The social biology of domiciliary cockroaches: colony structure, kin recognition and collective decisions

A substantial body of research on eusocial insects seen in the last decades has gone hand-in-hand with the development of social evolution theory. In contrast, little attention has been given to the non-eusocial insect species that nevertheless exhibit a rich spectrum of social behaviours, thus effectively skewing our vision of insect sociality. Recent studies on the behaviour, ecology and genetic of “gregarious” cockroaches (Blattodea) have revealed a diversity of social structures and group dynamics unique among insects, providing an important comparative model for the broader understanding of insect social evolution. Here, we present an overview of the social biology of the domiciliary cockroaches (ca. 25 species adapted to human habitats) based on research on two model species, Blattella germanica and Periplaneta americana. We discuss the evolution of these domiciliary cockroaches, considering them in the context of “social herds” within the insect sociality framework.     

Patterns of aggregation behaviour in six species of cockroach: comparing two experimental approaches

Cockroaches (Blattaria) are considered universally gregarious insects. In this study, we tested the hypothesis that even related species and/or males and females of the same species may exhibit different aggregation patterns. We compared aggregation behaviour in six cockroach species belonging to three groups: Eublaberus, Blaberus, and Gromphadorhina‐Princisia (all Blaberidae). For comparison, binary choice and free aggregation arena tests were used. The results revealed that blaberid clades clearly differed in their aggregation patterns. Females of Eublaberus species aggregated, whereas only a low aggregation tendency was recorded in conspecific males. Both sexes of Blaberus species aggregated, especially when tested in the arena. The Madagascar hissing cockroaches of the Gromphadorhina‐Princisia clade are not gregarious – the aggregation tendency was entirely absent in males of the two species studied and in females of Princisia vanwaerebeki van Herrewege, and was only weakly demonstrated in females of Gromphadorhina portentosa (Schaum). The observed patterns of aggregation are compared with previous reports concerning social behaviour of blaberid species.     

Jaburetox,a urease-derived peptide: Effects on enzymatic pathways of the cockroach Nauphoeta cinerea

Jaburetox is a recombinant peptide derived from one of the Canavalia ensiformis urease isoforms. This peptide induces several toxic effects on insects of different orders, including interference on muscle contractility in cockroaches, modulation of UDP-N-acetylglucosamine pyrophosphorylase (UAP) and nitric oxide synthase (NOS) activities in the central nervous system of triatomines, as well as activation of the immune system in Rhodnius prolixus. When injected, the peptide is lethal for R. prolixus and Triatoma infestans. Here, we evaluated Jaburetox toxicity to Nauphoeta cinerea cockroaches, exploring the effects on the central nervous system through the activities of UAP, NOS, acid phosphatases (ACP), and acetylcholinesterase (AChE). The results indicated that N. cinerea is not susceptible to the lethal effect of the peptide. Moreover, both in vivo and in vitro treatments with Jaburetox inhibited NOS activity, without modifying the protein levels. No alterations on ACP activity were observed. In addition, the enzyme activity of UAP only had its activity affected at 18 hr after injection. The peptide increased the AChE activity, suggesting a mechanism involved in overcoming the toxic effects. In conclusion, our findings indicate that Jaburetox affects the nitrinergic signaling as well as the AChE and UAP activities and establishes N. cinerea as a Jaburetox-resistant model for future comparative studies.     

Laboratory and field evaluation of Beauveria bassiana bait against two cockroach species (Dictyoptera: Blattellidae,Blattidae) in Jinan City,East China

A Beauveria bassiana bait formulation was evaluated for its efficacy in the laboratory and field against German cockroaches, Blattella germanica (L.), and American cockroaches, Periplaneta americana (L.). The LT₅₀ values were 46 and 119?h, respectively, for B. germanica and P. americana. The cockroach species were lured and killed effectively when linear distance was about 5?m between B. bassiana bait and fresh mouse food. In the field, more than 90% relative decline in density was found 14?d after treatment, which lasted for 90?d. In conclusion, laboratory and field studies demonstrated the high efficacy of a B. bassiana bait formulation against common cockroach species.     

