Soldier Defense Secretions Research Articles

Variations in Worker Cuticular Hydrocarbons and Soldier Isoprenoid Defensive Secretions Within and Among Introduced and Native Populations of the Subterranean Termite, Reticulitermes flavipes

In social insects, cuticular hydrocarbons (CHCs) play a central role in nestmate recognition. CHCs have proved to be useful for identifying species and differentiating populations. In combination with CHCs, isoprenoid soldier defensive secretions (SDSs) have been previously used in some termite species for chemotaxonomic analyses. This study compared the levels of chemical variation within and among introduced (French) and native (U.S.) populations of the subterranean termite, Reticulitermes flavipes. Worker CHCs and soldier SDSs from termites collected from colonies in nine populations in Florida, Louisiana, and France were analyzed. Discriminant analyses revealed that both localities and populations can be distinguished by using the variation in CHC profiles. Principal component analyses of CHC profiles as well as the calculation of two distance parameters (Nei and Euclidean) revealed remarkable chemical homogeneity within and among French populations. These analyses also showed that the CHC profiles of French populations were closer to termite populations from Louisiana than to those from Florida. Of the six distinct SDS chemotypes, one was common to populations in France and Louisiana. The possibility that populations in France originated from Louisiana, and the potential causes and consequences of chemical homogeneity within introduced populations are discussed.

Cuticular Hydrocarbons and Soldier Defense Secretions of Reticulitermes in Southern California: A Critical Analysis of the Taxonomy of the Genus in North America

Cuticular hydrocarbons (CHC) and soldier defense secretions (SDS) were characterized for collections of Reticulitermes from six counties (Los Angeles, Orange, Riverside, San Bernardino, San Diego, and Santa Barbara) in southern California. Collection sites included the type locality for R. hesperus, Lake Arrowhead (formerly known as Little Bear Lake) in the San Bernardino Mountains. In southern California, there are two CHC phenotypes, SC-A and SC-B, which are easily distinguished by the presence or absence of 5-methyl pentacosane, 5-methyl heptacosane, 5,17-dimethyl pentacosane, and 5,17-dimethyl heptacosane. These phenotypes are similar, but not identical, to previously designated phenotypes of Reticulitermes from northern California. The SDS of termites collected from southern California were characterized; (-)-germacrene A was abundant in all but the four samples from Lake Arrowhead. Soldiers of phenotype SC-A produced >79% germacrene A. The four samples from Lake Arrowhead produced no germacrene A, but contained >78% gamma-cadinene. The SDS from the Lake Arrowhead samples were more similar to those of CA-A/CA-A' from northern California than to any of the CHC phenotypes from southern California. Soldiers of CHC phenotype SC-B produced germacrene A, with the proportion varying from 16.2 to 98.7%. The SDS of SC-B were more similar to those of SC-A than to any of the phenotypes from northern California. The CHC phenotype SC-A found in southern California likely represents R. hesperus and SC-B appears to be a new, as yet undescribed species. We discuss the state of current taxonomic research on Reticulitermes.

Intraspecific variation in soldier defense secretions of Longipeditermes longipes (Isoptera, Nasutitermitinae)

The chemical defense secretions of major and minor soldiers of 46 colonies of the free-ranging termite Longipeditermes longipes (Isoptera, Termitidae, Nasutitermitinae) were isolated and analyzed by chromatography and spectroscopy. The colonies, collected from rainforest sites in Malaysia, showed monoterpene patterns rich in pinenes and limonene but with few qualitative differences among colonies. In marked contrast, the diterpene chemistry is highly variable that includes bicyclic (secotrinervitane), tricyclic (trinervitane), tetracyclic (rippertane), and a spirotetracyclic (longipane) skeleton. This paper presents the compositional and structural diversity, and calculates genetic distances among sympatric and allopatric colonies from a single species.

