Escape Behavior of the Formosan Subterranean Termite (Isoptera: Rhinotermitidae) in Response to Disturbance

Disturbances to monitors and baits have often been noted as a confounding factor for the success of termite baiting systems. We studied escape behavior of Coptotermes formosanus Shiraki to a food source disturbance in the laboratory. Four peripheral chambers connected to the central food chamber through tubes served as escape routes from which escape patterns, escape and return times and differences in soldier and worker responses were recorded. We used two sand conditions: moist and dry to test whether favorable versus unfavorable conditions change escape behavior. The post disturbance responses were video recorded for 1 h. C. formosanus quickly escaped from the central chamber when disturbed. The time taken to reach the maximum number of termites that escaped ranged from 20 s to 210 s followed by a gradual return to the disturbed chamber. The majority of the escapees returned to the disturbed area within a few minutes after evacuation in both dry and wet sand conditions. A significantly higher percentage of termites escaped in wet sand arenas (43%) than in the dry sand arenas (31%). Our experiments demonstrate that C. formosanus is highly unlikely to abandon bait stations for extensive periods of time as a result of mechanical disturbances regardless of habitat quality.     

Distribution of Formosan subterranean termite (Isoptera: Rhinotermitidae) in Mississippi

An extensive monitoring and survey program in Mississippi was conducted from 2000 to 2004 to investigate the distribution of the Formosan subterranean termite, Coptotermes formosanus Shiraki (Isoptera: Rhinotermitidae). Seventy-two towns from 22 counties in southern Mississippi were monitored with a total of 3914 traps that catch alates during the swarming season. In addition, 259 licensed pest management professionals in Mississippi were surveyed to determine the locations of termite infestations treated. The alates of C. formosanus were recovered in 12 counties with light traps, and termite infestations were documented in an additional 13 from data collected in the termite survey. Infestations of C. formosanus have been documented in urban, urban cluster, rural, and forested areas of Mississippi. However, the distribution in mean total capture of alates for 4 yr differed significantly among the four ecological areas with the highest percentage in forested areas (31%) and the lowest percentage in urban cluster areas (17%). Most of the infestations of C. formosanus were geographically distributed along the coastal areas of southern Mississippi from Gulfport to Pascagoula. The greatest total number of alates captured in light traps was documented in Pearl River County. Mass swarming of C. formosanus occurred primarily in May or June, depending on weather conditions. The number of documented counties with the evidence of large and widely dispersed swarms of C. formosanus in different ecological areas, and the increase in total annual alate captures from 2000 to 2003, suggest that this invasive termite species is now firmly established in Mississippi.     

Trail Communication During Foraging and Recruitment in the Subterranean Termite Reticulitermes santonensis De Feytaud (Isoptera, Rhinotermitidae)

The search for food in the French subterranean termite Reticulitermes santonensis De Feytaud is organized in part by chemical trails laid with the secretion of their abdominal sternal gland. Trail-laying and -following behavior of R. santonensis was investigated in bioassays. During foraging for food termites walk slowly (on average, 2.3 mm/s) and lay a dotted trail by dabbing the abdomen at intervals on the ground. When food is discovered they return at a quick pace (on average, 8.9 mm/s) to the nest, laying a trail for recruiting nestmates to the food source. While laying this recruitment trail the workers drag the abdomen continuously on the ground. The recruitment trail is highly attractive: it is followed within a few seconds, by more nestmates, and at a quicker pace (on average, 6.4 mm/s) than foraging trails (on average, 2.9 mm/s). The difference between foraging and recruitment trails in R. santonensis could be attributed to different quantities of trail pheromone. A caste-specific difference in trail pheromone thresholds, with workers of R. santonensis being more sensitive to trails than soldiers, was also documented: soldiers respond only to trails with a high concentration of trail pheromone.     

Formosan subterranean termite swarming behavior and alate sex-ratio (Isoptera: Rhinotermitidae)

Summary The Formosan subterranean termite (Coptotermes formosanus Shiraki) was first sighted in the cities of New Orleans and Lake Charles, Louisiana, in 1966. Its distribution in the historic French Quarter of New Orleans 23 years later (1989) and in 1991 and 1992 is reported here using counts of alates from light-traps. Light-trappings revealed a Quarter-wide distribution of the termite.C. formosanus shows an extended pause in flight behavior in the middle of the two-month flight season that is predictable and independent of weather. A colony-level mechanism, namely, developmental inhibition of nymphs by alates, may regulate this intermission in flight. The sex-ratio of light-trapped alates varies over time, but is generally female-biased, especially at the height of the flight season.     