Latitudinal pattern in body size in a cockroach,Eupolyphaga sinensis

Body size of insects with flexible life cycles is expected to conform to the saw‐tooth model, a model in which the relationship between size and developmental time depends on length of the growing season. In species with high variability in terms of voltinism, however, more complex patterns can be expected. Few empirical studies have demonstrated the existence of such relationships, or whether climatic factors contribute to these relationships. In this study, we investigated the geographic variation in body size of the Chinese cockroach, Eupolyphaga sinensis Walker (Blattaria: Polyphagidae), which has a variable life cycle length. The sizes of adults – collected from 14 localities ranging from temperate to subtropical zones in China – were measured, using body length, body width, and pronotum width as parameters. The relationship between size, latitude, and climate factors (encompassing 10 variables) was then investigated. We found that the body size of E. sinensis varied considerably with latitude: cockroaches were larger at low and high latitudes, but smaller at intermediate latitudes. Thus, the relationship between climate and body size conformed to a saw‐tooth pattern. Results indicate that two factors were significantly associated with body size clines: season length and variability in life cycle length. Our results also demonstrated that climatic factors contribute to latitudinal clines in body size, which has important ecological and evolutionary implications. It can be expected that global climate change may alter latitudinal clines in body size of E. sinensis.     

Comparative application of the coefficient of variation and range-based statistics for assessing the taxonomic composition of fossil samples

Dental variation has been used commonly to assess taxonomic composition in morphologically homogeneous fossil samples. While the coefficient of variation (CV) has been used traditionally, range-based measures of variation, such as the range as a percentage of the mean (R%) and the maximum/minimum index (I_max/min) have recently become popular alternatives. The current study compares the performance of these statistics when applied to single- and pooled-species dental samples of extant Cercopithecus species. A common methodology for such problems of species discrimination has been to simply compare the maximum value of a variation statistic observed in extant samples with that observed in the fossil sample. However, regardless of what statistic is used, this approach has an unknowable Type I error rate, and usually has low power to detect multiple species. A more appropriate method involves a formal hypothesis test. The null hypothesis is that the level of variation in the fossil sample does not exceed what might be expected in a sample drawn randomly from a reference population, taking into account sampling error and the size of the fossil sample. Previous research using this method with the CV has indicated that it offers considerable power at an acceptable Type I error rate. In the current study, the data of primary interest were posterior dental dimensions for single- and pooled species samples from extant Cercopithecus species. In addition, the study also investigated the relative performance of variation statistics when applied to highly dimorphic canine dimensions, since much recent work has employed sexually dimorphic dental dimensions for assessing single-species hypotheses. The results indicate that the CV consistently out-performed the range-based statistics when using posterior dental dimensions to test a single-species hypothesis. Regardless of which statistic was used, tests on sexually dimorphic dimensions offered minimal power. In consideration of these results and the problem of studywise Type I error rates, we recommend against the use of multiple measures of variation to test for multiple species composition, and advocate the CV as the statistic of choice when using the method of Cope & Lacy (1992). For similar reasons, we argue for careful selection of dental variables for inclusion in such analyses, and in particular recommend against including sexually dimorphic dimensions when testing for multiple species composition.     

Cookeolus spinolacrymatus sp. nov., a Late Pliocene priacanthid fish from Okinawa, Japan

A new fossil priacanthid fish,Cookeolus spinolacrymatus, is described on the basis of a single specimen from the Late Pliocene Shinzato Formation, Shimajiri Group, Miyagi-shima, Okinawa, Japan. This species is distinguished from the Recent speciesC. japonicus (Cuvier, 1828) by having a strong spine on the anterior portion of the lacrymal. The specimen was identified as a small benthic adult, or large juvenile, based on comparison with pelvic fin length development in the Recent species.     

