Endogenous synchronization of the chemical signature of Reticulitermes (Isoptera: Rhinotermitidae) worker termites

Termites of the genus Reticulitermes are characteristic of temperate regions. Their colonies comprise various castes, the most numerous being that of workers which can develop into soldiers or secondary reproductives (neotenics). Each caste has a mixture of hydrocarbons (HCs) on the cuticle forming a chemical signature. The primary aim of this study was to compare the changes in the chemical signature of a population of worker termites fed on paper with juvenile hormone to differentiate them into soldiers with a control population of termites fed only on paper or wood for one month. Gas chromatography was used to analyze the cuticular profiles of Reticulitermes flavipes termites to determine whether they changed, and, if so, when and how. The data collected over one month showed that the workers fed with JH did not differentiate into soldiers but that there were progressive changes in the hydrocarbon profile independent of the treatment. These results indicate that the differentiation of the chemical signature of the worker caste is a dynamic process, depending only on time and not on colony membership, confirming that, for these termites, this signature has a lesser role in colony membership than caste membership, unlike the chemical signatures of other social insects. The temporal process of this cuticular change is also associated with a change in the alkene/methyl-branched alkane ratio.     

Experimental manipulation of termites (Isoptera,Macrotermitinae) foraging patterns in a Sahelo-Sudanese savanna: effect of litter quality

Summary The foraging pattern of four sympatric fungus-growing termite species was followed for five months of the dry season in a semi-arid savanna (Senegal). Experimental plots were established with and without termites (exclusion with insecticide) and amended with different litter quality: Acacia leaves, millet canes, ground millet (<500 µm), Combretum wood and cattle manure. Results showed different foraging patterns between the fungus-growing species. Odontotermes nilensis, for example, rapidly developed its foraging activity on the litter. In contrast, Macrotermes subhyalinus delayed its foraging activity but, extensively collected the food after its discovery. Differences in food preference were evident between the termite species: Odonto-termes nilensis foraged preferentially on millet, canes and Acacia leaves, Macrotermes subhyalinus preferred cane and ground millet. Ancistrotermes guineensis selected the millet, and Microtermes sp. was mostly recorded foraging on pieces of dead wood. Relationships between different foraging parameters were established to assess the relative intensity of this activity over the dry season. The comparison between untreated plots and plots treated with Fipronil® (Aventis) clearly demonstrated that the exclusion of termites from part of the savanna is possible and that this possibility can be used to study their role on the soil and organic matter cycle in situ.     

Re-examination of the phylogeny of Rhacophoridae (Anura) based on mitochondrial and nuclear DNA

The phylogenetic relationships among rhacophorid frogs are under dispute. We use partial sequences of three mitochondrial (12S rRNA, 16S rRNA, and cytochrome b) and three nuclear protein-coding (Rag-1, rhodopsin exon 1, and tyrosinase exon 1) genes from 57 ingroup taxa and eight outgroup taxa to propose a hypothesis for phylogenetic relationships within Rhacophoridae. Our results support recognition of the genus Feihyla, and Chiromantis is the sister taxon to the clade formed by Feihyla, Polypedates and Rhacophorus. We place Aquixalus odontotarsus within Kurixalus, and the remaining species of Aquixalus and Philautus jinxiuensis into the genus Gracixalus. We give Philautus (Kirtixalus) the rank of genus and place Philautus menglaensis within it. The division of species groups among Chinese Rhacophorus needs revision, and a cryptic species is revealed within Rhacophorus nigropunctatus. Rhacophorus pingbianensis is considered a synonym of Rhacophorus omeimontis. The validity of Rhacophorus hui is confirmed by present molecular evidence.     

Molecular phylogeny of Asian termites (Isoptera) of the families Termitidae and Rhinotermitidae based on mitochondrial COII sequences

The families Termitidae and Rhinotermitidae are the most evolved and diverse groups of the social insects, termites (Order Isoptera), showing elaborated morphology and complex behavior. Molecular phylogeny of termites with the emphasis on these families was examined by Bayesian and maximum-likelihood analyses based on DNA sequence of mitochondrial cytochrome oxidase II (COII) gene of 31 genera sampled in Asia (mainly Thailand and Japan) along with those reported previously. Termitidae was monophyletic and originated from within polyphyletic Rhinotermitidae. Among the four subfamilies of Termitidae, Macrotermitinae was monophyletic suggesting a single common origin of fungus-growing habit characteristic for this subfamily, and was placed in the basal position in the family. A group consisting of other subfamilies Termitinae and Nasutitermitinae, though some important groups were still untouched, was the most apical but neither Termitinae nor Nasutitermitinae formed a monophyletic lineage. It was implied that, as defense systems of the soldier castes, the appearance of snapping mandibles has occurred at a single event, but the development of nasus for chemical secretion has probably not. Our tree provides some evidence concerning contradictions in the previously proposed phylogeny of termites.     

