Phylogenetic analysis of Blattabacterium,endosymbiotic bacteria from the wood roach,Cryptocercus (Blattodea: Cryptocercidae), including a description of three new species

Members of the cockroach genus Cryptocercus are wood-feeding, subsocial insects that live in temperate forests of the Nearctic and Palaearctic. At present, nine species are recognized: Cryptocercus relictus and Cryptocercus kyebangensis in eastern Asia and Russia, Cryptocercus primarius and Cryptocercus matilei in southwestern China, Cryptocercus clevelandi in the western USA, and Cryptocercus darwini, Cryptocercus garciai, Cryptocercus punctulatus, and Cryptocercus wrighti in the eastern USA. Like all extant cockroaches, Cryptocercus harbor endosymbiotic bacteria, Blattabacterium, in their fat bodies. The endosymbionts in all cockroaches have been considered a single species, Blattabacterium cuenoti, since their discovery about a century ago. However, a recent analysis of DNA sequences from representatives of four cockroach families has indicated that there is considerable DNA sequence divergence among B. cuenoti from different host species. As a part of our studies on the evolution of Cryptocercus, we examined DNA sequence divergence among B. cuenoti from six of the nine known Cryptocercus species. Specifically, we sequenced approximately 2,400 bp of the 16S rRNA and 23S rRNA genes of B. cuenoti from six species of Cryptocercus. We found that B. cuenoti in Cryptocercus has differentiated into multiple monophyletic lineages distinguishable by DNA sequence of rRNA genes and host association. Our sequence divergence estimates were consistent with those reported for other, congeneric bacterial species. We propose the recognition of three new species of Blattabacterium within Cryptocercus species as follows: Blattabacterium relictus sp. nov. in C. relictus, Blattabacterium clevelandi sp. nov. in C. clevelandi, and Blattabacterium punctulatus sp. nov. in C. darwini, C. garciai, C. punctulatus, and C. wrighti.     

Evidence for cocladogenesis between diverse dictyopteran lineages and their intracellular endosymbionts

Bacteria of the genus Blattabacterium are intracellular symbionts that reside in specialized cells of cockroaches and the termite Mastotermes darwiniensis. They appear to be obligate mutualists, and are transmitted vertically in the eggs. Such characteristics are expected to lead to equivalent phylogenies for host and symbiont, and we tested this hypothesis using recently accumulated data on relationships among termites and cockroaches and their Blattabacterium spp. Host and symbiont topologies were found to be highly similar, and various tests indicated that they were not statistically different. A close relationship between endosymbionts from termites and members of the wood-feeding cockroach genus Cryptocercus was found, supporting the hypothesis that the former evolved from subsocial, wood-dwelling cockroaches. The majority of the Blattabacterium spp. sequences appear to have undergone similar rates of evolution since their divergence from a common ancestor, and an estimate of this rate was determined based on early Cretaceous host fossils. The results support the idea that the stem group of modern cockroaches radiated sometime between the late Jurassic and early Cretaceous-not the Carboniferous, as has been suggested on the basis of roach-like fossils from this epoch.     

Evolution and phylogenetic information content of the ribosomal DNA repeat unit in the Blattodea (Insecta)

The organization, structure, and nucleotide variability of the ribosomal repeat unit was compared among families, genera, and species of cockroaches (Insecta:Blattodea). Sequence comparisons and molecular phylogenetic analyses were used to describe rDNA repeat unit variation at differing taxonomic levels. A reverse similar 1200 bp fragment of the 28S rDNA sequence was assessed for its potential utility in reconstructing higher-level phylogenetic relationships in cockroaches. Parsimony and maximum likelihood analyses of these data strongly support the expected pattern of relationships among cockroach groups. The examined 5' end of the 28S rDNA is shown to be an informative marker for larger studies of cockroach phylogeny. Comparative analysis of the nucleotide sequences of the rDNA internal transcribed spacers (ITS1 and ITS2) among closely related species of Blattella and Periplaneta reveals that ITS sequences can vary widely in primary sequence, length, and folding pattern. Secondary structure estimates for the ITS region of Blattella species indicate that variation in this spacer region can also influence the folding pattern of the 5.8S subunit. These results support the idea that ITS sequences play an important role in the stability and function of the rRNA cluster.     

