New trace fossils of termites (Insecta: Isoptera) from the late Eocene‐early Miocene of Egypt,and the reconstruction of ancient isopteran social behavior

Three new ichnogenera and five new ichnospecies are described for new trace fossils of termitaria (including associated gallery systems) of subterranean termites from upper Eocene through lower Miocene rocks of northern Egypt. All but two ichnospecies (Krausichnus trompitus, ichnogen, and ichnosp. nov., and A’, altus, ichnosp. nov.) show varying degrees of affinity to nests of the extant subterranean termite species Sphaerotermes sphaerothorax (Termitidae, Macro‐termitinae).

Termitichnus qatranii (Bown, 1982) is divided into two ichnospecies, the previously named T. qatranii, and a more generalized form, T. simplicidens, ichnosp. nov. Vondrichnus obovatus, ichnogen. and ichnosp. nov., is named for simple, possibly macrotermitine nests with oblate form, and Fleaglellius pagodus, ichnogen. and ichnosp. nov., records a nest form similar to V. obovatus, but one in which successive vertical growth by chamber apposition has produced a subterranean, tower‐like structure, reminiscent of that built by extant, epigeous Cubitermes. Krausichnus trompitus, ichnogen. and ichnosp. nov., and A’, altus, ichnosp. nov., record two unique nest architectures probably produced by unknown but related species of humivorous termites. The nest architecture expressed by Krausichnus is of wholly unknown affinity and is only distantly related in form to the ichnofossils of other termite nests known from the Tertiary of Egypt.

Study of the architecture of the nests of these ancient termites reveals details important in reconstructing the phylogeny of termite nests and documents two novel blueprints for chamber expansion and society budding in what were probably primitive Macrotermi‐tinae. It also indicates that at least two constructed edifices, earlier known only as epigeous manifestations of extant termite species, were almost certainly first developed by species living underground.

The extant Macrotermitinae are a subfamily of termites originally believed to have evolved in the post‐Eocene of the Ethiopian biogeo‐graphic region. We offer fossil evidence of four distinct, possibly macrotermitine structures from upper Eocene rocks. This evidence suggests that this termite subfamily (or at least their peculiar mode of nest construction) might have evolved considerably earlier, as it is already well established and exhibits several variants by the late Eocene in Egypt.     

<Emphasis Type="Italic">Myrothecium dimorphum</Emphasis>, sp. nov., a soil fungus from beach sand in the Bonin (Ogasawara) Islands,Japan

A new Myrothecium species isolated from beach sand in the Bonin Islands, Japan, is characterized by dark green sporodochia composed of conidiophores with verticillate phialides and ovate and ellipsoidal, often curved conidia on their apexes mixed with erect, straight setae, and seta-like conidiophores with terminal polytomous structures composed of 2–8 digitate polyphialides bearing single globose conidia at each apex.     

Influence of Feed Components on Symbiotic Bacterial Community Structure in the Gut of the Wood-Feeding Higher Termite Nasutitermes takasagoensis

The influence of carbon sources on bacterial community structure in the gut of the wood-feeding higher termite Nasutitermes takasagoensis was investigated. 16S rRNA gene sequencing and terminal-restriction fragment length polymorphism (T-RFLP) analyses revealed that the bacterial community structure changed markedly depending on feed components at the phylum level. Spirochaetes was predominant in the clone libraries from wood- and wood powder-fed termites, whereas Bacteroidetes was the largest group in the libraries from xylan-, cellobiose-, and glucose-fed termites, and Firmicutes was predominant in the library from xylose-fed termites. In addition, clones belonging to the phylum Termite Group I (TG1) were found in the library from xylose-fed termites. Our results indicate that the symbiotic relationship between termite and gut microorganisms is not very strong or stable over a short time, and that termite gut microbial community structures vary depending on components of the feeds.     