Effect of Average Worker Size on Tunneling Behavior of Formosan Subterranean Termite Colonies

The effect of a colony's average worker size on tunneling behavior of Coptotermes formosanus Shiraki (Isoptera: Rhinotermitidae) in laboratory foraging arenas was investigated. Large groups of foragers from four colonies were added to arenas and allowed to tunnel for a period of 22 days. Various aspects of the resulting tunnel galleries were analyzed using ANOVA and regression analysis. After the exclusion of one colony due to high mortality, the only parameters shown to differ significantly were tunnel width and tunnel segmentation. Larger workers tended to construct less segmented galleries with wider tunnels. Tunnel length was positively correlated to size on the first 2 days, but the relationship reversed by day 4. Tunneling on day 1 was positively correlated to the discovery of food items. Our results indicate that a foraging group's average worker size can affect tunnel pattern and that groups composed of larger workers are better tunnelers, but groups of smaller sized workers are able to make up the difference over time. Possible explanations for size-related variation in tunneling behavior are discussed.     

Behavioral Response of Two Subterranean Termites (Isoptera: Rhinotermitidae) to Vibrational Stimuli

Behavioral responses to vibrational stimuli were examined in two subterranean termite species, Coptotermes formosanus Shiraki and Reticulitermes flavipes (Kollar). Termites habituated to vibrational stimulation when in social groups, but failed to do so individually, indicating that habituation is a collective action. In assays on termite groups, both species demonstrated a similar pattern of behavioral responses to vibrational stimuli: evanescent cessation of activity and movement, followed by withdrawal from the vibration source. Groups of both species then gradually moved back toward the vibration source as a consequence of continuous exposure. However, it took a significantly shorter period for C. formosanus to return (57 s) to the test arena and to resume (80 s) normal foraging activities in the test arena compared with R. flavipes, which took 97 and 227 s, respectively, when exposed to the vibration frequency of 120 bmpm. High vibration frequency (240 bmpm) increased the time required to return (C. formosanus, 80 s; R. flavipes, 153 s) and to resume regular locomotion (C. formosanus, 186 s; R. flavipes, 263 s). Our experiments demonstrate that workers play a crucial part in adjusting groups of termites to distressful vibrations. Soldiers of R. flavipes demonstrated similar behavioral responses as workers, however, C. formosanus soldiers exhibited a transient positive response before withdrawal. An erratum to this article is available at .     

Division of Labour among Workers in the Termite,Reticulitermes fukienensis (Isoptera: Rhinotermitidae)

To investigate division of labour in Reticulitermes fukienensis, worker and larval subgroups were established in laboratory cultures and task performance patterns of different-sized workers and larvae were recorded. Five categories of subgroups were established based on head width differences: small larvae (SL); large larvae (LL); small workers (SW); medium workers (MW); and large workers (LW). Subgroups were compared for their ability to carry out the following tasks: tunnel construction; covered gallery construction; gallery repair; and feeding. Larval subgroups were found to carry out none of the tasks investigated. They did not feed or build covered galleries or underground tunnels. Temporal polyethism occurred among workers of different size groups. Covered foraging galleries were exclusively built and tunnels were predominantly built by older workers (i.e. MW and especially LW). Remarkably, most SW were apparently unable to burrow into the soil, something that all LW did within 3 d and 95% of MW did within 5 d. Surprisingly, LW ate 8.5 and 65.5 times as much food (i.e. filter paper) as MW and SW, respectively, although their average body weights were, respectively, only approximately two and four times that of MW and SW. MW carried out all four tasks that LW performed, although MW were typically less efficient. This contrasts with the hypothesis that different instars of termites should carry out different non-overlapping tasks. Large workers carried out most of all tasks compared with other worker sizes. The only exception was for gallery repair where repair rate by LW did not differ significantly from MW. The predominance of tasks carried out by larger (i.e. older) workers indicates a possible new pattern for division of labour in these lower termites. This contrasts with the pattern of organization of division of labour in the social Hymenoptera.     

