationship between diazotrophic bacteria and subterranean termites may primarily be trophic rather than symbiotic. Such reliance of Rhinotermitidae on soil microbial decomposition activity for optimal colony growth may also have had a critical mechanistic role in the initial emergence of Termitidae.

Social Behavior, Ovary Size, and Population of Origin Influence Cuticular Hydrocarbons in the Orchid Bee Euglossa dilemma.

AbstractCuticular hydrocarbons (CHCs) are waxy compounds on the surface of insects that prevent desiccation and frequently serve as chemical signals mediating social and mating behaviors. Although their function in eusocial species has been heavily investigated, little is known about the evolution of CHC-based communication in species with simpler forms of social organization lacking specialized castes. Here we investigate factors shaping CHC variation in the orchid bee Euglossa dilemma, which forms casteless social groups of two to three individuals. We first assess geographic variation, examining CHC profiles of males and females from three populations. We also consider CHC variation in the sister species, Euglossa viridissima, which occurs sympatrically with one population of E. dilemma. Next, we consider variation associated with female behavioral phases, to test the hypothesis that CHCs reflect ovary size and social dominance. We uncover a striking CHC polymorphism in E. dilemma spanning populations. In addition, we identify a separate set of CHCs that correlate with ovary size, social dominance, and expression of genes associated with social behavior, suggesting that CHCs convey reproductive and social information in E. dilemma. Together, our results reveal complex patterns of variation in which a subset of CHCs reflect the social and reproductive status of nestmates.

Fused Colonies of the Formosan Subterranean Termite (Blattodea: Rhinotermitidae) for Laboratory Experiments.

Laboratory studies of Coptotermes formosanus Shiraki (Blattodea: Rhinotermitidae) often employ the use of field-collected foraging populations of individuals as defined colonies. The biological relevance of this practice is often called into question, because these colonies lack a full composition of reproductive castes and brood, which may have physiological and behavioral consequences. Rearing intact laboratory colonies can be done; however, it is time-consuming and labor-intensive. The artificial fusion of field-collected foraging populations with a young, laboratory-reared incipient colony may provide whole, intact colonies for laboratory research. The current study measures survivorship of fused colonies using laboratory-reared complete incipient colonies ranging in age from 0 to 5 mo, fused with 100 workers and 10 soldiers from field-collected populations of different colonial origin. Results indicate that 60% of colony fusion was successful when the incipient colony introduced is 5 mo of age. This method of colony fusion will provide researchers with intact colonies using minimal resources.

Field Evaluations of Fluid Baits against Colonies of the Formosan Subterranean Termite (Blattodea:Rhinotermitidae).

Fluid baits comprised of 10% dry medium (impregnated with 0.5% hexaflumuron) and 90% methylcel solution were injected into foraging galleries of the Formosan subterranean termite, Coptotermes formosanus Shiraki (Blattodea: Rhinotermitidae), in trees and buildings. Elimination times at two of the six baited sites were longer than expected, possibly due to the application disturbance or the presence of an old and declining colony, but fluid bait treatments eliminated all baited colonies. Fluid baits were injected through drill holes behind infested wood surface, and bypassed the difficulties for installing above-ground (AG) bait stations on uneven surface or at tight corners. Because a large quantity of fluid baits could be applied at once, termite control professionals may be able to avoid revisiting the site to add more baits. The absence of unsightly AG station in a house will also make fluid baits more acceptable to homeowners than AG baits. Currently, local applications of liquid termiticides are common practices because termite control professionals are often urged by homeowners to take remedial actions when termites are found in a house, but such treatments only drive termites away from the active loci without affecting the termite colony(s). Remedial treatments with fluid baits would result in immediate bait consumption and shorten colony elimination time with or without the use of in-ground baiting systems. Introduction of fluid baits will add another tool to the termite control industry and will make it more plausible for the homeowners to accept baiting technology.

Parental Nitrogen Transfer and Apparent Absence of N2 Fixation during Colony Foundation in Coptotermes formosanus Shiraki.

Colony foundation and early growth is a critical period in the life-cycle of a termite colony, as the initial family unit is resource limited. One such resource is nitrogen, which is essential for initial colony growth. This study examined the whole-colony nitrogen inventory during foundation and early growth of Coptotermes formosanus Shiraki colonies. It was hypothesized that termite colonies would go through an initial period of parental investment, representing a transfer of nitrogen to the first brood, and that once a functional worker caste was present, further provisioning in the form of intrinsic N2 fixation would occur. Our results showed that, when in nitrogen-poor rearing conditions, the king and queen initially transferred half of their nitrogen reserves to their first brood. However, the total nitrogen content in colonies did not increase over a 12 month period, despite the presence of functional workers. Furthermore, colonies did not increase their biomass beyond the initial parental investment. Together, these results imply that nitrogen acquisition in incipient C. formosanus colonies relies on environmental or dietary sources, rather than the putative fixation through symbiotic diazotrophs.

Flight Phenology of Two Coptotermes Species (Isoptera: Rhinotermitidae) in Southeastern Florida.