Directional selection by the subterranean termite Coptotermes formosanus Shiraki (Isoptera: Rhinotermitidae) at tunnel intersections

Subterranean termites construct complicated tunnel networks for foraging below ground. Thus, they often encounter tunnel intersections during foraging activity. Directional selection by termites at intersections is likely to affect foraging efficiency because depending on their selection, the path length from food resources to the nest can vary significantly. In order to understand how termites are guided to choose the most economical path without the use of pheromones, we artificially excavated two tunnels of varying widths (W₁ and W₂; 2, 3, or 4 mm) that intersected at a 90° angle in each of nine soil‐filled arenas. We observed the directional selection of termites at intersections in arenas with combinations of W₁ (mm) and W₂ (mm) (W₁, W₂). For (W₁, W₂) = (2, 2) and (4, 4), termites statistically equally chose the three directions left, right, and straight, while for the (3, 3) arena combination, termites preferentially decided to go straight. On the other hand, for (W₁, W₂) = (2, 3) (2, 4), and (3, 4), termites advancing from narrower tunnels into intersections tended to turn right or left, while termites coming from the wider tunnel were more inclined to go straight. On the assumption that a wider tunnel is more frequently used by termites in the field, we can deduce that the wider tunnel may represent the most efficient/shortest path. Thus, this simple selection mechanism can prevent termites from deviating from the most economical path.     

Studies on hyphomycetes from the pacific-north west-II./Sporophiala-A new member of dematiaceae

Sporophiala a new genus of Dematiaceae characterized by producing phaeophragmophialospores in acropetal, catenate, simple or branched chains arising from phialides, withS. prolifica as type is described.     

<Emphasis Type="Italic">Hirsutella proturicola</Emphasis> sp. nov. isolated from a proturan, <Emphasis Type="Italic">Baculentulus</Emphasis> densus (Protura,Hexapoda)

A new species of Hirsutella, H. proturicola, isolated from a subterranean proturan (Baculentulus densus; Protura, Hexapoda), is described and illustrated. Hirsutella proturicola is characterized by producing monoblastic phialides of 24–51.5 × 2.5–5 μm with a slightly roughened neck, fusiform and curved conidia of 9–18 × 2.5–4 μm that have a truncate base and a papillate projection often capped with sheath-like mucilage, and pluricellular, globose to subglobose chlamydospores of 21–48 × 21–41.5 μm. This species is morphologically and phylogenetically close to H. rostrata, an acaropathogenic species, but can be distinguished from the size of the phialides and the size and shape of the conidia.     

Foraging,recruitment and predation by Decamorium uelense (Sanstchi) (Formicidae: Myrmicinae) on termites in Southern Guinea Savanna,Nigeria

Summary The myrmicine ant Decamorium uelense is a predator of small termites which feed inside their food sources of roots and plant debris. The ant uses a specialized recruitment regime to secure its prey. A scout ant searches for foraging termites and returns to the nest where it recruits a column of 10–30 ants which attack and immobilize the termites. A mass recruitment phase is then instigated, with larger numbers of ants retrieving the prey. The major prey item is Microtermes (consumed at an annual rate of 632 termites m^-2), with other small termites accounting for 0.5% of the total predation. The annual predation of Microtermes in primary savanna woodland by D. uelense removes 74% of the standing population.     

Mammalian predation on termites: tests with the burrowing mouse (Oxymycterus roberti) and its prey

Summary A laboratory study of mammalian predation on termites was conducted using the burrowing mouse (Oxymycterus roberti) and eight species of central Brazilian termites. The results of preference trials demonstrate that Oxymycterus discriminate between termite species and that the eight prey species fall into three groups: least preferred, most-preferred and a group of intermediate preference.Preferences of Oxymycterus are not explained by the size or the nutritional quality of the termite species. However, three measures of termite soldier-based defense do correlate highly with preference. Termite species with soldiers exhibiting a mechanical or mixed defense are greatly preferred by Oxymycterus over species with soldiers exhibiting a chemical based defense. This chemical defense by termites is effective in severely limiting predation by small mammals.     