The bacterial community in the gut of the Cockroach Shelfordella lateralis reflects the close evolutionary relatedness of cockroaches and termites

Termites and cockroaches are closely related, with molecular phylogenetic analyses even placing termites within the radiation of cockroaches. The intestinal tract of wood-feeding termites harbors a remarkably diverse microbial community that is essential for the digestion of lignocellulose. However, surprisingly little is known about the gut microbiota of their closest relatives, the omnivorous cockroaches. Here, we present a combined characterization of physiological parameters, metabolic activities, and bacterial microbiota in the gut of Shelfordella lateralis, a representative of the cockroach family Blattidae, the sister group of termites. We compared the bacterial communities within each gut compartment using terminal-restriction fragment length polymorphism (T-RFLP) analysis and made a 16S rRNA gene clone library of the microbiota in the colon-the dilated part of the hindgut with the highest density and diversity of bacteria. The colonic community was dominated by members of the Bacteroidetes, Firmicutes (mainly Clostridia), and some Deltaproteobacteria. Spirochaetes and Fibrobacteres, which are abundant members of termite gut communities, were conspicuously absent. Nevertheless, detailed phylogenetic analysis revealed that many of the clones from the cockroach colon clustered with sequences previously obtained from the termite gut, which indicated that the composition of the bacterial community reflects at least in part the phylogeny of the host.     

Detection and distribution patterns of telomerase activity in insects.

Telomeres of most insects consist of pentanucleotide (TTAGG)n repeats, although the repeats are absent in Diptera and some other insect species, where the telomere regions are perhaps maintained without telomerase. To understand various and unusual telomere formation in insects, we have studied the characteristic features of a putative insect telomerase that has not been previously described. Using a modified telomeric repeat amplification protocol (TRAP), we first detected the telomerase activity in crickets, cockroaches and two Lepidopteran insects. The telomerase from crickets and cockroaches required dATP, dGTP and dTTP but not dCTP as a substrate and sequence analyses of the products of TRAP revealed that the (TTAGG)n repeats are synthesized by telomerase. The cockroach telomerase was detected both in somatic (fat body, muscle and neural tissues) and germ line (testis) cells, suggesting that expression of this enzyme is not regulated in a tissue-specific manner at an adult stage. While we detected high levels of telomerase activity in crickets and cockroaches, we could not detect activity in all tissues and cell cultures of the silkworm, Bombyx mori and in two Drosophila and one Sarcophaga cell lines. This supports the theory that Dipteran insects maintain their telomeres without telomerase.     

Phylogeny of cockroaches (Insecta,Dictyoptera, Blattodea), with placement of aberrant taxa and exploration of out‐group sampling

We addressed the phylogeny of cockroaches using DNA sequence data from a broad taxon sample of Dictyoptera and other non‐endopterygotan insect orders. We paid special attention to several taxa in which relationships are controversial, or where no molecular evidence has been used previously: Nocticolidae, a family of small, often cave‐dwelling cockroaches, has been suggested to be the sister group of the predaceous Mantodea or of the cockroach family Polyphagidae; Lamproblatta, traditionally placed in Blattidae, has recently been given family status and placed as sister to Polyphagidae; and Saltoblattella montistabularis Bohn, Picker, Klass & Colville, a jumping cockroach, which has not yet been included in any phylogenetic studies. We used mitochondrial (COI + COII and 16S) and nuclear (18S and 28S) genes, and analysed the data using Bayesian inference (BI) and maximum likelihood (ML). Nocticolidae was recovered as sister to Polyphagidae. Lamproblatta was recovered as sister to Blattidae, consistent with the traditional placement (not based on phylogenetic analysis). However, because of the limited support for this relationship and conflict with earlier morphology‐based phylogenetic hypotheses, we retain Lamproblattidae. S. montistabularis was consistently placed as sister to Ectobius sylvestris Poda (Blaberoidea: Ectobinae), indicating that the saltatorial hindlegs of this genus are a relatively recent adaptation. Isoptera was placed within Blattodea as sister to Cryptocercidae. Nocticolidae + Polyphagidae was sister to Isoptera + Cryptocercidae, and Blaberoidea was sister to the remaining Blattodea.     