The dispersal flight activity ("swarming") of two invasive subterranean termite species, Coptotermes gestroi (Wasmann) and Coptotermes formosanus Shiraki, was monitored in metropolitan southeastern Florida, where both species are now sympatric and major structural pests. Historical records of alates collected in the area showed that the two species have distinct peaks of flight activity, from mid-February to late April for C. gestroi, and from early April to late June for C. formosanus. However, an overlap of the two dispersal flight seasons has been observed since at least 2005. The daily monitoring of dispersal flight events in southeastern Florida in 2014, 2015, and 2016 confirmed that simultaneous flights occurred several times each year. In addition, environmental conditions for favorable flights were identified, and it was established that low temperature was the primary factor inhibiting both species from dispersal flights, while all other factors had little impact on the occurrence of major dispersal flight events. However, both species shared similar temperature requirements for favorable dispersal flight conditions despite distinct peaks of activity over time. The analysis of sex ratios and average weights of the alates suggests that intrinsic colony factors are important for the timing of the maturation of alates, and that once a cohort of individuals is ready to disperse, a flight may occur as soon as the environmental conditions are favorable.

Foraging Distance and Population Size of Juvenile Colonies of the Formosan Subterranean Termite (Isoptera: Rhinotermitidae) in Laboratory Extended Arenas.

The relationship between colony size and foraging distance was examined in extended foraging arenas with juvenile colonies of the Formosan subterranean termite, Coptotermes formosanus Shiraki. Our results showed that as long as royal pairs are present, larger colonies foraged at longer distances, and the oldest workers distributed farther away from the central nest. The results agree with the scaling model that predicts a large foraging range for animals of larger body size. An analysis of published data from population survey studies and field trials of bait toxicants showed that field colonies of the eastern subterranean termite, Reticulitermes flavipes (Kollar), follow the scaling model, while C. formosanus colonies were inconsistent with the model prediction. Reasons for the inconsistency with field data of C. formosanus are discussed.

Reinvasion Dynamics of Subterranean Termites (Isoptera: Rhinotermitidae) Following the Elimination of All Detectable Colonies in a Large Area.

Following the elimination of all detectable termite colonies in the 32-acre Louis Armstrong Park in New Orleans, LA, in 2002­2003, termite activity was monitored by using 808 Sentricon stations. Between January 2004 and July 2005, termites were found in 8­11 stations. In August 2005, the Park was flooded by Hurricane Katrina, but termites remained active. Post-Katrina termite activity levels of 16­21 stations were recorded throughout 2006, and in October 2007, the activity drastically increased to 43 stations. This rapid increase of termite activity continued into 2008, and a total of 94 stations harbored termite activities by July 2008. Termite activity peaked at 109 stations in September 2008 and then leveled down to 64 stations in March 2009. Termite activity in the Park between 2004 and 2009 was described by a Sigmoid model with a carrying capacity of 76 stations, and a Sigmoid mid-point of 1,202 d. In April 2009, a total of 14 colonies of Coptotermes formosanus Shiraki and one colony of Reticulitermes flavipes (Kollar) were delineated by using microsatellite genotyping and mark­recapture protocol. Of the 15 colonies, eight near the Park border probably originated from existing colonies from outside, and seven C. formosanus colonies found inside the Park were probably initiated by alate pairs. Our results showed that, if surrounded by high population pressure of termites and no control measures are applied, an area cleared of termite populations by baits can be completely re-populated by termites from outside in 53 mo.

Dispersal Flights of the Formosan Subterranean Termite (Isoptera: Rhinotermitidae).

The Formosan subterranean termite, Coptotermes formosanus Shiraki, is a pest of major economic concern. This termite is particularly known for its tendency to establish populations in nonendemic areas via maritime vessels as well as human-aided transport of infested materials. The natural spread of this species after new introductions occurs in part by dispersal flights originating from mature colonies. Dispersal flight activity is also the primary variable for the evaluation of area-wide management programs. Few studies exist describing the dynamics and distribution of a typical dispersal flight for this species. The present study used data collected by mark-recapture of C. formosanus alates over 12 individual evenings of dispersal flights in the New Orleans French Quarter. In this study, we found that for one selected flight dispersal location, which was not affected by a high density of trap locations nearby, alates flew on average 621 m from their parent colony. A new record of a 1,300-m dispersal flight was recorded. Spatial analysis showed that neither wind nor light affected the direction of flight, which may, however, be attributed to scarce light and wind measurements in the study region.

Reinvasion and colony expansion of Coptotermes formosanus (Isoptera: Rhinotermitidae) after areawide elimination.

The population recovery of Coptotermes formosanus Shiraki (Isoptera: Rhinotermitidae) colonies were monitored after an areawide elimination of all detectable colonies from September 2003 to August 2005 in Louis Armstrong Park, New Orleans, LA. Six colonies reinvaded the vacant niche created by the full elimination. These colonies expanded their territories throughout the study period. This represented 43% of the original number of colonies present in the park before the elimination. To determine the mode of the reinvasion, nuptial pair establishment was monitored during the C. formosanus dispersal flight seasons. Nuptial pairs were discovered up to 1 yr after the elimination. Morphological and genetic data were collected from field colonies before the full elimination in 2002 and again in 2005 after the reinvasion of these territories by new colonies. These data were used to estimate the relative age of reinvading colonies as compared with their predecessors. It is proposed that the first three reinvading colonies detected were smaller colonies that were undetectable before the full elimination, or were older, established colonies present outside of the park, that expanded their foraging territories into the park in the absence of competition from the eliminated populations. The subsequent three colonies to reinvade seemed to be small colonies founded during or just before the study period by an imago pair after a dispersal flight into the park from outlying areas. The implications of this study on subterranean termite areawide integrated pest management strategies are discussed.