Fungal Infection of Mantis Shrimp (<Emphasis Type="Italic">Oratosquilla oratoria</Emphasis>) Caused by Two Anamorphic Fungi Found in Japan

Two fungal pathogens of the mantis shrimp (Oratosquilla oratoria) in Yamaguchi and Aichi Prefectures, Japan are described as the new species Plectosporium oratosquillae and Acremonium sp. (a member of the Emericellopsis marine clade). Both fungi infect the gills of the mantis shrimp, which become brown or black due to melanization. The former species is characterized by its slow growth on artificial seawater yeast extract peptone glucose (PYGS) agar, pale yellow to pale orange or grayish yellow colonies, short cylindrical solitary phialides with a wavy tip, and one-celled ellipsoidal conidia. Although lacking the two-celled conidia demonstrated by the type species Plectosporium tabacinum, the taxonomic placement of the new species was confirmed by DNA sequence analysis of the internal transcribed spacer (ITS) region of ribosomal DNA (ITS1, 5.8S rDNA and ITS2). Acremonium sp., the other causal pathogen, differs from P. oratosquillae by its fast growth on PYGS agar, pale orange to salmon-colored colonies, long, slender conidiophores consisting of solitary phialides with tips lacking an undulate outline, and typically cylindrical conidia. Analysis of ITS and β-tubulin gene sequences placed this fungus within the phylogenetically distinct Emericellopsis (anam. Acremonium) marine clade. Various physiological characteristics of both pathogens were also investigated. This is the first report of a fungal infection found on the mantis shrimp in Japan.     

The effect of ergosterol on the allogrooming behavior of termites in response to the entomopathogenic fungus Metarhizium anisopliae

Termites have physiological and behavioral immunities that make them highly resistant to pathogen infections, which complicates biocontrol efforts. However, the stimuli that trigger the pathogen-avoidance behaviors of termites are still unclear. Our study shows that workers of Coptotermes formosanus exposed to the conidia of Metarhizium anisopliae exhibited a significantly higher frequency and longer duration of allogrooming behaviors compared with untreated termites. Volatile compounds in the cuticle of control termites and termites previously exposed to a suspension of M. anisopliae conidia were analyzed and compared using a gas chromatography-mass spectrometer (GC-MS). Our results showed that the amount of ergosterol differed between the fungus-exposed and control termites. Choice tests showed that termites significantly preferred to stay on filter paper treated with ergosterol (0.05, 0.1, or 1.0 mg/mL) compared with control filter paper. In addition, termites exposed to ergosterol followed by M. anisopliae conidia were allogroomed at a significantly higher frequency and for a longer duration than termites exposed to alcohol (the solvent used with the ergosterol in the ergosterol trials) alone followed by M. anisopliae conidia. These results showed that ergosterol may enhance the allogrooming behavior of termites in the presence of entomopathogenic fungi.     

Methane and hydrogen production in a termite-symbiont system

Methane and hydrogen emission rates and the ¹³C of CH₄ were observed for various termites in Australia, Thailand and Japan. Combined with the already reported emission rates of CH₄ in the literature, the phylogenetic trend was examined. Emission rates of the observed termites were categorized into five groups: group I with high CH₄ and low H₂ emission rates with a CH₄/H₂ ratio of typically 10/1; group II with high CH₄ and high H₂ emissions with a CH₄/H₂ ratio of 4/1–1/2; group III with low emission rates of CH₄ and H₂; group IV with high H₂ and insignificant CH₄ emissions; and group V with insignificant emissions for both CH₄ and H₂. In lower termites, there are both colonies infected and uninfected with methanogens even in the same species, and no specific trend in CH₄ and H₂ emissions was observed within a genus. Whether protozoa in the hindgut of termites are infected with methanogens or not and the differences in species compositions of protozoa are possibly responsible for the inter-colonial variations. The proportions of infected colonies were possibly small for the family Kalotermitidae (dry wood feeders), and relatively large for families of wet or damp wood feeders. The hydrogen emission rate possibly depends on the locality of methanogens: namely, whether they are intracellular symbionts of protozoa or whether they are attached to the hindgut wall. Emission rates of higher termites were classified into groups according to genera and the diet. Most species of soil or wood/soil interface feeders classified into group I, while the soil feeders Dicuspiditermes in Thailand and Amitermes in Australia were classified into groups with high H₂ emission rates. Typical wood-feeding termites and fungus-growing termites were classified into group III. The results indicate that higher termites tend to increase the CH₄ emission rate during dietary evolution from wood- to soil-feeding, and two types of the system with different efficiencies of interspecies transfer of H₂ have been formed. The ¹³C of CH₄ was discernible with a difference in the decomposition process in the termite–symbiont system among lower termites, fungus-growing termites and other higher termites.     

