Brian T. Forschler

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.

Correlation of mitochondrial haplotypes with cuticular hydrocarbon phenotypes of sympatric Reticulitermes species from the southeastern United States.

Three sympatric Reticulitermes species have been identified in Georgia, USA, based on morphological characters from alates and soldiers: R. flavipes, R. virginicus, and R. hageni, but species identification at individual collection sites is often difficult because alate production is seasonal and soldiers comprise 1–3% of the colony. We therefore set up an experiment to determine if chemical phenotypes and mtDNA haplotypes can be used together to separate species of subterranean termites. Subterranean termites of the order Reticulitermes (Isoptera: Rhinotermitidae) were collected from 20 inspection ports across four soil provinces in Georgia. Each collection was identified to species using dichotomous keys. Two collections, HH11 and BH25, however, could not be unequivocally keyed to species and were classified as unknown. The mitochondrial cytochrome oxidase II (COII) gene was sequenced from individual members of each collection and the variation in cuticular hydrocarbon phenotypes from these same collections was characterized. The cuticular hydrocarbon and mtDNA phylogenetic analyses show agreement with both unknown collections falling out in a separate clade. Specimens from HH11 nad BH25 are different morphologically, chemically, and genetically from the three known sympatric species in Georgia. Our results suggest that these two collections may represent at least one new taxon in Reticulitermes. Furthermore, the association of cuticular hydrocarbon phenotypes and mtDNA haplotypes demonstrates that, when combined with morphological characters, they are useful in separating known species, determining new species, and understanding termite evolution.

Grooming Behavior as a Mechanism of Insect Disease Defense.

Grooming is a well-recognized, multipurpose, behavior in arthropods and vertebrates. In this paper, we review the literature to highlight the physical function, neurophysiological mechanisms, and role that grooming plays in insect defense against pathogenic infection. The intricate relationships between the physical, neurological and immunological mechanisms of grooming are discussed to illustrate the importance of this behavior when examining the ecology of insect-pathogen interactions.

Correspondence of soldier defense secretion mixtures with cuticular hydrocarbon phenotypes for chemotaxonomy of the termite genus Reticulitermes in North America

Soldier defense secretions from samples of Reticulitermes collected in California, Nevada, Arizona, New Mexico, and Georgia were characterized and correlated with cuticular hydrocarbon phenotypes. Twenty-seven cuticular hydrocarbon phenotypes have been defined, and soldier defense secretion (SDS) phenotypes have been described for 25 of these. Forty-five terpenoid compounds were found, including monoterpenes, sesquiterpenes, and a few diterpenes. The monoterpenes include (−)-α-pinene, (−)-β-pinene, (−)-camphene, myrcene, (Z)- and (E)-ocimene, and (−)-limonene. The major sesquiterpenes produced are (+)-γ-cadinene, (+)-γ-cadinene aldehyde, (−)-germacrene A, germacrene B, γ-himachalene, and β-bisabolene. Some SDS phenotypes pair with more than one cuticular hydrocarbon phenotype; however, with two exceptions, each hydrocarbon phenotype is associated with only one SDS phenotype. These chemical characterizations lend support to the conclusion that there are numerous undescribed species of Reticulitermes in North America.

Cuticular hydrocarbons suggest three lineages in Reticulitermes (Isoptera: Rhinotermitidae) from North America.

Cuticular hydrocarbon mixtures can be used to discriminate insect taxa. They have utility for determining phylogenetic relationships where they are independent characters with discrete states and represent a hierarchical distribution of shared, derived characters. We report inferred degrees of relatedness among the chemical phenotypes of Reticulitermes from PAUP (phylogenetic analysis using parsimony) analyses of cuticular hydrocarbon characters. One hundred and forty-one Reticulitermes colonies collected from California, Georgia, New Mexico, Arizona and Nevada were used. Initial maximum parsimony analyses sorted the 141 colonies into 26 chemical phenotypes. Subsequent analyses, using the ancestral species Coptotermes formosanus and Heterotermes sp. as outgroups, sorted Reticulitermes taxa into three major lineages, each characterized by a different set of dominant methyl-branched or unsaturated hydrocarbon components. Reticulitermes in lineage I have cuticular hydrocarbon mixtures with a preponderance of internally branched monomethylalkanes and 11,15-dimethylalkanes. Those in lineage II are defined by a preponderance of 5-methylalkanes and 5,17-dimethylalkanes. Taxa in lineage III are characterized by the predominance of olefins and a relative paucity of n-alkanes and methyl-branched alkanes. Bootstrap analyses and decay indices provided statistical support and robustness for these chemical-based relationships.