Interspecific competition among urban cockroach species

The aim of this study was to quantify and to compare the effects of intraspecific and interspecific competition among pairs of urban cockroaches [Blatta orientalis L., Periplaneta americana (L.), and Periplaneta australasiae Fabricius (Dictyoptera: Blattodea)] in relation to the limitation of resources such as shelter or food. Our approach was to assess whether the presence of one species affected the resource exploitation of another. A reduction in access to shelters or to food for one species revealed dominant/subordination relationships among species and induced spatial segregation in shelters and temporal segregation during food exploitation. The fragmentation of available resources facilitated spatial segregation and the access of more individuals of the subordinate species. Individuals of each species were aware of the presence of non‐conspecifics. The outcome of interspecific competition under laboratory conditions should help us to understand how segregated spatial distributions occur under natural conditions in urban areas.     

Five Recent Mya species, including three new species and their fossil connections

Three new species of Mya Linné are described from Greenland. Mya eideri sp. nov. is figured and compared with boreal and fossil species with which it has been confused, and the known distribution of the species is indicated. The validity in Recent faunas of the fossil species M. pseudoarenaria and M. uddevallensis is discussed; two new Recent species, M. neoovata sp. nov. and M. neouddevallensis sp. nov., from Ikka Fjord, southwest Greenland, and Scoresbysund, East Greenland, respectively, are described for Recent material of the comparable two fossil taxa. Accepted: 2 May 1999     

Polymorphic population genetic markers for the Australian wood cockroach Panesthia australis

Primers for five polymorphic nuclear DNA (nDNA) markers and one mitochondrial DNA (mtDNA) gene (COI) were developed for the Australian wood cockroach Panesthia australis using a range of techniques. Eight mitochondrial haplotypes and four to 36 alleles per nuclear locus were detected in 744 cockroaches. Observed heterozygosity ranged from 0.117 to 0.816 in a sample of 30 animals from one population. The markers will be useful for population biology and for the measurement of the effects of habitat fragmentation on this ecologically important forest‐dependent species.     

Impacts of Paleo-Oxygen Levels on the Size, Development, Reproduction, and Tracheal Systems of Blatella germanica

Atmospheric oxygen has varied substantially over the Phanerozoic (the last 500 million years) with periods of both hyperoxia and hypoxia relative to today. Unlike some insect groups, cockroaches have not been reported to exhibit gigantism during the late Paleozoic period of hyperoxia. Studies with modern insects have shown a diversity of developmental responses to oxygen, suggesting that evaluation of historical hypotheses should focus on groups most closely related to those present in the Paleozoic. Here we investigated the impacts of Paleozoic oxygen levels (12–31%) on the development of Blatella germanica cockroaches. Body size decreased strongly in hypoxia, but was only mildly affected by hyperoxia. Development time, growth rate and fecundity were negatively impacted by both hypoxia and hyperoxia. Tracheal volumes were inversely proportional to rearing oxygen, suggesting developmental responses aimed at regulating internal oxygen level. The results of these experiments on a modern species are consistent with the fossil record and suggest that changes in atmospheric oxygen would be challenging for many insects, despite plastic compensatory responses in the tracheal system.     

Phylogeny of ensign scale insects (Hemiptera: Coccoidea: Ortheziidae) based on the morphology of Recent and fossil females

The Ortheziidae (ensign scale insects) is a morphologically well‐defined family. The morphology and occurrence in the fossil record suggests a probable early origin of the family in scale insect evolution. The present phylogenetic analysis – based on 69 morphological characters of female ortheziids, using 39 exemplar Recent species – provides the first analytical assessment of relationships among living genera of the family, as well as the relationships of eight fossil species, based on complete, well‐preserved specimens in amber. Monophyly of the subfamilies Newsteadiinae, Ortheziinae and Ortheziolinae is supported, but Nipponortheziinae is found to be paraphyletic by inclusion of the Ortheziolinae. Thus, the subfamily Ortheziolinae is reduced in rank to tribe Ortheziolini stat.n. , which now includes Matileortheziola Kozár & Foldi, Ortheziolacoccus Kozár, Ortheziolamameti Kozár and Ortheziola?ulc. Consequently, the tribes Matileortheziolini, Ortheziolacoccini and Ortheziolamametini are synonymized ( syn.n. ) here under Ortheziolini. Five new species and one new genus of fossil ensign scales are described from three amber deposits: Burmorthezia gen.n. with type species Burmorthezia kotejai sp.n. and also B. insolita sp.n ., both in mid‐Cretaceous Burmese amber (98 Ma) and Arctorthezia baltica sp.n. in Eocene Baltic amber (c. 43 Ma) based on second‐instar nymphs; Mixorthezia kozari sp.n . and M. dominicana sp.n . in Miocene Dominican amber (c. 17 Ma) based on adult females. Fossil placements are unambiguous, with Burmorthezia forming a stem to crown‐group (Recent and Tertiary) Ortheziidae. A summary of described fossil ortheziids is provided.     