Morphology,Phylogeny, and Diversity of Trichonympha (Parabasalia: Hypermastigida) of the Wood‐Feeding Cockroach Cryptocercus punctulatus

ABSTRACT. Trichonympha is one of the most complex and visually striking of the hypermastigote parabasalids—a group of anaerobic flagellates found exclusively in hindguts of lower termites and the wood‐feeding cockroach Cryptocercus—but it is one of only two genera common to both groups of insects. We investigated Trichonympha of Cryptocercus using light and electron microscopy (scanning and transmission), as well as molecular phylogeny, to gain a better understanding of its morphology, diversity, and evolution. Microscopy reveals numerous new features, such as previously undetected bacterial surface symbionts, adhesion of post‐rostral flagella, and a distinctive frilled operculum. We also sequenced small subunit rRNA gene from manually isolated species, and carried out an environmental polymerase chain reaction (PCR) survey of Trichonympha diversity, all of which strongly supports monophyly of Trichonympha from Cryptocercus to the exclusion of those sampled from termites. Bayesian and distance methods support a relationship between Trichonympha species from termites and Cryptocercus, although likelihood analysis allies the latter with Eucomonymphidae. A monophyletic Trichonympha is of great interest because recent evidence supports a sister relationship between Cryptocercus and termites, suggesting Trichonympha predates the Cryptocercus‐termite divergence. The monophyly of symbiotic bacteria of Trichonympha raises the intriguing possibility of three‐way co‐speciation among bacteria, Trichonympha, and insect hosts.     

Termites as models of swarm cognition

Eusociality has evolved independently at least twice among the insects: among the Hymenoptera (ants and bees), and earlier among the Isoptera (termites). Studies of swarm intelligence, and by inference, swarm cognition, have focused largely on the bees and ants, while the termites have been relatively neglected. Yet, termites are among the world’s premier animal architects, and this betokens a sophisticated swarm intelligence capability. In this article, I review new findings on the workings of the mound of Macrotermes which clarify how these remarkable structures work, and how they come to be built. Swarm cognition in these termites is in the form of “extended” cognition, whereby the swarm’s cognitive abilities arise both from interaction amongst the individual agents within a swarm, and from the interaction of the swarm with the environment, mediated by the mound’s dynamic architecture. The latter provides large scale “cognitive maps” which enable termite swarms to assess the functional state of their structure and to guide repair efforts where necessary. The crucial role of the built environment in termite swarm cognition also points to certain “swarm cognitive disorders”, where swarms can be pushed into anomalous activities by manipulating crucial structural and functional attributes of the termite system of “extended cognition.”     

Inter- and intraspecific alarm response in the termiteMacrotermes subhyalinus (Rambur)

