Sociobiology: Volume 41, Number 1(A & B), 2003

Feature Articles (41A): Totally devoted to a Symposium on the Formosan Subterranean Termite

Feature Articles (41B): Devoted to an enumeration of the staphylinid myrmecophiles found with Dorylus (Dichthadia) laevigatus.


Overview of the Global Distribution and Control of the Formosan Subterranean Termite

By Nan-Yao Su

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In the United States, the number of areas infested with the Formosan subterranean termite has increased in recent years, but its worldwide distribution remains primarily in areas between ca. 35º north and south of the equator. Soil termiticide applications that have been in widespread use by the pest control industry for the half-century are intended for exclusion of soilborne termites from the structures, but their inability to kill vast colonies in soil may have allowed the establishment and increase of Coptotermes formosanus populations in many introduced areas. With the development of baits that can be used to eliminate termite colonies, there is hope that C. formosanus populations may be substantially reduced if area-wide programs are implemented.

KEY WORDS: C. formosanus, population control, area-wide management.

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Experience with Coptotermes formosanus in China (Isoptera: Rhinotermitidae)

By Zhong Junhong & Liu Liling

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Coptotermes formosanus Shiraki is by far the most destructive termite species which occurs widely in South, Central and East China. The northern limit approximates to Huai River. In Guangdong, Guangxi, Hunan, Fujian and Jiangxi provinces, it is responsible for severe damage to buildings. In addition to its attacks on timber in service such as window and door frames, wooden floors, ceilings and wallboards in buildings, it infests ornamental trees, bamboo, plastics, rubber, leather, cotton products, plastic-sheathed electric and telephone cables (including underground cables) and so on. In total the losses caused by C. formosanus annually are estimated to approach 0.8 billion RMB. Current control of C. formosanus in developed cities in southeast China rely on preconstruction soil treatment. After chlordane was severely restricted by our country in 2001, chlorpyrifos, phoxim, permethrin, cypermethrin and other synthetic pyrethroids as alternative insecticides for use in treated soil barriers were approved by the National Constructional Ministry of P.R. China in the beginning of 2001. Baiting is a simple, economical and effective method to destroy colonies of C. formosanus and has become more widely adopted by many firms in cities in recent years. Dusting with arsenic trioxide is one of the most common treatments for an active termite infestation and used by professionals (in particular in rural areas). Direct destruction of the colony of C. formosanus is sometimes done when the termite nest can be located.

KEY WORDS: C. formosanus, preconstruction soil treatment, baiting, dusting.

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Economic Importance of Formosan Termite and Control Practices in Japan (Isoptera: Rhinotermitidae)

By Kunio Tsunoda

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The Formosan subterranean termite, Coptotermes formosanus Shiraki, has threatened Japanese wooden construction for more than 300 years. The distribution of C. formosanus is generally limited to the warmer coastal regions, although this distribution has also been reported to extend to areas with a monthly mean temperature in January of >4ºC and to inland areas. It is estimated that the cost of preventing and controlling termite infestations is at least $US 800 million a year based on the amount of termiticidal formulations consumed in 1996. In addition to the conventional chemical soil and wood treatments, considerable attention has recently been paid to improving the environmental conditions in the crawlspace to reduce the risk of decay and termite attack in Japanese homes.

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Behavior and Reproductive Biology of the Primary Reproductives of the Formosan Subterranean Termite (Isoptera: Rhinotermitidae)

By Ashok Raina, Yong Ihl Park & Christopher Florane

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Primary reproductives or alates of the Formosan subterranean termite, Coptotermes formosanus, swarm from April-June in Louisiana. About 1200 pairs were set up during 1999 and 2000 using agar and birch sawdust medium. Mating was observed and recorded. The tail to tail mating lasted an average of 28 seconds. Over 400 of these pairs were observed for egg laying, egg hatch and incipient colony development. Four discrete oviposition cycles with intervening periods of rest were observed over a period of two years. Average number of eggs in the first batch was 39. Both ovaries and testes increased in size with age. After an initial slow growth total progeny increased over fivefold towards the end of the second year. Multiple matings were essential to obtain maximum reproductive potential. Removal of the progeny during active egg laying induced increased oviposition. Understanding the reproductive biology of the Formosan subterranean termite is a key to the development of novel approaches for management of this pest.