Evidence from multiple gene sequences indicates that termites evolved from wood-feeding cockroaches

Despite more than half a century of research, the evolutionary origin of termites remains unresolved [1] [2] [3]. A clear picture of termite ancestry is crucial for understanding how these insects evolved eusociality, particularly because they lack the haplodiploid genetic system associated with eusocial evolution in bees, ants, wasps and thrips [4] [5]. Termites, together with cockroaches and praying mantids, constitute the order Dictyoptera, which has been the focus of numerous conflicting phylogenetic studies in recent decades [6] [7] [8] [9] [10] [11] [12]. With the aim of settling the debate over the sister-group of termites, we have determined the sequences of genes encoding 18S ribosomal RNA, mitochondrial cytochrome oxidase subunit II (COII) and endogenous endo-beta-1, 4-glucanase (EG) from a diverse range of dictyopterans. Maximum parsimony and likelihood analyses of these sequences revealed strong support for a clade consisting of termites and subsocial, wood-feeding cockroaches of the genus Cryptocercus. This clade is nested within a larger cockroach clade, implicating wood-feeding cockroaches as an evolutionary intermediate between primitive non-social taxa and eusocial termites.     

Inherited Fungal and Bacterial Endosymbionts of a Parasitic Wasp and Its Cockroach Host

Bacterial endosymbionts of insects are increasingly being recognized as common, diverse, and integral to the biology of their hosts. Inherited fungal symbionts have been largely overlooked, however, even though insect guts appear to be a key habitat for an incredible array of fungal diversity. Like bacteria, fungal symbionts also likely play important roles in the ecology and evolution of their insect associates. The objective of this study was to lay the foundations for understanding the roles of the vertically transmitted fungal and bacterial associates of both the brownbanded cockroach, Supella longipalpa, and its parasitic wasp, Comperia merceti. We used culture-dependent and culture-independent molecular methods and phylogenetic analyses in order to identify the symbionts. Two fungal associates of brownbanded cockroaches were found. To our knowledge, this is the first record of vertically transmitted fungal symbionts in the order Blattaria. The wasp was found to house a close relative of one of the cockroach fungi but no bacterial symbionts. Finally, the brownbanded cockroaches also harbored three lineages of bacterial symbionts: Blattabacterium and two lineages of Wolbachia, indicating the number of vertically transmitted symbionts in this insect may be as many as five.     

Susceptibility of cockroaches (Gromphadorhina portentosa,Nauphoeta cinerea and Blaptica dubia) exposed to entomopathogenic nematodes

Cockroaches are major pests, vectors of pathogenic bacteria and induce allergies. Current control methods use chemical pesticides, but they can be ineffective and costly and there are reports of resistance in the field; hence new control methods are needed. There are conflicting reports about the susceptibility of cockroaches to entomopathogenic nematodes (EPNs); so we investigated if EPNs could kill several diverse cockroach species, including the Madagascan hissing roach (Gromphadorhina portentosa), the Lobster roach (Nauphoeta cinerea) and Blaptica dubia. Female adult cockroaches were exposed to either commercial products containing Steinernema kraussei or a combination of Heterorhabditis spp. and Steinernema spp. at 50 and 150 nematodes per cm² for 21 days. We also monitored feeding and the numbers of infective juveniles that were produced from each cockroach corpse. We found that S. kraussei were harmless to all cockroach species (at both doses) but when exposed to a mixture of Heterorhabditis spp. and Steinernema spp. B. dubia died after 6 days and its feeding was strongly inhibited. We also found that the mixture of Heterorhabditis spp. and Steinernema spp. could proliferate in the cadavers of B. dubia whilst S. kraussei could only reproduce in G. portentosa and B. dubia but not N. cinerea. In conclusion, S. kraussei was harmless to all three cockroach species but B. dubia was killed when exposed to Heterorhabditis spp. and Steinernema spp., highlighting the differences in the host range of EPNs.     

Relationship between the para-homologous sodium channel point mutation (g --> c at nucleotide 2979) and knockdown resistance in the German cockroach using multiplex polymerase chain reaction to discern genotype

Extensive use of pyrethroid insecticides for urban pest control has led to widespread pyrethroid resistance in the German cockroach. A mutation at nucleotide position 2979 (G to C, causing a leucine to phenylalanine change) in the S6 transmembrane segment of domain II of the para-homologous voltage-gated sodium channel has been previously identified in knockdown-resistant cockroaches and demonstrated by site-directed mutagenesis to reduce channel sensitivity to pyrethroids. In a recent survey, 83% of pyrethroid-resistant German cockroach populations were found to possess this mutation. A German cockroach strain with a low incidence of the L993F mutation was subjected to selection pressure with cypermethrin and subsequently evaluated over several generations for the knockdown resistance phenotype. Correspondingly, we determined the genotype of individual cockroaches of each population at the 2979 position of the para-homologous gene. Genotype was discerned by development of a polymerase chain reaction method that employed a mismatched primer-template set. A direct relationship was observed between mean knockdown time and the presence of the kdr mutation. Furthermore, individuals homozygous for the kdr mutation exhibited a significantly higher mean knockdown time than heterozygotes or wildtype cockroaches. This is the first report demonstrating the progressive expression of the kdr allele in response to insecticide selection pressure.     