Summary In laboratory studies we analysed the reaction of aMacrotermes subhyalinus (Rambur) colony in contact with a small group of test termites placed at the head of a gallery system. As test termites we usedMacrotermes subhyalinus of the same nest,Macrotermes subhyalinus of a foreign nest and the related sympatric speciesMacrotermes bellicosus (Smeathman).Contact with an introduced group of foragers evokes a significant recruitment of workers and minor soldiers. The intensity of the response depends on the test groups we introduced. The highest recruitment results withM. bellicosus andM. subhyalinus from a foreign nest. During the recruitment the increase in the number of minor soldiers is greater than the increase in the number of workers, and as a result, the relative proportion of the soldiers in the population leaving the nest is significantly higher than before.Direct observations at the place of contact show thatM. bellicosus individuals are immediately attacked and killed. In the case ofM. subhyalinus from a foreign nest, not all individuals are killed; some are only intensively examined with the antennae or seized with the mandibles.M. subhyalinus from the same nest are rarely attacked. Mostly they are antennally inspected or enter the gallery system unhindered.The recruitment of termites from the nest is released by major workers which, after contact with the test termites, run back into the nest and provoke an alarm by tactile stimuli. On their way back into the nest the major workers, in all probability, lay a pheromone trail which leads the outgoing alarmed termites towards the place of contact.     

Nests of subterranean fungus-growing termites (Isoptera, Macrotermitinae) as nutrient patches for grasses in savannah ecosystems

Savannah are ecosystems in which mineral nitrogen is considered as a limiting factor for plant productivity. They are heterogeneous and spatially structured in patches, or islands, where mineral nitrogen content is concentrated. Among the soil macrofauna, termites of the Macrotermitinae subfamily are major determinants of soil heterogeneity through the biogenic underground nest structures (fungus‐comb chambers) they produce. To study the role of the heterogeneity created by termites on Pennisetum pedicellatum, an herbaceous grass species was grown in greenhouse. This was carried out using an homogeneous soil poor in mineral nitrogen, and an heterogeneous soil with patch, made of (i) Ancistrotermes cavithorax fungus‐comb chamber wall and (ii) soil with the same mineral nitrogen content as the termite handled soil. Plants exhibited a better growth on patch of termite‐modified soil whereas no significant differences were shown with the supply of mineral nitrogen. The presence of fungus‐comb chamber wall material resulted in an increase of fine root biomass and root/shoot ratio. We conclude that termites, through their building activities, may create nutrient patches available to grasses. Concurrently, our data illustrate that the higher mineral nitrogen content in termite‐built structures is not the only factor responsible for plant growth.     

Ant- and termite-eating in Australian mammals and lizards: a comparison

Abstract Relative abundance is one factor that influences selection of prey by insectivorous mammals and lizards. Ants and termites are extremely abundant over most of inland Australia. Their patterns of abundance are also broadly similar across climatic gradients, being most and least abundant in seasonally arid (tropic and sub-tropic) and temperate mesic regions, respectively. All else being equal (e. g. mechanisms of prey defence, palat-ability, availability), animals that eat many termites should also eat many (adult) ants. The present study asks three questions: (i) What is the diversity of specialized ant-eaters (>50% volume)?; (ii) Does specialization vary with climate?; and (iii) Are ants and termites eaten in broadly similar proportions (using an earlier study on termites). Of the mammals, only the echidna (Tachyglossus aculeatus) in mesic regions, and probably the marsupial mole (Notoryctes typhlops) in the arid zone and the striped possum (Dactylopsila trivirgata) in mesic regions are ant-specialized. Ant-specialization in mammals shows no pattern with regard to climate. Of the lizards, only four agamid lizards are ant-specialized: Moloch horridus (arid, semi-arid), Ctenophorus fordi (arid, semi-arid), Ctenophorus isolepis (arid) and Ctenophorus maculatus (arid). Specialization on ants by lizards is greatest in the arid zone (4 spp.); no lizard species were found to be ant-specialists in mesic regions. In the arid and semi-arid zone, two mammals each specialize on termites and one on ants; in mesic regions, two mammals specialize on ants and one on termites. Specialized insectivorous mammals thus demonstrate no marked preference for either termites or ants. Lizards, in contrast, are markedly termite-specialized (4 ant-specialist spp., 16 termite-specialist spp.), and specialization is greatest in the arid zone (16 spp.). Greater specialization on termites than on (adult) ants in lizards is explained with reference to differences in prey defence and palatability between ants and termites. Consumption of ant brood (eggs, larvae, pupae) appears to be associated with a fossorial foraging mode (the marsupial mole N. typhlops; spp. of Aprasia lizards; spp. of blindsnakes Ramphotyphlops).     