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Factors Regulating Caste Differentiation in the Formosan SubterraneanTermite With Emphasis on Soldier Formation

By Yong Ihl Park & Ashok Raina

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Regulation of caste systems in the Formosan subterranean termite, Coptotermes formosanus, as in other species of termites, is controlled by external factors such as nutrition and internal factors including age and hormone(s). Here we report primarily on soldier formation. The role of various such factors in regulating pre-soldier formation was examined. Removal of existing soldiers resulted in additional pre-soldier formation. Better nutrition among workers contributed to higher pre-soldier formation. Among regulator hormones, juvenile hormone (JH) is known to play a key role in soldier formation. Newly molted workers were competent to form pre-soldiers if treated with JH. Methoprene, a JH analog induced soldier formation in workers up to a maximum of 40 percent indicating the possibility of an inhibitory feedback signal from existing soldiers. Workers had to be in direct physical contact with pre-soldiers or soldiers to cause the inhibition. Corpora allata, that produced JH, were larger in size in the pre-soldiers as compared to workers and soldiers.

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Describing the Spatial and Social Organization of Formosan Subterranean Termite Colonies in Armstrong Park, New Orleans

By C. Husseneder, J. K. Grace, M. T. Messenger, E. L. Vargo & N-Y. Su

ABSTRACT


Colonies of the Formosan subterranean termite, Coptotermes formosanus, possess a complicated spatial and social organization comprising interconnected foraging sites and nests containing variable numbers of reproductives. This complex colony structure may affect the distribution of bait toxicants through the colony. Due to the cryptic life of subterranean termites, their colony organization cannot be directly observed, but needs to be inferred from the genetic structure. We investigated the genetic structure of C. formosanus in Armstrong Park, New Orleans, employing multilocus DNA fingerprinting and microsatellite genotyping. We applied DNA fingerprinting to differentiate colonies according to their genetic profiles. We assigned termites from different sites to colonies through diagnostic bands and genetic similarities. In general, foraging areas of colonies identified by their DNA profiles correlated with results of mark-release-recapture studies. The wide intra- and intercolonial range of genetic similarities indicates a variety of social organizations as well as different proliferation strategies of colonies (budding, swarming). For more detailed analyses of colony organization we are currently using microsatellite genotyping to reveal the numbers of reproductives and degree of inbreeding within colonies. The combined information on genetic structure and social organization of colonies will tell (1) if the majority of colonies is headed by unrelated pairs (colony foundation through swarming) or by multiple related reproductives, and (2) if the presence of multiple reproductives results in a genetic substructure within colonies, which might lead to subsequent colony proliferation through budding. We also intend to apply genetic information to examine how termites invade areas where baiting has eliminated all prior activity.

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Colony and Population Genetic Structure of Formosan Subterranean Termites from Hawaii and Louisiana

By Edward L. Vargo, Claudia Husseneder, J. Kenneth Grace, Gregg Henderson & Dennis Ring

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Formosan Subterranean Termite Feeding Preference as Basis for Bait Matrix Development (Isoptera: Rhinotermitidae)