Monophyletic origin of Caryophyllia (Scleractinia,Caryophylliidae), with descriptions of six new species

The genus Caryophyllia Lamarck, 1816 is the most diverse genus within the azooxanthellate Scleractinia comprising 66 Recent species and a purported 195 nominal fossil species. Examination of part of the deep-sea scleractinian collection made by the Paris Museum off New Caledonia and part of the material collected by CSIRO off Australian waters revealed the occurrence of 23 species of Caryophyllia, of which six are new to science. All new records, including the new species, are described, and synonyms, distribution, type locality, type material and illustration are provided for each species. An identification key to all Recent species of Caryophyllia is presented. In addition, the validity of the genus Caryophyllia was investigated by phylogenetic analyses of a dataset consisting of partial mitochondrial 16S rRNA sequences from 12 species assigned to this genus together with seven species representing some of the most morphologically similar caryophylliid genera, and 14 non-caryophyllid species representing 14 scleractinian families. Irrespective of the method of analysis employed, all of the Caryophyllia species formed a well-supported clade together with Dasmosmilia lymani and Crispatotrochus rugosus. Although based on a subset of the Recent Caryophyllia species, these results are consistent with Caryophyllia being a valid genus, but call for a reexamination of Dasmosmilia and Crispatotrochus.     

Detection of cockroach faeces: consumption of fluorescent bait and production of UV‐light‐detectable faeces from German cockroach,Blattella germanica

Cockroaches have long been recognized as a major source of allergens in the human environment. As some allergens produced by cockroaches are accumulated within their tiny faeces and may persist unseen in the environment, development of a reliable method for monitoring cockroach faeces distribution and abundance is crucial. Our aim was to explore the rate of consumption of fluorescent bait and defecation of UV‐light‐detectable fluorescent faeces from Blattella germanica (L.) (Blattodea: Ectobiidae) as a prerequisite for the establishment of faeces monitoring in the field. Laboratory experiments revealed that (1) the fluorescent bait was accepted by cockroaches as a food source and faeces produced after its consumption were fluorescent as well; (2) there was a positive relationship between the fluorescent bait consumed and production of fluorescent faeces; (3) single and brief consumption of fluorescent bait led to continuous fluorescent faeces production, giving cockroaches enough time (ca. 27 h) to deposit fluorescent faeces in an entire area of cockroach activity as well as in shelters. The amount of fluorescent bait consumed and the amount of fluorescent faeces produced was nevertheless different among males, non‐gravid females, and gravid females. We subsequently validated the new fluorescent faeces‐based monitoring technique in a semi‐field experiment with human observers, which revealed that fluorescent faeces were reliably identified and distinguished from environmental dirt when illuminated with a UV flashlight. We further discuss the biological perspectives of our findings and recommendations for practical use.     

Influence of carbon dioxide gas on German cockroach (Dictyoptera: Blattellidae) knockdown, recovery, movement and feeding

Abstract. Carbon dioxide anaesthesia differentially affects the knockdown and recovery of cockroaches, depending on the strain and the length of time that the colony has been subjected to a CO₂ regime. Adult males from two laboratory and two field-collected strains of German cockroaches, Blattella germanica, are knocked down within 7–45 s after exposure to CO₂. After 5 min of CO₂ exposure, presumptive recovery (i.e. the time for the cockroach to right itself after knockdown) for laboratory strains occurs significantly sooner than for field-collected strains. Control cockroaches, exposed to compressed air rather than CO₂, exit harbourage cups rapidly (≤3.20 min). However, although allowed a recovery period of 5 min, significant movement impairment occurs for all cockroach strains anaesthetized with CO₂. Carbon dioxide exposure significantly reduces consumption of 2.15% hydramethylnon bait and delays mortality even when 24 h is allowed for recovery before bait placement. Cockroaches allowed to recover for 48 h after 5 min of CO₂ exposure consume significantly more bait and die significantly faster than CO₂ exposed groups allowed 24 h of recovery, and mortality is not significantly different from nonanaesthetized bait-fed controls.     

