J. Kenneth Grace

Biology of Invasive Termites: A Worldwide Review

The number of recognized invasive termite species has increased from 17 in 1969 to 28 today. Fourteen species have been added to the list in the past 44 years; 10 have larger distributions and 4 have no reported change in distribution, and 3 species are no longer considered invasive. Although most research has focused on invasive termites in urban areas, molecular identification methods have answered questions about certain species and found that at least six species have invaded natural forest habitats. All invasive species share three characteristics that together increase the probability of creating viable propagules: they eat wood, nest in food, and easily generate secondary reproductives. These characteristics are most common in two families, the Kalotermitidae and Rhinotermitidae (which make up 21 species on the invasive termite list), particularly in three genera, Cryptotermes, Heterotermes, and Coptotermes (which together make up 16 species). Although it is the largest termite family, the Termitidae (comprising 70% of all termite species) have only two invasive species, because relatively few species have these characteristics. Islands have double the number of invasive species that continents do, with islands in the South Pacific the most invaded geographical region. Most invasive species originate from Southeast Asia. The standard control methods normally used against native pest termites are also employed against invasive termites; only two eradication attempts, in South Africa and New Zealand, appear to have been successful, both against Coptotermes species.

Cuticular Hydrocarbons of Termites of the Hawaiian Islands

Seven species of termites (Isoptera) belonging to three families are found in the Hawaiian Islands. The Kalotermitidae include Neotermes connexus Snyder, Cryptotermes brevis (Walker), Cryptotermes cynocephalus Light, Incisitermes immigrans Snyder, and the recently introduced Incisitermes minor (Hagen). Zootermopsis angusticollis (Hagen), a native of the Pacific Coastal region of North America has become established on Maui and is the sole representative of the Termopsidae. The only rhinotermitid known to be established in the Hawaiian Islands is Coptotermes formosanus Shiraki. A closely related species, Coptotermes vastator Light, has been reported from the Hawaiian Islands, but not recently documented. Cuticular hydrocarbon mixtures were characterized for each of the established and introduced species, as well as for C. vastator from Guam. The diversity of the hydrocarbon mixtures was extreme. At least half the hydrocarbons of C. brevis, C. cynocephalus, I. immigrans, and N. connexus are olefins. C. formosanus and C. vastator make no olefins, but methyl-branched alkanes comprise ca. 95% and 85% of their hydrocarbon mixtures, respectively. Blends of abundant hydrocarbons are species-specific and can be used to identify a given taxon without the diagnostic castes, soldiers, or imagoes. Cuticular hydrocarbon mixtures appear to correlate with habitat requirements.

Identification of Termite Species by the Hydrocarbons in their Feces

Blends of abundant cuticular hydrocarbons are species-specific for termites (Isoptera) and can be used to identify a given taxon without the diagnostic castes, soldiers or adults. We demonstrate that hydrocarbon extracts of termite fecal pellets from damaged wood can also be characterized and used to identify termites responsible for damage, even though termites are no longer present or easily recovered. In structures infested by drywood termites, it is common to find fecal pellets, but difficult to extract termites from the finished wood in service. Nine species belonging to two families (Kalotermitidae and Termopsidae) were examined to compare the hydrocarbon composition of termites and their fecal pellets. Diversity was extensive: at least one half of the amount of the hydrocarbons from Neotermes connexus, Incisitermes immigrans, Cryptotermes brevis, Cryptotermes cynocephalus, Procryptotermes corniceps, and Zootermopsis nevadensis nuttingi was olefins. Incisitermes minor and Pterotermes occidentis incorporated only small amounts of olefins in cuticular hydrocarbons; Marginitermes hubbardi had no detectable olefins. Hydrocarbons extracted from fecal pellets were qualitatively and quantitatively similar to cuticular extracts and can be used to determine the termite species responsible without the termites present.

Cuticular permeability of two species of Coptotermes Wasmann (Isoptera: Rhinotermitidae)

Cuticular permeability (CP) values of worker and soldier castes of Coptotermes formosanus Shiraki and Coptotermes vastator Light were determined using gravimetric techniques. Comparisons were made between castes, species, and between colonies of C. formosanus and C. vastator. CP values did not differ among either species or colony, however they did differ between castes. Data reported here and that of established data in the literature for C. formosanus are inconsistent, with the current report having values as small as half that of previously published data. Examining mean CP data by colony indicated that there is a range for Coptotermes spp. termites, from 6.84+/-1.2 to 25.61+/-1.49 microg H(2)Ocm(-2)mmHg(-1)h(-1) dependent upon caste. A novel method of examining intercaste CP variability is the use of worker:soldier CP ratios. Ratios for C. formosanus were consistent between the current study and previous reports. Mean percentage total body water (%TBW) lost at the time of death was 33.75+/-1.15% (soldiers) and 54.24+/-1.43% (workers) for C. formosanus, and 37.69+/-3.31% (soldiers) and 52.87+/-3.65% (workers) for C. vastator. Consistently, %TBW (and fresh mass) was greater in worker than in soldier castes. These data suggest the use of water storage mechanisms in worker termites, as shown for other rhinotermitids.

Preferences of Coptotermes formosanus Shiraki and Coptotermes gestroi (Wasmann) (Blattodea: Rhinotermitidae) among Three Commercial Wood Species.