Biosystematics of Reticulitermes termites in Europe: morphological, chemical and molecular data

In Europe the most abundant naturally residing termite is the subterranean genus Reticulitermes (Rhinotermitidae). Six phenotypes of Reticulitermes have been identified on the basis of morphological, chemical (cuticular hydrocarbons and soldier defensive secretions), and molecular (enzymatic alleles and mitochondrial ND1 sequence) features. They are R. santonensis in western France, R. grassei in southwestern France, northwestern and southern Spain and Portugal, R. banyulensis in northeastern Spain, central area of the Iberian Peninsula and southwestern France, R. lucifugus in Italy and southeastern France, R. balkanensis in the Balkans and R. sp. nov., a recently identified urban phenotype resembling R. balkanensis, in northern Italy and southeastern France. R. santonensis is close kin to the American species R. flavipes. R. grassei, R. banyulensis and R. lucifugus belong to the same species complex. R. balkanensis and the new phenotype R. sp. nov. are close to R. santonensis regarding cuticular hydrocarbons, to the lucifugus complex regarding DNA and to R. clypeatus from Israel regarding morphology. The species status of these genotypes has been confirmed by the mechanisms of species isolation. Prevention of hybridization depends on the method of colony formation in each species. Swarming dates, differences in pheromones, and infertility prevent hybridization by sexual alates. Interspecific aggression between workers prevents hybridization by neotenics. Behavioral and molecular studies have provided many data on the genetic structure of nests, which varies according to species and location. All colonies of R. santonensis are open all year. The colonies of R. grassei in southern areas and all colonies of R. banyulensis are closed families with generally a single reproductive couple. The colonies of R. grassei in northern areas and the colonies of R. lucifugus are open in the summer and closed in the winter. Based on the here presented data, the taxonomy and the speciation of the Reticulitermes genus in Europe are discussed.

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 (-)-alpha-pinene, (-)-beta-pinene, (-)-camphene, myrcene, (Z)- and (E)-ocimene, and (-)-limonene. The major sesquiterpenes produced are (+)-gamma-cadinene, (+)-gamma-cadinene aldehyde, (-)-germacrene A, germacrene B, gamma-himachalene, and beta-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.

Inhibition of fungal spore germination by Nasutitermes: evidence for a possible antiseptic role of soldier defensive secretions.

The antifungal property of two of the principal components of the terpenoid frontal gland secretions of nasute termite soldiers was studied by incubating spore suspensions of the fungus Metarhizium anisopliae with α-pinene and limonene singly or in combination at different concentrations. In vitro assays showed that these substances reduced spore germination through direct and indirect (vapor) contact. To determine if the frontal gland secretions protected termites from fungal infection in vivo, the effect of M. anisopliae on the time course of survival of Nasutitermes costalis and N. nigriceps was studied by exposing termites to either a 4.3 × 107 spores/ml or a control sporeless suspension. The caste composition of experimental groups was manipulated to create mixed-caste subcolonies and monocaste groups. Relative to Coptotermes formosanus, a species that relies on the mechanical defenses of soldiers, N. costalis and N. nigriceps were less susceptible to fungal infection. Spore-exposed N. costalis and N. nigriceps termites had 1.2 times the hazard ratio of death of controls, while the hazard ratio of death of spore-exposed C. formosanus was 11.4 times that of controls. Although the lower susceptibility to infection in Nasutitermes may be explained in part by the antifungal properties of α-pinene and limonene, group composition also played a major role in the survival of spore-exposed termites. Mixed-caste and soldier monocaste groups had 3.4 and 4.7 times the hazard ratio of death, respectively, relative to the worker monocaste treatment. These results suggest that although Nasutitermes terpenoid secretions may have antifungal properties, the caste composition of groups and the social interactions of termites also play a role in determining susceptibility to fungal infection.