First molecular genetic evidence for automictic parthenogenesis in cockroaches

Parthenogenesis is an asexual mode of reproduction that plays an important role in the evolution of sex, sociality, and reproduction strategies in insects. Some species of cockroach exhibit thelytoky, a type of parthenogenesis in which female offspring are produced without fertilization. However, the cytological and genetic mecha? nisms of parthenogenesis in cockroaches are not well understood. Here we provide the first molecular genetic evidence that cockroaches can reproduce through automixis. Using the American cockroach Periplaneta aniericana, we performed microsatellite analysis to investigate the genetic relationship between parthenogenetically produced nymphs and the parent virgin females, and found that all parthenogenetic offspring were homozygous for autosomal microsatellite markers, whereas the female parents were heterozygous. In addition, flow cytometry analysis revealed that the parthenogenetic offspring were diploid. Taken together, our results demonstrate that P. americana exhibits automixis-type thelytoky, in which diploidy is restored by gamete duplication or terminal fusion. These findings highlight the unique reproduction strategies of cockroaches, which are more varied than was previously recognized.     

Structural data on hypertrehalosaemic neuropeptides from Cryptocercus punctulatus and Therea petiveriana: how do they fit into the phylogeny of cockroaches?

Hypertrehalosaemic neuropeptides from the corpora cardiaca of the cockroaches Cryptocercus punctulatus and Therea petiveriana were structurally analysed to gather phylogenetic information independent from that provided by morphoanatomical data. Isolation of the peptides by liquid chromatography and structural elucidation by Edman degradation and mass spectrometry revealed an identical octapeptide for both species: pGlu-Leu-Asn-Phe-Ser-Pro-Asn-Trp-NH₂. This peptide, denoted Tem-HrTH, was previously found in tenebrionid beetles and in the cockroach Polyphaga aegyptiaca. Using this information for phylogenetic analysis yielded a peptide tree that supports the previous morphoanatomical data and thus places the woodroach Cryptocercus inside the cockroach subfamily Polyphaginae.     

The molecular evolution of the allatostatin precursor in cockroaches

Allatostatins (ASTs) of the Tyr/Phe-Xaa-Phe-Gly Leu/Ile-NH2 family are a group of insect neuropeptides that inhibit juvenile hormone biosynthesis by the corpora allata. We have obtained genomic DNA sequences that specify the preproallatostatin precursor for the cockroaches, Blatta orientalis, Blattella germanica, Blaberus (cranufer and Supella longipalpa. The sequences obtained are similar to those of Diploptera punctata and Periplaneta americana reported previously. The precursors of all these cockroach species are similar in size, and the organization of the ASTs that they contain (there are 13 or 14, depending on the species) have been conserved. With the sequences of these precursors, and using the homologous sequence in the orthopteran Schistocera gregari as an outgroup, a phylogenetic analysis using parsimony was carried out. The dendrograms obtained from these analyses. using the amino acid as well as the nucleotide sequences, are comparable with current models for cockroach phylogeny. Parsimony analysis was also used to study the genealogy of the different ASTs within the same precursor. Results suggest that the AST sequences were generated through a process of internal gene duplication which occurred before these species diverged from each other in evolutionary time.     

Metabolic stasis in an ancient symbiosis: genome-scale metabolic networks from two <Emphasis Type="Italic">Blattabacterium cuenoti</Emphasis> strains,primary endosymbionts of cockroaches