The influence of subterranean termites on the hydrological characteristics of a Chihuahuan desert ecosystem

Summary Rainfall simulation at an average intensity of 124 mm·h^-1 was used to compare infiltration and run off on arid areas where subterranean termites had been eliminated four years prior to the initiation of the study (termite free) with adjacent areas populated by subterranean termites (termites present). Infiltration rates on termite free plots with less than 5% perennial plant cover were significantly lower 51.3±6.8 mm·h^-1 than rates on comparable termites present plots 88.4±5.6 mm·h^-1. On plots centered on Larrea tridentata shrubs, there were no differences in infiltration rates with or without termites. Plots with shrub cover had the highest infiltration rates 101±6 mm·h^-1. Highest run-off volumes were recorded from termite free <5% grass cover plots and the lowest from plots with shrubs. There were no differences in suspended sediment concentrations from termites present and termite free plots. Average bed load concentration was more than three times greater from termite free, <5% cover plots than from termites present, <5% cover plots.The reduction in infiltration, high run-off volumes and high bedloads from termite free areas without shrub cover is related to increased soil bulk density resulting from the collapse of subterranean galleries of the termites that provide avenues of bulk flow into the soil. Subterranean termites affect the hydrology of Chihuahuan desert systems by enhancing water infiltration and retention of top soil. The presence of a shrub canopy and litter layer cancels any effect of subterranean termites on hydrological parameters. Since approximately 2/3 of the area is not under shrub canopies, subterranean termites are considered to be essential for the maintenance of the soil water characteristics that support the present vegetation.     

Materials for the fungus flora of Japan (56)Mariannaea camptospora andM. elegans var.punicea from Japan

One additional species and a variety ofMariannaea, M. camptospora andM. elegans var.punicea, were recorded for the first time in Japan.Mariannaea camptospora formed two types of conidiophores. One type was characterized by simple verticillate phialides sometimes with punctuate walls at the base, producing long oblique conidial chains, and symmetrical spindle-shaped conidia. The other type was characterized by more crowded and shorter phialides with small conidial droplets and hemispherical to concave smaller conidia.Mariannaea elegans var.punicea was characterized by distinct red purple pigmentation in agar media. (55): Udagawa, S. and Uchiyama, S., Mycoscience41: 263–267, 2000.     

Electron microscopy of selectively stimulated microconidiogenesis in wild type Neurospora crassa

Summary Microconidiogenesis in Neurospora crassa wild type is selectively stimulated by iodoacetate. The ultrastructural features of various stages of microconidiogenesis have been studied. The conidiogenous cells or phialides first bear a protuberance which then erupts into a conspicuous collarette surrounding, above a lamellated cushion, the successively budded, often elongated microconidia.     

Nitrogen Fixation by Termites in Tropical Forests, Thailand

Nitrogen (N) fixed by termites was evaluated as a N input to decomposition processes in two tropical forests, a dry deciduous forest (DDF) and the neighboring dry evergreen forest (DEF), Thailand. A diverse group of termite species were assayed by acetylene reduction method and only the wood/litter-feeding termites were found to fix N. More intensive samplings of two abundant species, Microcerotermes crassus and Globitermes sulphureus, were done across several seasons, suggesting N fixation rates of 0.21 and 0.28 kg ha⁻¹ y⁻¹ by termites in the DDF and DEF, respectively. Also, estimates of asymbiotic N fixation rates were 0.75 and 3.95 kg ha⁻¹ y⁻¹. N fixed by termites and by asymbiotic fixers is directly supplied to decomposers breaking down dead plant material and could be a major source of their N. N fixed by termites was 7–22% of that fixed by termites and asymbiotic fixers. Although N fixed by termites is a small input compared to other inputs, this N is likely important for decomposition processes.