By J. A. Morales-Ramos & M. G. Rojas

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The chemical composition of the most preferred wood species by the Formosan subterranean termite, Coptotermes formosanus Shiraki, was used as basis for the development of a nutritionally-based bait matrix against subterranean termites. Feeding preferences of the Formosan subterranean termite were determined in a multiple-choice feeding test. Twenty-four commercially available wood species were exposed simultaneously in a circular arena to groups of 2,500 Formosan termites for a period of 40 days at 27ºC and 95% RH. The most preferred wood species were yellow birch (Betulla alleghaniensis), red gum (Liquidambar styraciflua), Parana pine (Araucaria angustifolia), sugar maple (Acer saccharum), pecan (Carya illinoensis), and red oak (Quercus rubra) in order of preference by C. formosanus. High performance liquid chromatography (HPLC) was used to determine the contents of sugars, amino acids, and inorganic salts present in these wood species. Gas chromatography was used to determine the content of fatty acids and esterols in the same wood species. A bait matrix was formulated based on these chemical analyses using pure cellulose as the basic ingredient. Formosan subterranean termites significantly preferred the bait matrix over southern yellow pine wood (Pinus taeda L.) in laboratory and field evaluations.

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Field Evaluation of Nutritionally-Based Bait Matrix against Subterranean Termites (Isoptera: Rhinotermitidae)

By M. Guadalupe Rojas & J. A. Morales-Ramos

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A novel nutritionally based bait matrix was evaluated under field conditions in New Orleans Louisiana and Poplarville, Mississippi against infestations of the Formosan subterranean termite (Coptotermes formosanus Shiraki) and the Easter subterranean termite (Reticulitermes flavipes (Kollar)). The active ingredients tested, included diflubenzuron, chlorfluazuron and some natural products. The Quarterra interception and baiting system was used to deliver the bait matrix. In New Orleans, LA the test area consisted of 3 buildings and wooden areas within the SRRC campus. A total of 230 stations were installed in the wooden areas spaced 20ft (7.29m) apart and 66 stations around the 3 buildings spaced 15ft (5.47m) apart. In Poplarville, MS 396 stations were installed around 31 houses in 3 different neighborhoods spaced as above. These tests were designed to evaluate the areawide management concept. Temperature, termite activity and bait consumption were measured monthly from October 1999 in NOLA and from July 2000 in Poplarville to the present date. Termite activity stopped in the 2 termite infested buildings in New Orleans within 9 months of bait placement and no activity has been observed for the past 8 months. Evidence of heavy termite mortality was observed in both buildings and wooden areas. The rest of the studies are still in progress.

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Detection of Termite Cuticular Hydrocarbons by Solid-Phase Microextraction (SPME)

By John M. Bland, Weste L. A. Osbrink, Mary L. Cornelius, Alan R. Lax & Craig B. Vigo

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Solid phase microextraction (SPME)-gas chromatography-mass spectrometry was used as an alternative method for the detection and identification of termite cuticular hydrocarbons. In compari-son to the hexane extraction method, two SPME methods, headspace-SPME and direct contact-SPME were used to show the variation in cuticular hydrocarbon profiles between Coptotermes formosanus Shiraki, Reticulitermes flavipes, and Kalotermes opproximatus Snyder. Differences between caste members (workers, soldiers, and alates) of C. formosanus are also shown. The sexes of alates and four colonies of C. formosanus were found to have little differences. The headspace-SPME method was also shown to identify other compounds of interest such as fatty acids. Using the direct contact-SPME method, termites were continuously studied over time to monitor chemical changes, showing a periodicity in the production of cuticular hydrocarbons with a two hour cycle.

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Foraging Behavior of Coptotermes formosanus and Reticulitermes flavipes (Isoptera: Rhinotermitidae)

By Mary L. Cornelius

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The foraging behavior of two serious structural pests, the Formosan subterranean termite, Coptotermes formosanus Shiraki and the eastern subterranean termite, Reticulitermes flavipes (Kollar) was examined. Comparative laboratory studies of the trail following, feeding, tunneling behavior, and interactions with wood decay fungi of Formosan and eastern subterranean termites, and of the interspecific interactions between these two termite species were conducted. By increasing our understanding of the foraging behavior and interactions of Formosan and eastern subterranean termites, this research will have an impact on the development of novel control methods to eliminate termite populations that are threatening structures.