BACKGROUND: Cockroaches are terrestrial insects that strikingly eliminate waste nitrogen as ammonia instead of uric acid. Blattabacterium cuenoti (Mercier 1906) strains Bge and Pam are the obligate primary endosymbionts of the cockroaches Blattella germanica and Periplaneta americana, respectively. The genomes of both bacterial endosymbionts have recently been sequenced, making possible a genome-scale constraint-based reconstruction of their metabolic networks. The mathematical expression of a metabolic network and the subsequent quantitative studies of phenotypic features by Flux Balance Analysis (FBA) represent an efficient functional approach to these uncultivable bacteria. RESULTS: We report the metabolic models of Blattabacterium strains Bge (iCG238) and Pam (iCG230), comprising 296 and 289 biochemical reactions, associated with 238 and 230 genes, and 364 and 358 metabolites, respectively. Both models reflect both the striking similarities and the singularities of these microorganisms. FBA was used to analyze the properties, potential and limits of the models, assuming some environmental constraints such as aerobic conditions and the net production of ammonia from these bacterial systems, as has been experimentally observed. In addition, in silico simulations with the iCG238 model have enabled a set of carbon and nitrogen sources to be defined, which would also support a viable phenotype in terms of biomass production in the strain Pam, which lacks the first three steps of the tricarboxylic acid cycle. FBA reveals a metabolic condition that renders these enzymatic steps dispensable, thus offering a possible evolutionary explanation for their elimination. We also confirm, by computational simulations, the fragility of the metabolic networks and their host dependence. CONCLUSIONS: The minimized Blattabacterium metabolic networks are surprisingly similar in strains Bge and Pam, after 140 million years of evolution of these endosymbionts in separate cockroach lineages. FBA performed on the reconstructed networks from the two bacteria helps to refine the functional analysis of the genomes enabling us to postulate how slightly different host metabolic contexts drove their parallel evolution.     

Classical olfactory conditioning in the cockroach Periplaneta americana

We established a classical conditioning procedure for the cockroach, Periplaneta americana, by which odors were associated with reward or punishment. Cockroaches underwent differential conditioning trials in which peppermint odor was associated with sucrose solution and vanilla odor was associated with saline solution. Odor preference of cockroaches was tested by allowing them to choose between peppermint and vanilla sources. Cockroaches that had undergone one set of differential conditioning trials exhibited a significantly greater preference for peppermint odor than did untrained cockroaches. Memory formed by three sets of differential conditioning trials, with an inter-trial interval of 5 min, was retained at least 4 days after conditioning. This conditioning procedure was effective even for cockroaches that had been harnessed in plastic tubes. This study shows, for the first time in hemimetaborous insects, that both freely moving and harnessed insects are capable of forming olfactory memory by classical conditioning procedure. This procedure may be useful for future electrophysiological and pharmacological studies aimed at elucidation of neural mechanisms underlying olfactory learning and memory.     

Pterygodermatites nycticebi infections in golden lion tamarins (Leontopithecus rosalia rosalia) and aye‐ayes (Daubentonia madagascariensis) from a German zoo

In a golden lion tamarin (Leontopithecus rosalia rosalia) colony kept indoors in a German zoo, two animals presented a sudden onset of reduced general condition, lethargy, and diarrhea. At animal capture for clinical examination, adult nematode stages were observed after stress‐induced defecation. Despite treatment, two golden lion tamarins died in the following 2 days. At necropsy, spirurid stages were found in the lungs and intestine. Additionally, adult Pterygodermatites spp. were identified in histopathological samples of intestine and pancreas, confirming the previous diagnosis. Upon diagnosis, all animals were treated with ivermectin (0.2 mg/kg; SC). Thereafter, the general condition of the golden lion tamarins improved, whereby some of them excreted spirurid nematodes over 3 days. Four weeks after treatment, 20 fecal samples from the colony were examined and proved negative for parasitic stages. Given that common German cockroaches (Blattella germanica) are suitable intermediate hosts of Pterygodermatites nycticebi, 30 specimens were collected from seven different locations around the golden lion tamarins housing. Third‐stage larvae of Pterygodermatites spp. were recovered from those cockroaches. Regular anthelmintic treatments, coprological screenings, and controls for intermediate hosts were recommended. More than 2 years later, P. nycticebi infection was diagnosed again histopathologically in an aye‐aye (Daubentonia madagascariensis) which suddenly died. Coprological analysis confirmed the presence of spirurid eggs. Due to prosimian primates' cockroach‐eating habits and given that total cockroach eradication proved impossible, continuous cockroach control strategies and regular treatments of primates are currently performed to prevent further P. nycticebi infections.     

The evolutionary transition from subsocial to eusocial behaviour in Dictyoptera: phylogenetic evidence for modification of the "shift-in-dependent-care" hypothesis with a new subsocial cockroach