First records of the parthenogenetic Surinam cockroach Pycnoscelus surinamensis (Insecta: Blattodea: Blaberidae) for Central Europe

Sixteen species of cockroaches have been reported for Austria so far. This study is the first record of the parthenogenetic Surinam cockroach, Pycnoscelus surinamensis (L.) for Austria (and thus Central Europe). The species is natively distributed in Indo‐Malaysia but has been unintentionally introduced in many, mainly tropical, countries throughout the world. Sequencing the DNA barcoding region revealed that all Austrian P. surinamensis samples had the same haplotype, which they shared with samples from the United States of America, Guyana and French Polynesia, indicating that all these samples/populations belong to the same clonal lineage. Even though in temperate regions, the occurrence of P. surinamensis is currently limited to greenhouses, we advocate proper monitoring of the populations with respect to global warming and the expected increasing independence of this species from greenhouses that comes along with it.     

Differential consumption of baits by pest blattid and blattellid cockroaches and resulting direct and secondary effects

Consumption was measured of three commercially available cockroach gel baits (0.01 and 0.05% fipronil and 0.6% indoxacarb) by two pest blattellid (German cockroach, Blattella germanica L., and brownbanded cockroach, Supella longipalpa Fabricius) and three pest blattid [oriental cockroach, Blatta orientalis L., American cockroach, Periplaneta americana L., and smokybrown cockroach, Periplaneta fuliginosa (Serville)] species (Dictyoptera), and direct and secondary effects were quantified. All three baits were greatly preferred for consumption over dog food; however, virtually all consumption (ca. 98%) by pest blattids was gel baits containing sugar feeding stimulants and water. Pest blattid greater preference for gel baits was probably due to their greater need for nutrients in baits due to their greater cuticular water permeability and higher metabolism than the pest blattellids. Brownbanded cockroaches had lowest percentage gel bait selection. Pest blattellids consumed greater amounts of bait per g body weight than pest blattids. Cockroaches consumed more active ingredient than needed to cause mortality; however, based on bait consumption, a 30‐g tube of gel bait potentially killed from 394 to 6 966 adult cockroaches, depending on species. Mortality for all cockroach species was faster for adults (≥3 days) than for nymphs (≥7 days); however, most brownbanded cockroaches exposed to indoxacarb survived despite consuming 1.5‐ to >3‐fold more than other baits, suggesting low enzyme production by brownbanded cockroaches and consequently lower conversion of indoxacarb into its toxic form. Besides direct mortality, German cockroaches died from indirect effects: exposure to debris from other cockroaches that had direct access to the gel baits or bait contact without ingestion. Although maximization of bait consumption is important, factors that enhance secondary mortality and contact toxicity should also be considered.     

Sense organs on the antennal flagellum of a giant cockroach, Gromphadorhina portentosa, and a comparison with those of several other species (Dictyoptera, Blattaria)

Two kinds of tactile hairs, plain and wavy, thick-walled chemoreceptors and two types of thin-walled chemoreceptors are present on the antennal flagellum of Gromphadorhina portentosa males. These are described and their location on the antenna noted. Females of this species have plain tactile hairs and the same types of chemoreceptors as do the males but wavy tactile hairs are absent. The antennal sense organs of a few specimens of five other species of cockroaches — Periplaneta americana, Blatta orientalis, Supella longipalpa, Pycnoscelus surinamensis and Diploptera punctata — were also examined. All lacked tactile hairs but were provided with thick-walled chemoreceptors and with two types of thin-walled chemoreceptors similar to those described for Gromphadorhina portentosa.