The Formosan subterranean termite, Coptotermes formosanus Shiraki, and the Asian subterranean termite, Coptotermes gestroi (Wasmann), are both pests of wood in service in Hawaii and Florida. We conducted a laboratory study using method modified from those described in standard E1-09 of the American Wood Protection Association (AWPA 2009) to assess the termite resistance of three commercially available wood species used in regions of the USA where both termite species occur: Douglas fir, Pseudotsuga menziessii, southern yellow pine, Pinus spp. and redwood, Sequoia sempervirens. A multiple-choice (three-choice) assay was used for four weeks (28 days) in order to simulate field conditions of food choice and assess termite feeding preferences under 28 °C and 72–80% RH. 400 termites (360 workers: 40 soldiers) were released into each test jar. Five replicates and two controls of each wood species were used with each termite species. Termite mortality was recorded at the end of the test; and wood wafers were oven-dried and weighed before and after termite exposure to determine the mass loss due to termite feeding, and rated visually on a 0 (failure) to 10 (sound) scale. There were significant differences in mean mass loss values among the three wood species and between two termite species. The mean mass loss value for redwood was significantly lower than Douglas fir and southern yellow pine with both termite species. However, C. formosanus showed increased feeding on Douglas fir and southern yellow pine compared to C. gestroi.

Comparative Study of the Resistance of Six Hawaii-Grown Bamboo Species to Attack by the Subterranean Termites Coptotermes formosanus Shiraki and Coptotermes gestroi (Wasmann) (Blattodea: Rhinotermitidae)

Bamboo is widely grown and utilized as a construction material around the world, particularly in the tropics. At present, there are about 70 bamboo species and varieties recorded from Hawaii. The objective of our study was to determine the relative resistance of six Hawaii-grown bamboo species to attack by Coptotermes formosanus Shiraki and Coptotermes gestroi (Wasmann). Four-week laboratory feeding trials were performed as described in standard E1-09 of the American Wood Protection Association (AWPA 2009). Samples of each of the six bamboo species were individually exposed to 200 termites (with 10% soldiers); and termite mortality, wood mass loss, and visual appearance of the samples (on a scale of 0–10) were recorded at the conclusion of the trail. Mean mass losses of the six species as a result of termite feeding ranged from 13–29%; with the two most resistant bamboo species, Gigantocholoa pseudoarundinacea and Bambusa oldhamii, demonstrating significantly greater resistance to termite attack than the most susceptible bamboo species, Guadua anguistifolia, with both termite species. Dendrocalamus brandisii, Dendrocalamus latiflorus, and Bambusa hirose were intermediate in their termite resistance. Overall, we observed very little difference in wood preference between C. formosanus and C. gestroi. Although bamboo is a very promising construction material, and species clearly differ in their susceptibility to termite attack, all six species evaluated in the present study would require additional protection for use under conditions of high termite pressure.

Coptotermes formosanus and Coptotermes gestroi (Blattodea: Rhinotermitidae) Exhibit Quantitatively Different Tunneling Patterns

Tunneling behavior and the spatial dispersion of tunnels constructed by the subterranean termites Coptotermes formosanus Shiraki and Coptotermes gestroi (Wasmann) (formerly known as C. vastator Light) (Blattodea: Rhinotermitidae) were examined in foraging arenas. The results indicated that these two termite species construct quantitatively different tunnel systems, supporting visual observations made in earlier studies. Coptotermes gestroi constructed thin, highly branched tunnels, while C. formosanus tended to construct wider and less branched tunnels. Tunnels of C. gestroi showed more spatial dispersion than those of C. formosanus, and this species constructed a larger number of tunnels compared to C. formosanus. The presence or absence of food (wood) within the arena did not influence the tunneling pattern of either species. Although previous observations have suggested that these two termite species exhibit different tunneling behaviors; this is the first quantification of the differences. Comparative studies of the foraging behavior of subterranean termite species contribute to our understanding of their distribution and ecology and may help to improve pest management programs, particularly those based on placement of toxic baits. Moreover, differences in tunneling patterns may reflect different foraging strategies optimized for either tropical (C. gestroi) or subtropical/temperate (C. formosanus) environments.

Termite Species Distribution and Flight Periods on Oahu, Hawaii

Termites are economically-important structural pests, costing residents of Hawaii over

Effects of Exposure Duration on Transfer of Nonrepellent Termiticides Among Workers of Coptotermes formosanus Shiraki (Isoptera: Rhinotermitidae)

100 million annually. On Oahu, the last published termite swarming survey occurred from 1969 to 1971, and the last termite hand-collection survey occurred from 1998 to 2000. To contribute data on termite occurrences on Oahu, a light-trap survey took place from February 2011 to September 2012, and a hand-collection survey occurred from September to November 2012. Formosan subterranean termite, Coptotermes formosanus Shiraki, swarming was compared over the duration of the study, finding peak swarming in May 2011. C. formosanus alate activity density was regressed with environmental factors, finding a negative correlation with average wind speed and a positive correlation with average rainfall. Coptotermes gestroi (Wasmann) alates were observed in April, June, and July 2011 and in June 2012. Four species of termites were found in the hand-collection survey of 44 sites: Incisitermes immigrans (Snyder) (n = 8/44), C. formosanus (n = 2/44), Cryptotermes cynocephalus Light (n = 1/44), and Neotermes sp. (n = 1/44). This study contributes to distribution data for termite species on Oahu and records alate activity for two important termite pests.