Dual origin of gut proteases in Formosan subterranean termites (Coptotermes formosanus Shiraki) (Isoptera: Rhinotermitidae)

Cellulose digestion in lower termites, mediated by carbohydrases originating from both termite and endosymbionts, is well characterized. In contrast, limited information exists on gut proteases of lower termites, their origins and roles in termite nutrition. The objective of this study was to characterize gut proteases of the Formosan subterranean termite (Coptotermes formosanus Shiraki) (Isoptera: Rhinotermitidae). The protease activity of extracts from gut tissues (fore-, mid- and hindgut) and protozoa isolated from hindguts of termite workers was quantified using hide powder azure as a substrate and further characterized by zymography with gelatin SDS-PAGE. Midgut extracts showed the highest protease activity followed by the protozoa extracts. High level of protease activity was also detected in protozoa culture supernatants after 24 h incubation. Incubation of gut and protozoa extracts with class-specific protease inhibitors revealed that most of the proteases were serine proteases. All proteolytic bands identified after gelatin SDS-PAGE were also inhibited by serine protease inhibitors. Finally, incubation with chromogenic substrates indicated that extracts from fore- and hindgut tissues possessed proteases with almost exclusively trypsin-like activity while both midgut and protozoa extracts possessed proteases with trypsin-like and subtilisin/chymotrypsin-like activities. However, protozoa proteases were distinct from midgut proteases (with different molecular mass). Our results suggest that the Formosan subterranean termite not only produces endogenous proteases in its gut tissues, but also possesses proteases originating from its protozoan symbionts.     

Colony breeding system influences cuticular bacterial load of Formosan subterranean termite (Isoptera: Rhinotermitidae) workers

The goal of this study was to test whether the breeding system and/or the degree of inbreeding of field colonies of the Formosan subterranean termite, Coptotermes formosanus, Shiraki (Isoptera: Rhinotermitidae) influences bacterial load on the cuticle of foraging workers. We enumerated bacterial load on the cuticle of groups of workers foraging in 20 inground monitoring stations surrounding the French Market in New Orleans, LA, and identified bacteria species using 16S rRNA gene sequencing. We used microsatellite genotyping to assign the 20 worker groups to seven simple family colonies (headed by a single pair of reproductives) and four extended family colonies (headed by multiple inbreeding reproductives) with a wide range of degrees of inbreeding. Workers from extended family colonies had a higher bacterial load than those from simple family colonies; however, bacterial load was not significantly correlated to the degree of inbreeding, possibly because of confounding factors in colony life history, such as age and/or size of colonies. Colonies with high bacterial load did not have a higher proportion of entomopathogens, and thus, bacterial load is not necessarily an indicator for disease risk. The majority of bacteria cultured from the cuticle of termites were soil bacteria with no known pathology against termites.     

Evaluation of treatment success and patterns of reinfestation of the Formosan subterranean termite (Isoptera: Rhinotermitidae)

Spread of the Formosan subterranean termite, Coptotermes fornwsanus Shiraki (Isoptera: Rhinotermitidae), is connected with the transport of infested wood, in particular, railroad ties. Therefore, efficient treatment of infestations, especially along railroads, is imperative to prevent further termite damage and spread. Evaluation of treatment success hinges on the ability to assign infestation sites to colonies and compare colony identity before and after treatment. Because colonies of the Formosan subterranean termite can be headed by a pair of reproductives (simple families) or by multiple reproductives (extended families), the question arises whether the breeding system of a colony influences treatment success and whether treatment of an area might have an impact on the breeding system. We used microsatellite genotyping to compare colony affiliation and breeding systems of Formosan termites infesting the Riverfront Railroad, New Orleans, LA, before and after treatment with 0.5% noviflumuron. Before treatment, four colonies were simple families, and 11 colonies were extended families. A year after treatment began, all treated colonies had vanished and did not reappear during this study. One colony from an untreated monitoring station moved into a nearby station after treatment. Colonies that were detected after treatment consisted of 12 simple families and six extended families; extended families found after treatment contained a higher number of reproductive neotenics than the extended families found before treatment. Extended families were more likely than simple families to move into inground stations that had been previously occupied by termite colonies.     

Effect of tolerance to insecticides on substrate penetration by Formosan subterranean termites (Isoptera: Rhinotermitidae)

Seven selected insecticides were applied to three substrates and bioassayed for barrier efficacy and toxicity against the Formosan subterranean termite, Coptotermes formosanus Shiraki, originating from colonies that differed in their tolerance to the insecticides. A profound substrate effect was seen with all insecticides tested. Sand was the most difficult substrate to penetrate and caused faster and greater mortality of C. formosanus. Increased tolerance in C. formosanus was accompanied by a decrease in penetration of treated substrata with chlordane, methoxychlor, chlorpyrifos, and deltamethrin. The opposite was true in the case of permethrin and bendiocarb. More tolerance C. formosanus displayed decreased mortality in the presence of chlorpyrifos, deltamethrin, bendiocarb, and fipronil. The opposite was true for permethrin.     