KEY WORDS: Coptotermes formosanus, Reticulitermes flavipes, foraging behavior, wood consumption, wood decay fungi, species interaction.

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Installation Guidelines for the Basaltic Termite Barrier: A Particle Barrier to Formosan Subterranean Termites (Summary)

By Julian R. Yates III, J. Kenneth Grace & James N. Reinhardt

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The Formosan subterranean termite, Coptotermes formosanus Shiraki (Isoptera: Rhinotermitidae), is the most economically important insect pest in Hawaii, and an increasing problem in North America.

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Approaches to Biological Control of Termites

By J. Kenneth Grace

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Biological control refers to the application or manipulation of predators, parasitoids, or pathogens in order to suppress and manage insect populations. The cryptic habits of subterranean termites limit their susceptibility to predators. Ants are generally considered to be the most effective predators of termites, and under some conditions may be able to exclude them from occupying feeding sites at the soil surface. However, ants are not able to penetrate far into the subterranean gallery system. To date, there is very little documentation of termite parasitism. Thus, the most promising area of biological control research with termites appears to lie with pathogens. Unfortunately, termite social and chemical defenses appear to be strong limiting factors in inhibiting disease outbreaks in termite colonies. Fungi and nematodes have received the greatest attention to date, and a few commercial formulations are either available or on the near horizon, but their field efficacy is not well documented. Bacteria are also currently of interest, and manipulation via molecular techniques may be a viable path to development of efficient microbial agents. Other approaches to enhancing the efficacy of microbial agents may be the use of combinations of pathogens, or insecticide plus pathogen combinations. Microbial control is a balancing act, in that pathogens must not be repellent, and must not have such rapid or dramatic effects upon the infected individuals that others will subsequently avoid contact with them or with the inoculum source. On the other hand, they must be capable of distribution through the colony and induction of either an epizootic or sufficient delayed mortality to essentially destroy the colony. A self-replicating time bomb, akin to a computer virus, would be the ideal microbial control agent.

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Termite Resistant Wood Products

By J. Kenneth Grace

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Use of either naturally-resistant woods, preservative treatments, or engineered wood products with enhanced insect/decay resistance represents a final line of defense in structural protection. Ideally, proper construction techniques, physical or chemical barriers, and baiting systems can be used to prevent termites from ever entering the structure. However, very few things in life work perfectly all of the time, so redundancy is valuable in termite prevention. If termites penetrate the outer defenses of the building, use of insect-resistant building materials ensures against structural collapse and allows sufficient time for the problem to be discovered and corrected. Laboratory and field tests against Coptotermes formosanus have demonstrated the preservative concentrations necessary for protection, and illustrate the different modes of action of different preservatives (toxicity vs. repellence). For example, borate and CCA (chromated copper arsenate) wood treatments are not repellent to termites, but a four-year field test in Hawaii indicates that only superficial cosmetic damage will occur even with long-term termite exposure and that the timbers remain structurally sound. In contrast, untreated timbers in this test were destroyed within a single year. Research on naturally-resistant woods indicates that these, too, are not immune to termite attack, but can be equivalent in resistance to preservative-treated wood. Steps must also be taken to protect engineered wood products, and viable approaches are incorporation of preservatives during manufacture or incorporation of wood or bark containing naturally-resistant extractives.