Tolerance of Aardwolves to Defense Secretions of Trinervitermes trinervoides

Trinervitermes trinervoides provides over 90% of the diet of aardwolves ( Proteles cristatus ) in South Africa. The soldiers of this termite possess highly effective chemical-defense secretions preventing most myrmecophagous mammals from specializing on them as a primary source of food. Soldiers of T. trinervoides were collected, liquidized in oil, and then added in different concentrations to the diets of captive aardwolves, African civets ( Civettictis civetta ), and bat-eared foxes ( Otocyon megalotis ). The foxes refused to feed on the termite mixture, whereas the civets and aardwolves tolerated concentrations of 1.0% and >7.5% respectively; this supports the hypothesis that ancestral aardwolves were phylogenetically preadapted, via their viverrid-like ancestors, to tolerate noxious chemicals and hence the defense secretions of soldiers of Trinervitermes .

Soldier defensive secretions of two rare nearctic desert termite species

Termite defensive secretions were analysed by GC-MS from hexane extracts of the heads of soldiers from five colonies of A. emersoni and a single foraging group of A. coachellae collected in southern Califomia. The sesquiterpene ether, 4,11-epoxy-cis-eudesmane, was the major component (ca 90%) in soldier secretions of both species. Limonene, three sesquiterpene hydrocarbons, and additional unknown compounds, including a possible diterpene hydrocarbon, were also detected. Intracolony variability of secretions among individual soldiers from one colony of A. emersoni was not observed; however, some minor interspecific differences were found. The occurrence of 4,11-epoxy-cis-eudesmane in four Nearctic and at least four Amitermes spp. from other zoogeographic regions suggests a pre-Cretaceous evolution and dispersal of ancestral Amiterrnes.

Chemical heterogeneity of soldier defensive secretions in the desert subterranean termite, Amitermes wheeleri

Soldier defensive secretions were analyzed by GC/MS in eight spatially separated groups of Amitermes wheeleri collected in Arizona and California. Eleven sesquiterpenoids, four of known structure, were isolated. Quantitative and qualitative differences among the groups of termites were extensive; composition of the defensive secretions among colonies varied from one to six components. Intraspecific differences between pairs of sympatric colonies at six sites were much less pronounced. The variability found in A. wheeleri suggests that soldier defensive secretions in the genus Amiterines are not reliable markers for interspecific systematic comparisons.

Soldier defense secretions of the genusHospitalitermes in Peninsular Malaysia

Soldier defense secretions of the genus Hospitalitermes (Isoptera, Termitidae, Nasutitermitinae) were chemically analyzed by GC-MS and were found to contain volatile monoterpenes and polyoxygenated diterpenes. Interspecific and intraspecific chemical variations for H. umbrinus, H. hospitalis, H. flaviventris, and H. bicolor are described. Interspecific variations in monoterpene and diterpene structures and compositions were evident. A remarkable example of large intraspecific variation from a single species was observed inH. umbrinus, indicating that colonies of this species can be separable into two chemically distinct groups. The diterpenes found in one of these groups have unusually high molecular weights.

Chemical variation in defensive secretions of four species of Nasutitermes

Nineteen monoterpenes and 13 diterpenes were distinguished in soldier defensive secretions of four species of Nasutitermes. Samples of N. columbicus, N. corniger, N. ephratae and N. nigriceps were collected in Costa Rica and Panama. Interspecifically, these species can be differentiated by monoterpenes, which exhibit a complex pattern. Intraspecifically, there is detectable variation in monoterpenes and diterpenes, each of which allows discrimination of four populations of N. corniger. Similarity between two Pacific populations of N. corniger from different environments, a tropical dry forest and a rainforest, suggests there is no climatic influence on chemical compositions of defensive secretions. In addition, results from Costa Rica provide evidence that the Atlantic population of N. corniger is derived from Pacific populations.

Interspecific variation of diterpene composition ofCubitermes soldier defense secretions

Soldiers of the humivorous termite generaCubitermes andCrenetermes (Isoptera: Termitidae: Termitinae) secrete a mixture of unusual diterpene hydrocarbons from the fontanellar opening of the frontal gland. The defense secretions of sixCubitermes species and oneCrenetermes species show species-specific distributions of the five chemically identified major components and 12 unidentified minor components. The secretion is also present with its characteric terpene distribution in white presoldiers and newly molted soldiers, but is completely absent in workers.