Resolving the evolution of sterile worker castes: a window on the advantages and disadvantages of monogamy

Many social Hymenoptera species have morphologically sterile worker castes. It is proposed that the evolutionary routes to this obligate sterility must pass through a ‘monogamy window’, because inclusive fitness favours individuals retaining their reproductive totipotency unless they can rear full siblings. Simulated evolution of sterility, however, finds that ‘point of view’ is critically important. Monogamy is facilitating if sterility is expressed altruistically (i.e. workers defer reproduction to queens), but if sterility results from manipulation by mothers or siblings, monogamy may have no effect or lessen the likelihood of sterility. Overall, the model and data from facultatively eusocial bees suggest that eusociality and sterility are more likely to originate through manipulation than by altruism, casting doubt on a mandatory role for monogamy. Simple kin selection paradigms, such as Hamilton''s rule, can also fail to account for significant evolutionary dynamics created by factors, such as population structure, group-level effects or non-random mating patterns. The easy remedy is to always validate apparently insightful predictions from Hamiltonian equations with life-history appropriate genetic models.     

The effect of nonnestmates on foraging trail caste ratios ofNasutitermes corniger (Isoptera: Termitidae)

The mean number of workers on foraging trails ofNasutitermes corniger Motschulsky decreases after the introduction of nonnestmate conspecifics or congeneric heterospecifics. The mean number of soldiers on foraging trais does not vary when nestmates, conspecific nonnestmates or heterospecific workers are introduced. When heterospecific soldiers are introduced to the trail, however, the number ofN. corniger soldiers increases. Consequently, the ratio of soldiers to workers increases. These findings demonstrate that the addition of nonnestmates temporarily alters caste ratios on foraging trails, representing a shift in the distribution of castes in response to environmental perturbation.     

Development of non-reproductive castes in the neotropical termite generaCornitermes,Embiratermes andRhynchotermes (Isoptera,Nasutitermitinae)

Summary The developmental pattern of the neuter castes was studied in the mandibulate nasute generaCornitermes, Embiratermes andRhynchotermes. InCornitermes walkeri, all the workers and soldiers are male. There are two larval and a single worker instar. Workers can molt into presoldiers. InEmbiratermes chagresi andRhynchotermes perarmatus, both sexes are present among the neuters. A slight sexual dimorphism (males > females) is discernible among both larval instars and among workers ofE. chagresi; female workers can molt into presoldiers. InR. perarmatus, the sexual dimorphism is conspicuous from the first larval instar on. Male larvae go through two instars, then give rise to workers, which do not molt. InR. perarmatus, there is no worker stage in females, but a third larval instar, preceding the presoldier. Hypotheses are proposed as to the evolution of these caste patterns, attempting to conciliate present knowledge of Nasutitermitinae phylogeny and known evolutionary trends affecting termite caste patterns, according to the species' ecology.Research Associate: National Fund for Scientific Research (Belgium).     

Developmental pathways of Glossotermes oculatus (Isoptera, Serritermitidae): at the cross-roads of worker caste evolution in termites

SUMMARY The onset of a specialized ("true") worker caste is a crucial step in the evolution of termite societies. Such workers, permanently excluded from wing development, repeatedly evolved from totipotent immatures, called "false" workers or pseudergates. In the family Rhinotermitidae, the presence of true workers and the level of specialization of this caste are highly variable, and key taxa illustrate transitional situations providing clues about worker evolution. Here we focused on the status of working immatures of Glossotermes oculatus , from the family Serritermitidae, now thought to represent either the sister-group of the Rhinotermitidae or a basal lineage nested within them. Contrary to previous assumptions, we show that the apterous immatures performing worker tasks in G. oculatus are the source of the single wing-budded nymphal instar preceding the alate. Consequently, they qualify as pseudergates rather than true workers. However, the sex ratio is strongly male biased in pseudergates and soldiers, which is a trait usually restricted to termites with true workers. We therefore argue that pseudergates of G. oculatus are close to a point where the species could easily shift toward the differentiation of a true worker caste, and that G. oculatus pinpoints a new possible route for the evolution of true workers from pseudergates.     