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U.S. Forest Service Termiticide Tests

By Terence L. Wagner

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The U.S. Forest Service has been testing chemicals for termite control since 1939. Today its termiticide testing program is nationally recognized for providing unbiased efficacy data for product registration using standardized tests, sites, and evaluation procedures. Virtually all termiticides undergo Forest Service testing before being registered by EPA. Termiticides undergo 18-24 months of laboratory screening before going to the field. Based on the concrete slab test, termiticides are considered effective in the field at the lowest concentration(s) that prevent termites from penetrating treated soil in 10 plots at each site for at least five years. Sites are located in Arizona, Mississippi, Florida, and South Carolina. Results provide a benchmark to compare and assess new and existing products. Because termite control is no longer limited to repellent or contact chemical barriers, tests are also performed on non-repellent delayed-action termiticides, barriers, and wood products. Some of the new products have novel effects on termite biology, ecology, and behavior that require new evaluation procedures. The Forest Service presently has 26 funded agreements with industry involving laboratory screening of three termiticides and field evaluations of 20 termiticides and four impregnated barriers. We also continued to monitor plots on five termiticides and two physical barriers from past (expired) agreements. Marketed termiticides being tested by the Forest Service include bifenthrin (Biflex®), chlorpyrifos, cypermethrin, fenvalerate, permethrin (Dragnet® and Torpedo®), imidacloprid (Premise®), and fipronil (Termidor®). Candidate termiticides include chlorfenapyr, cyfluthrin (Tempo®), and deltamethrin.

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Putative Resistance to Insecticides in the Formosan Subterranean Termite - An Overview

By Weste L. A. Osbrink & Alan R. Lax

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Cytochrome P450 monooxygenase activity, using aldrin as a substrate, has been shown to significantly vary between Reticulitermes virginicus (Banks) colonies. Increased monooxygenase activity was accompanied by significantly increased lethal time to mortality when termites were exposed to chlordane. Lethal times to mortality responses were also established for workers and soldiers of the Formosan subterranean termite, Coptotermes formosanus Shiraki, exposed to chlordane and deltamethrin. There were significant differences in the tolerance ratio of workers between colonies. One colony was 16X more tolerant than another to deltamethrin. C. formosanus soldiers also had significant differences in tolerance ratios. A diagnostic dose of deltamethrin did not kill soldiers from one the C. formosanus colonies. Effect of C. formosanus colony tolerance differences on penetration of insecticide treated sand was also investigated.

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Natural Products as Pesticidal Agents for Control of the Formosan Termite (Coptotermes formosanus Shiraki)

By Mario R. Tellez, Weste Osbrink & Mozaina Kobaisy

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An overview of the literature on investigations of natural products for use against the termite pest Coptotermes formosanus. Our current results in this field are also presented. Results obtained from components of natural sources known to be associated with antitermite activity are presented. Results from additional studies on groups of compounds with structural features known or suspected to be associated with antitermite activity, quinones (benzo- naphtho- andanthra-), are also presented.

KEY WORDS: Coptotermes formosanus, natural products, bioactivity, antitermite.

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IMPASSE™ Termite Barrier: New Pre-Construction Termite Barrier Technology

By P.J. Wege, W.D. McClellan, A.F. Bywater & M.A. Hoppé

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IMPASSE™ termite barrier is designed to protect structures from termite damage. It is intended for installation by trained personnel on to the soil surface before the foundation slab is poured. IMPASSE™ termite barrier is a novel laminate polymer membrane in which an interior layer containing lambda-cyhalothrin is surrounded on either side by layers of polymer impervious to the active ingredient. These layers prevent migration of the active ingredient to the laminate surface, ensuring minimal or no risk of exposure to workers. Isolating the lambda-cyhalothrin from the soil also permits it to degrade at a rate that is independent of soil conditions, ensuring a protracted period of termite protection. The smooth and tough polymer sheet provides a degree of physical protection against termite penetration, whilst the interior termiticide-containing layer enables the membrane to resist direct penetration by termites and provides additional protection in the event of minor accidental damage during installation. Computer modeling, supported by interim data from field trials in the USA and Asia suggest that the duration of termite protection will greatly exceed that of current liquid termiticide treatments.