Extreme intraspecific chemical variability in soldier defense secretions of allopatric and sympatric colonies ofLongipeditermes longipes

The chemical defense secretions of major and minor soldiers of over 18 colonies of the primitive glue-squirting nasute termiteLongipeditermes longipes (Isoptera, Termitidae, Nasutitermitinae) were analyzed chormatographically. The colonies, collected from four rainforest sites in peninsular Malaysia, showed monoterpene patterns rich in pinenes and limonene but with few quantitative differences between colonies. In marked contrast, the diterpene chemistry is high variable, and includes tricyclic (trinervitane), tetracyclic (rippertane), bicyclic (secotrinervitane), and a new spirotetracyclic (longipane) skeleton. Three new natural products are included in this remarkable and unprecedented example of structural diversity among different colonies of a single species.

Soldier defense secretions of the Malaysian termite,Hospitalioermes umbrinus (Isoptera, Nasutitermitinae)

Trinervita-1(15),8(19)-dien-2β,3α-diol and the new trinervita-1(15),8(19)-dien-2β, 3α-diol 2-O-acetate constitute the major diterpene constituents of the soldier defense secretions of the NasutitermiteHospitalitermes umbrinus of Malaysian region. Studies on the intraspecific variations on sympatric and allopatric populations indicate two distinct patterns of chemical composition.

Sesquiterpenes from termite soldiers. Structure of amiteol, a new 5β, 7β, 10β-eudesmane from [formula omitted][formula omitted

The defense secretions of termite soldiers in the genera Amitermes , Noditermes , and Prorhinotermes contain sesqueterpene hydrocarbons, alcohols and ethers. Three new eudesmanes possessing the hitherto unknown 5β, 7β, 10β configuration were identified from Amitermes excellens .

Chemotaxonomy of Subulitermes and Nasutitermes termine soldier defense secretions. Evidence against the hypothesis of diphyletic evolution of the Nasutitermitinae

The soldier defense secretions of advanced neotropical termites of the genus Subulitermes (Isoptera, Nasutitermitinae) contain diterpenes identicalto those found in Nasutitermes species, while soldiers of the primitive mandibulate nasute genera Cornitermes, Armitermes and Rhynchotermes lack diterpenoid compounds. It now appears improbable that the elongate nasus and intimately associated terpenoid defense secretions evolved from a primitive mandibulate ancestor independently along two phyletic branches. An alternate hypothesis is proposed in which morphological divergence from a common diterpene-producing ancestor may have occurred.

Soldier defense secretions ofTrinervitermes bettonianus (Isoptera, Nasutitermitinae): Chemical variation in allopatric populations

The soldier defense secretions of major and minor soldiers of the East African nasute termiteTrinervitermes bettonianus were analyzed chromatographically and spectrometrically, and the diterpene and monoterpene constituents were determined for three allopatric populations. Incipient colonies from one population raised on food from a second population produced their population-specific terpene pattern independent of food source. Secretion compositions are thus genetically rather than nutritionally determined, in agreement with biosynthetic studies in other nasutes. A minor volatile constitutent, previously proposed to be 3-ethyl-2-octanol was re-identified as (R)-2-decanol.

Interspecific variation in the defence secretions of Nasutitermes soldiers

Termite soldiers of the genus Nasutitermes (Isoptera, Termitidae, Nasutitermitinae) eject a viscous, sticky defense secretion composed of non-polar monoterpene hydrocarbons and polyoxygenated diterpenes from the frontal gland. Chemical compositions are described in detail for the East African species, N. kempae and N. infuscatus, and for the Neotropical species N. ephratae. Additional comparisons with the new world species N. costalis, N. rippertii and N. octopilis are presented. The structure of a new monohydroxykempene from N. ephratae is described, and a physicochemical model for the stickiness of the glue is presented. Although the structures of the individual secretion components vary within the genus, the glue-like nature of the secretion remains essentially unchanged. Chemical analysis of soldier defense secretions may be useful in studying the systematic biology of termites.