四种白蚁的线粒体DNA限制性片段长度多态性研究(等翅目:鼻白蚁科,木白蚁科)

用６种限制性内切酶分析了４种白蚁的线粒体 Ｄ Ｎ Ａ 限制性片段长度多态性,根据限制性片段差异计算了４种白蚁之间的遗传距离,利用 Ｕ Ｐ Ｇ Ｍ Ａ 聚类分析法构建了分子聚类图。结果表明：尖唇异白蚁与散白蚁属关系很近,聚类分析结果显示应将其归于散白蚊属     

Subterranean termites (Isoptera: Rhinotermitidae): Exploitation of equivalent food resources with different forms of placement

Abstract Coptotermes gestroi and Heterotermes tenuis have been described as important urban pests in Brazil. The establishment of control technologies using baits that consider the social behavior of termites requires a better knowledge of their foraging behavior. Thus, this study analyzed the feeding behavior of these species with three different forms of food placement: food on the surface of a substrate and food either partially or completely buried in the substrate. Experimental arenas were composed of a central chamber connected to three food chambers. Each central chamber contained 550 foragers of C. gestroi or 517 foragers of H. tenuis. Blocks of Pinus elliottii were placed in the different food chambers. After 28 days, the consumption of each wood block and the percentage of foraging individuals recruited for food chambers were verified in relation to the total survival rate obtained for each one of the 20 replicates. Results showed that completely buried food was most consumed for H. tenuis and presented a higher recruitment rate of workers and soldiers for both species. Although the consumption had non‐significant differences for C. gestroi, these termites exhibited a tendency to prefer completely buried food. In these conditions, it can be concluded that the forms of food placement used in the present research influenced the recruitment of individuals for both species. Data also suggests that when the foraging subterranean termites find food resources in the tunneling substrate, they tend to concentrate their efforts on it, a behavior regime that reduces exposure to external environment.     

Factors that Affect Aggression Among the Worker Caste of Reticulitermes spp. Subterranean Termites (Isoptera: Rhinotermitidae)

Aggression was observed among both inter- and intraspecific combinations of four colonies of Reticulitermes flavipes (Kollar) and Reticulitermes hageni Banks in laboratory assays each month for 4 consecutive months. Termites were most frequently aggressive toward colonies of a different species. Number of individuals that displayed aggression decreased over the study period, from April to July. There was a slight trend toward reduced aggressive behavior as termites were maintained in the laboratory for 3 months. Passive and aggressive individuals were identified and reexamined for display of aggressive or passive behavior toward nonnestmates. Eighty-nine percent of previously aggressive termites displayed aggression a second time. Eighty-eight percent of previously passive termites were passive upon reexamination. Differences in head capsule size between passive and aggressive individuals provided no correlation between the presence of aggressiveness and the head capsule size in the worker caste.     

Chemical Identification and Ethological Function of Soldier-Specific Secretion in Japanese Subterranean Termite Reticulitermes speratus (Rhinotermitidae)

We identified the soldier-specific compounds in the Japanese subterranean termite, Reticulitermes speratus, to clarify their ethological roles. Silica gel column chromatography separated one major soldier-specific compound in the hexane fraction accounting for 70–80% of the total amount of the fraction, while cuticular hydrocarbons constituted the rest. We identified the compound as β-selinene by gas chromatography-mass spectrometry (GC-MS) and nuclear magnetic resonance (NMR) spectroscopy. Comparative GC analyses of the major exocrine glands detected the compound in the soldier’s frontal gland. Both soldiers and workers made aggregation to the hexane fraction, as well as to the crushed heads and head extract of the soldiers. They did not aggregate to cuticular hydrocarbons, making it likely that β-selinene was the aggregation pheromone in this species. The opportunistic predator of this termite, Lasius japonicus, was also attracted to the compounds. The ant workers, therefore, would use the termite aggregation pheromone as a kairomone for hunting them.     

Response of the Formosan subterranean termites (Isoptera: Rhinotermitidae) to baits or nonrepellent termiticides in extended foraging arenas

Distance effects of three treatments, noviflumuron, fipronil, and thiamethoxam, against laboratory populations of the Formosan subterranean termite, Coptotermesformosanus Shiraki, were tested in extended foraging arenas with foraging distances of 50 m. The results showed that during the 10-wk test period, all termites were killed by noviflumuron baits, whereas the nonrepellent termiticides fipronil and thiamethoxam divided the laboratory populations into two groups after causing 25-35% worker mortality. The horizontal transfer of lethal effects of fipronil was < or = 5 m. For thiamethoxam, the distance of transfer was substantially shorter. Because of their dose-dependent lethal time, the nonrepellent termiticides did not fulfill the requirements of a liquid bait model.