Molecular phylogeny of the tribe Sphodrini (Coleoptera: Carabidae) based on mitochondrial and nuclear markers

A phylogenetic analysis of 6.4 kb of nucleotide sequence data from seven genes (mitochondrial cox1-cox2 and tRNA(leu), and nuclear Ef-1alpha C0, Ef-1alpha C1, 28S, and 18S) was done to reconstruct the phylogenetic relationships of the ground-beetle tribe Sphodrini. Gene regions of variable nucleotide length were aligned using both a secondary structure model, Clustal W, and a combination of the two. Sensitivity analysis was performed in order to explore the effect of alignment methods. The ribosomal and protein-coding genes were largely congruent based on the ILD test and partitioned Bremer support measures. MtDNA analysis provided high resolution and high support for most clades. The tribe Sphodrini and the related tribes Platynini, Pterostichini and Zabrini made up monophyletic clades, but the relationship between them was weakly resolved and sensitive to alignment strategy. Previously suggested relationships between subtribes of Sphodrini were not corroborated, and only the subtribe Atranopsina revealed high support as the sister clade to the other subtribes. The analyses clearly demonstrated the importance of exploring effects of alignment methods that may become particularly important in resolving polytomies and nodes with low support.     

Molecular phylogeny of Moenkhausia (Characidae) inferred from mitochondrial and nuclear DNA evidence

Moenkhausia is one of the most speciose genera in Characidae, currently composed of 75 nominal species of small fishes distributed across South American hydrographic basins, primarily the Amazon and Guyanas. Despite the large number of described species, studies involving a substantial number of its species designed to better understand their relationships and putative monophyly are still lacking. In this study, we analysed a large number of species of Moenkhausia to test the monophyly of the genus based on the phylogenetic analysis of DNA sequences of two mitochondrial and three nuclear genes. The in‐group included 29 species of Moenkhausia, and the out‐group was composed of representatives of Characidae and other members of Characiformes. All species of Moenkhausia belong to the same clade (Clade C); however, they appear distributed in five monophyletic groups along with other different genera, which means that Moenkhausia is polyphyletic and indicates the necessity of an extensive revision of the group.     

A phylogeny of Diprotodontia (Marsupialia) based on sequences for five nuclear genes

Even though the marsupial order Diprotodontia is one of the most heavily studied groups of Australasian marsupials, phylogenetic relationships within this group remain contentious. The more than 125 living species of Diprotodontia can be divided into two main groups: Vombatiformes (wombats and koalas) and Phalangerida. Phalangerida is composed of the kangaroos (Macropodidae, Potoroidae, and Hypsiprymnodontidae) and possums (Phalangeridae, Burramyidae, Petauridae, Pseudocheiridae, Tarsipedidae, and Acrobatidae). Much of the debate has focused on relationships among the families of possums and whether possums are monophyletic or paraphyletic. A limitation of previous investigations is that no study to date has investigated diprotodontian relationships using all genera. Here, we examine diprotodontian interrelationships using a nuclear multigene molecular data set representing all recognized extant diprotodontian genera. Maximum parsimony, maximum likelihood, and Bayesian methods were used to analyze sequence data obtained from protein-coding portions of ApoB, BRCA1, IRBP, Rag1, and vWF. We also applied a Bayesian relaxed molecular clock method to estimate times of divergence. Diprotodontia was rooted between Vombatiformes and Phalangerida. Within Phalangerida, the model-based methods strongly support possum paraphyly with Phalangeroidea (Burramyidae + Phalangeridae) grouping with the kangaroos (Macropodiformes) to the exclusion of Petauroidea (Tarsipedidae, Acrobatidae, Pseudocheiridae, and Petauridae). Within Petauroidea, Tarsipedidae grouped with both Petauridae and Pseudocheiridae to the exclusion of Acrobatidae. Our analyses also suggest that the diprotodontian genera Pseudochirops and Strigocuscus are paraphyletic and diphyletic, respectively, as currently recognized. Dating analyses suggest Diprotodontia diverged from other australidelphians in the late Paleocene to early Eocene with all interfamilial divergences occurring prior to the early Miocene except for the split between the Potoroidae and Macropodidae, which occurred sometime in the mid-Miocene. Ancestral state reconstructions using a Bayesian method suggest that the patagium evolved independently in the Acrobatidae, Petauridae, and Pseudocheiridae. Ancestral state reconstructions of ecological venue suggest that the ancestor of Diprotodontia was arboreal. Within Diprotodontia, the common ancestor of Macropodidae was reconstructed as terrestrial, suggesting that tree kangaroos (Dendrolagus) are secondarily arboreal.     