Cockroaches have always been used to understand the first steps of social evolution in termites because they are close relatives with less complex and integrated social behaviour. Termites are all eusocial and ingroup comparative analysis would be useless to infer the origin of their social behaviour. The cockroach genus Cryptocercus was used as a so-called "prototermite" model because it shows key-attributes similar to the termites (except Termitidae): wood-feeding, intestinal flagellates and subsocial behaviour. In spite of these comparisons between this subsocial cockroach and eusocial termites, the early and remote origin of eusocial behaviour in termites is not well understood yet and the study of other relevant "prototermite" models is however needed. A molecular phylogenetic analysis was carried out to validate a new "prototermite" model, Parasphaeria boleiriana which shows a peculiar combination of these key-attributes. It shows that these attributes of Parasphaeria boleiriana have an independent origin from those of other wood-eating cockroaches and termites. The case of P. boleiriana suggests that a short brood care was selected for with life on an ephemeral wood resource, even with the need for transmission of flagellates. These new phylogenetic insights modify evolutionary hypotheses, contradicting the assumption made with Cryptocercus model that a long brood care is necessary for cooperation between broods in the "shift-in-dependent-care" hypothesis. An ephemeral wood resource is suggested to prompt generation overlap and the evolution of cooperation, even if brood care is shortened.     

Identification of novel periviscerokinins from single neurohaemal release sites in insects MS/MS fragmentation complemented by Edman degradation.

Three novel members of the periviscerokinin family could be identified directly from extracts of single abdominal perisympathetic organs of blaberoid cockroaches by means of electrospray ionization-quadrupole time of flight (ESI-QTOF) MS. Sequences of these periviscerokinins were confirmed by Edman degradation. Their primary structures are GSSGLIPFGRT-NH2 (Lem-PVK-1), GSSGLISMPRV-NH2 (Lem-PVK-2), and GSSGMIPFPRV-NH2 (Lem-PVK-3). Hitherto only known from the American cockroach, this neuropeptide family contains a highly conserved N-terminus whereas, at the C-terminus, only the penultimate amino-acid residue (Arg) has been found in all members of this peptide family. The identified periviscerokinins are the only abundant myoactive peptides in abdominal perisympathetic organs of blaberoid cockroches and they appear to be absent in the retrocerebral complex. Screening of extracts of single abdominal perisympathetic organs (70-90 microm in diameter), from five different species of the suborder Blaberoidea, revealed that they all contain the three neuropeptides which are described here for the first time.     

Juvenile hormone and methyl farnesoate production in cockroach embryos in relation to dorsal closure and the reproductive modes of different species of cockroaches

Juvenile hormone (JH), produced by the corpora allata (CA), is first detectable after dorsal closure, a conspicuous event in embryogenesis. The present research found that the timing of dorsal closure was consistently at about 45% of the total embryonic development time across most of the oviparous and ovoviviparous cockroach species examined. These included the ovoviviparous cockroaches Blaberus discoidalis, Byrsotria fumigata, Rhyparobia maderae, Nauphoeta cinerea, Phoetalia pallida, Schultesia lampyridiformis, and Panchlora nivea, as well as the oviparous cockroaches Blatta orientalis, Periplaneta americana, Eurycotis floridana, and Supella longipalpa. However, the only known viviparous cockroach Diploptera punctata completed dorsal closure at 20.8% of embryo development time. Methyl farnesoate (MF), the immediate precursor of JH III, is considered a functional molecule in crustaceans; however, in insects its function is still unclear. To understand the role of JH and MF in cockroach embryos, I compared JH and MF biosynthesis and release in several cockroach species of known phylogenetic relationships. Using a radiochemical assay, the present research showed that cockroach embryos representing all three reproductive modes produced and released both JH and MF, as previously shown for B. germanica, N. cinerea, and D. punctata. Members of a pair of embryonic CA from B. discoidalis, B. fumigata, R. maderae, and D. punctata were incubated with and without farnesol. MF accumulated in large amounts only in CA of R. maderae in the presence of farnesol, which indicates that control of the last step of biosynthesis of JH, conversion of MF into JH by MF epoxidase, is probably a rate-limiting step in this species.     

New case of true mimicry in cockroaches (Blattodea)

True mimicry is an extremely rare phenomenon in cockroaches, with beetles as their favorite models. So far, only very few Pseudophyllodromiinae could be identified that conform to all requirements for Batesian mimicry and similarly exist only isolated reports on Müllerian mimicry. The vast majority, however, of the so‐called beetle‐mimicking cockroaches only shows a high degree of generalized convergence regarding their outer appearance. Specific models or counterparts, respectively, are lacking. Here we report on a last instar nymph of an undescribed Blattinae (Blattodea: Blattidae) that accurately mimics a truly harmful, sympatric carabid beetle. Although no indications for inedibility of this cockroach became evident, thus making Batesian mimicry likely, Müllerian mimicry can not be excluded with certainty. At any rate, this represents both the first case of mimicry in Blattidae and the first case with a ground beetle (Carabidae) as model for a cockroach mimic.