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Baits as a Tool for Population Control of the Formosan Subterranean Termite

By Nan-Yao Su

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Localized application of arsenic dust was one of the preferred approaches for population control of subterranean termites in the late 1800s to early 1900s in the United States. In the 1950s when the long-lasting and inexpensive cyclodienes were registered, soil barrier treatments become the standard technique for the termite control industry. Because of the large colony size, application of soil termiticides beneath a structure usually does not impact C. formosanus colony, and the surviving colony continues to produce alates and foragers that further infest nearby areas. Following the discovery of molting inhibitory effects of hexaflumuron against subterranean termites in the late 1980s, numerous field studies demonstrated that colonies of C. formosanus could be eliminated by application of hexaflumuron baits. Although it has been established that baits can be used to kill colonies of C. formosanus, there is a need to implement a program to manage C. formosanus populations in a large area.

KEY WORDS: Population control, evaluation criteria, hexaflumuron bait, area-wide management.

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Evaluation of Commercial Termiticides and Baiting Systems for Pest Management of the Formosan Subterranean Termite, Coptotermes Formosanus (Isoptera: Rhinotermitidae)

By Grady J. Glenn & Roger E. Gold

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Increase in Biomass and Soldier Production for Coptotermes formosanus (Shiraki) Workers Maintained in the Laboratory up to Nine Months

By Deborah A. Waller

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The factors that influence rates of soldier production in subterranean termites are poorly understood. In the present study, foraging groups of Coptotermes formosanus Shiraki were collected from baldcypress trees in Lake Charles, Lousiana, in November 1985, and February, April/May and August 1986. Termites were maintained in the laboratory for nine months, seven months, four/five months and zero months, respectively, and then examined for survivorship, termite biomass and soldier production after incubation for an additional five weeks at 30ºC. Within collection periods, there was no association between survivorship and soldier production or between mean dry biomass and soldier production. However, termites maintained for longer periods in the laboratory had significantly greater survivorship, termite dry biomass and soldier production than recently collected termites. It is unclear whether termite age or season of collection influenced these results.

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Myrmecophilous Staphylinidae (Coleoptera) Associated with Dorylus (Dichthadia) laevigatus (Hymenoptera: Formicidae) in Malaysia with Studies of Their Behavior

By David H. Kistner, Stefanie M. Berghoff & Ulrich Maschwitz

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Three new genera are described here; Berghoffia (type species, B. spectabilis new sp.); Pseudolydorus (type species, Pseudolydorus linsenmaieri new sp.); and Pygoplanus (type species P. subterraneus new sp.) Berghoffia represents the first record of the tribe Dorylogstrini outside of Africa. The three new species above are all associated with Dorylus (Dichthadia) laevigatus in Perak, Malaysia as are other new species here listed. These are Doryloxenus coecus new sp.. Zyras (Rhychodonia) praedabunda new sp. and Trachydonia dichthadiaphila new sp. Micropolemon subahensis new sp. was collected from palm oil baits in Poring Hot Springs, Sabah associated with the same ant species. Additionally both Doryloxenus groveri and Micropolemon malayensis were collected associated with the same host. These species had been previously described from specimens associated with termites, most likely left behind after raids of the ants had subsided. A revised phylogeny of the Pygostenini is presented which incorporates the 2 new pygostenine genera. The behavior of most of the species is described. The range of behavior is from predation [Zyras (Rhynchodonia), and Trachydonia] to close association (all of the Pygostenini). Within the Pygostenini, the limuloid genera (Doryoxenus, Pseudolydorus, & Pygosplanus) seem to be more closely integrated into the ant colony than the more elongate species (Micropolemon). The progenitors of all the Pygostenini is in Africa where the more primitive genera are found with the more epigean species of doryline ants. The subgenus Zyras (Rhynchodonia) is presently known only from the Indomalayan zoogeographic region, but since the majority of the known species of Zyras (s. lat.) are from Africa, it is likely that the ancestral link will ultimately be with some of the larger bodied African Zyras. Trachydonia has far more species in Africa than in the Orient so the few species in Malaysia and Indonesia may be the result of invasion after the movement of the Indian subcontinent onto the rest of Asia.

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