Molecular phylogeny of Old World swifts (Aves: Apodiformes, Apodidae, Apus and Tachymarptis) based on mitochondrial and nuclear markers

We provide a molecular phylogeny for Old World swifts of genera Apus and Tachymarptis (tribe Apodini) based on a taxon-complete sampling at the species level. Phylogenetic reconstructions were based on two mitochondrial (cytochrome b, 12S rRNA) and three nuclear markers (introns of fibrinogen and glyceraldehyde 3-phosphate dehydrogenase plus anonymous marker 12884) while the myoglobin intron 2 did not show any intergeneric variation or phylogenetic signal among the target taxa at all. In contrast to previous hypotheses, the two genera Apus and Tachymarptis were shown as reciprocally monophyletic in all reconstructions. Apus was consistently divided into three major clades: (1) East Asian clade of A. pacificus and A. acuticauda, (2) African-Asian clade of A. caffer, A. batesi, A. horus, A. affinis and A. nipalensis, (3) African-Palearctic clade of eight currently accepted species among which sequences of A. apus and A. pallidus clustered in a terminal crown clade. Phylogenetic signal of all four nuclear markers was extremely shallow within and among species of tribe Apodini and even among genera, such that intra- and intergeneric relationships of Apus, Tachymarptis and Cypsiurus were poorly resolved by nuclear data alone. Four species, A. pacificus, A. barbatus, A. affinis and A. caffer were consistently found to be paraphyletic with respect to their closest relatives and possible taxonomic consequences are discussed without giving particular recommendations due to limitations of sampling. Incomplete mitochondrial lineage sorting with cytochrome-b haplotypes shared among species and across large geographic distances was observed in two species pairs: A. affinis/A. nipalensis and A. apus/A. pallidus. Mitochondrial introgression caused by extant or past gene flow was ruled out as an explanation for the low interspecific differentiation in these two cases because all nuclear markers appeared to be highly unsorted among Apus species, too. Apparently, the two extant species pairs originated from very recent dispersal and/or speciation events. The currently accepted superspecies classification within Apus was not supported by our results.     

The origin of a 'true' worker caste in termites: mapping the real world on the phylogenetic tree

The evolution of the 'true' worker caste in termites is not decisively inferred by coding and mapping both this character and the foraging behaviour on a phylogenetic tree. Answering to Thompson et al. (2000, 2003), and with reference to Grandcolas and D'Haese (2002), we show that this indecisive inference depends on the correct consideration paid to the outgroups. These last ones could be non subsocial cockroaches, or some wood-eating subsocial cockroaches often considered misleadingly as living ancestors, or even any hemimetabolous insects, all of them would be unambiguously lacking 'true' worker caste and pseudergate caste and not showing the 'one-piece' life type foraging behaviour. These statements derive from observing, coding and mapping the real world on the tree without making ad hoc assumptions. In that respect, because termites do not exist in isolation, apart from the tree of life, mapping the character of interest on the tree must be applied to the outgroups as well.     

Microsatellite markers for two species of dampwood termites in the genus Zootermopsis (Isoptera: Termopsidae)

Dampwood termites in the genus Zootermopsis inhabit forested areas in western North America. To better understand the colony composition and breeding structure of Zootermopsis, we identified polymorphic microsatellite loci to use in population analysis. Microsatellite loci were isolated from Zootermopsis nevadensis nevadensis (Hagen); however, all primers amplified homologous loci in Zootermopsis angusticollis (Hagen) and Zootermopsis nevadensis nuttingi (Hagen). Twelve loci were polymorphic in one or more of the above subspecies and species. The number of alleles per locus ranged from one to six, with some allelic differences among subspecies and species. We are currently utilizing the microsatellite markers to investigate the population genetics of Zootermopsis.     

Characterization of gut bacterial community associated with worker and soldier castes of Globitermes sulphureus Haviland (Blattodea: Termitidae) using 16S rRNA metagenomic

Globitermes sulphureus is a major pest in coconut plantations and also a secondary pest in the urban, suburban and rural areas of building structures, as well as in tropical agriculture areas. Although the literature revealed microbiome in termite guts, the bacterial community in G. sulphureus intestinal tracts remain largely unidentified. Here, we aimed to characterize the bacterial community associated with the worker and soldier castes of G. sulphureus using 16S metagenomic. The Illumina HiSeq 2500 sequencing machine was used to amplify the V3 and V4 regions of the 16S rRNA gene. The sequencing output was analyzed using Qiime pipeline v1.9.1. The result of the analysis showed that Spirochaetes, Fibrobacteres, Firmicutes, Proteobacteria, and Bacteroidetes were the dominant phyla that reside in the guts of the worker and soldier castes. We found that the worker and soldier castes shared similar bacterial phyla in their guts but different bacterial genera. Our findings provided a baseline information on microbial community inhabiting G. sulphureus guts up to the genus level for some phyla.     