Protist Communities from Four Castes and Three Species of Reticulitermes (Isoptera: Rhinotermitidae)

Abstract Previous studies indicate subterranean termite protist communities are qualitatively similar within termite species but differ in relative species abundance between castes. We investigated protist communities from four castes of Reticulitermes flavipes (Kollar), Reticulitermes virginicus (Banks), and Reticulitermes hageni Banks. We used a standardized counting technique and found termite workers and early stage nymphs had the largest population of protists, followed by soldiers and alates. R. flavipes workers averaged 59,000 flagellates compared with 21,000 in R. hageni and 14,000 in R. virginicus. We recorded two new genera, Microjoenia Grassi and Monocercomonas Grassi, from R. virginicus. We identified eight protist species from R. hageni, whereas only four have been previously noted. This is the first report of Dinenympha fimbriata Kirby, Holomastigotes elongatum Grassi, Monocercomonas Grassi, Pyrsonympha minor Powell, Spirotrichonympha flagellata (Grassi), and Trichonympha agilis Leidy from R. hageni. Across all termite species, we found that workers, nymphs, and soldiers had similar relative protist species abundances, with alates being different. The results also demonstrated the utility of using the relative abundance of indicator protist species to identify these three subterranean termite species. The presence of Dinenympha gracilis can be used to distinguish R. flavipes from R. virginicus and R. hageni. R. virginicus has a greater abundance of T. agilis, compared with the other two termite species. The relative abundance of D. fimbriata and Pyrsonympha from R. hageni is greater than those found in both R. flavipes and R. virginicus.

Molecular characterization of neuropeptide F from the eastern subterranean termite Reticulitermes flavipes (Kollar) (Isoptera: Rhinotermitidae)

Neuropeptide F (NPF)-like immunoreactivity was previously found to be abundant in the eastern subterranean termite, Reticulitermes flavipes. Purification of the NPF from a whole body extract of worker termites was accomplished in the current study by HPLC and heterologous radioimmunoassay for an NPF-related peptide, Helicoverpa zea Midgut Peptide-I. A partial amino acid sequence allowed determination of the corresponding cDNA that encoded an open reading frame deduced for authentic R. flavipes NPF (Ref NPF): KPSDPEQLADTLKYLEELDRFYSQVARPRFa. Effects of synthetic NPFs on muscle contractions were investigated for isolated foreguts and hindguts of workers, with Drm NPF inhibiting spontaneous contractions of hindguts. Phylogenetic analysis of invertebrate NPF sequences reveals two separate groupings, with Ref NPF occurring within a clade composed exclusively of arthropods.

Influence of Guidelines and Passageways on Tunneling Behavior of Reticulitermes flavipes (Kollar) and R. virginicus (Banks) (Isoptera: Rhinotermitidae)

Tunneling behavior of laboratory-maintained cultures of Reticulitermes flavipes (Kollar) and R. virginicus (Banks) was examined to determine (1) if the termites build tunnels along preexisting wires or tunnels, and (2) whether tunnels are arranged to optimize search efficiency. Tunnel patterns were considered optimal if, for the number of tunnels present, the maximum area was explored. Termites entered either control arenas or arenas in which they encountered a wire or a pre-formed tunnel. Analyses revealed that R. flavipes and R. virginicus almost always follow pre-formed tunnels, but do not follow wires as readily. Within each species, the distributions of tunnels in treatment arenas were different from distributions in control arenas, most often when pre-formed tunnels were the treatment. Optimal tunnel arrangements in control arenas were found in 62% of R. flavipes patterns with 2 tunnels and in 43% of R. virginicus patterns with 2 tunnels. None of the 3-tunnel patterns from control arenas of R. flavipes and 29% of those of R. virginicus had optimal arrangements. Overall, the spatial arrangement of tunnels in control arenas was significantly different between R. flavipes and R. virginicus.

DNA Technology, Interstate Commerce, and the Likely Origin of Formosan Subterranean Termite (Isoptera: Rhinotermitidae) Infestation in Atlanta, Georgia

Abstract This study was the first to use DNA sequencing data to infer the origin of a Formosan subterranean termite, Coptotermes formosanus Shiraki, infestation. Cytochrome oxidase II gene sequence was used to define eight specific maternal lineages from 14 geographic locations across the southeastern United States, Hawaii, and China. One maternal lineage, A, was unique to all the Atlanta collections and one New Orleans collection, which indicated that termites at these sites shared a common maternal ancestor. The impact of interstate commerce in terms of Formosan subterranean termite introductions is addressed as is the industry response needed to arrest the spread of these termites via commercial traffic in used railroad cross ties.

Subterranean Termite, Reticulitermes spp. (Isoptera: Rhinotermitidae), Colony Response to Baiting with Hexaflumuron Using a Prototype Commercial Termite Baiting System

Six subterranean termite colonies, representing two species, Reticulitermes flavipes and R. virginicus were characterized and baited during the spring and summer of 1993 at three locations in the P...