Cockroach mating behaviors,sex pheromones,and abdominal glands (Dictyoptera: Blaberidae)

Two chemical signals are essential in all cockroach sexual behavioral sequences: the sex pheromone released by one partner, generally the female (for long distance attraction), and an aphrodisiac sex pheromone produced exclusively by male tergal glands (for female mounting and tergal contact or feeding behavior). Unlike the other cockroach groups, the males of the Oxyhaloinae species produce both chemical signals: the pheromone and the aphrodisiac. The occurrence of three patterns of mating behavior (A, B, and C), the production of male sex pheromones, and the existence in the male of developed sternal and tergal glands in seven related Oxyhaloinae species, make these cockroaches a useful model for studying the evolution of mating behavior patterns. The various types of mating behavior were not classified in the previous studies by Roth and Barth. In this report, they have been named type A (female in upper position), B (male in upper position), and C (male and female end to end). In type A mating, the male tergal glands, which are licked by the females, are well developed, whereas in types B and C, there is no licking of the male's tergal secretion by the females and the tergal glands are much less developed; the aphrodisiacs secreted by the tergal glands may no longer act in this case through contact chemoreception, but through an olfactory process involving volatile components. One common sex pheromone component seems to be acetoin. I suggest that the mating behavior tends from A toward B and C during the evolutionary process with a concomitant regression of the tergal glands and changes in the aphrodisiac emission levels. The mating behavioral sequences of cockroaches (Dictyoptera) and crickets (Orthoptera) show a striking degree of similarity and are probably examples of convergent evolution.     

Molecular systematics of racers, whipsnakes and relatives (Reptilia: Colubridae) using mitochondrial and nuclear markers

Four protein-encoding mitochondrial genes (cytochrome b , NADH-dehydrogenase subunits 1, 2 and 4) and one nuclear (c-mos) gene were sequenced to infer phylogenetic relationships among Old and New World representatives of racers and whipsnakes, Coluber (sensu lato). New World Coluber ( Coluber sensu stricto, including Masticophis ) and Salvadora proved to have affinities with the Old World non-racer colubrine genus Ptyas (and possibly Elaphe s.l. and Coronella ), whereas Old World ' Coluber ' form several basally related clades; these are (1) Hemorrhois -( Spalerosophis-Platyceps ); (2) Hierophis , with Eirenis nested within this paraphyletic genus and (3) ' Coluber ' dorri as the sister taxon to Macroprotodon cucullatus . The position of ' Coluber ' zebrinus along with Hemerophis socotrae located at the base of the Old World racer radiation forming the possible sister group to all remaining Palearctic racers and whipsnakes remains less well supported. Nevertheless, inter- and subgeneric relationships among many of the Old World racer groups have been resolved.     

Phylogenetic relationships among families of Gadiformes (Teleostei, Paracanthopterygii) based on nuclear and mitochondrial data

Phylogenetic hypotheses among Gadiformes fishes at the suborder, family, and subfamily levels are controversial. To address this problem, we analyze nuclear and mitochondrial DNA (mtDNA) sequences for the most extensive taxonomic sampling compiled to date, representing all of the recognized families and subfamilies in the order (except the monotypic family Lyconidae). Our study sampled 117 species from 46 genera, comprising around 20% of the species described for the order (more than 60% of all genera in the order) and produced 2740 bp of DNA sequence data for each species. Our analysis was successful in confirming the monophyly of Gadiformes and most of the proposed families for the order, but alternative hypotheses of sister-group relationships among families were poorly resolved. Our results are consistent with dividing Gadiformes into 12 families in three suborders, Muraenolepidoidei, Macrouroidei, and Gadoidei. Muraenolepidoidei contains the single family Muraenolepididae. The suborder Macrouroidei includes at least three families: Macrouridae, Macruronidae and Steindachneriidae. Macrouridae is deeply divided into two well-supported subfamilies: Macrourinae and Bathygadinae, suggesting that Bathygadinae may be ranked at the family level. The suborder Gadoidei includes the families: Merlucciidae, Melanonidae, Euclichthyidae, Gadidae, Ranicipitidae, and Bregmacerotidae. Additionally, Trachyrincinae could be ranked at family level including two subfamilies: Trachyrincinae and Macrouroidinae within Gadoidei. Further taxonomic sampling and sequencing efforts are needed in order to corroborate these relationships.