Periplaneta americana (American ), Germany. Larva or Nymph. Finally, crown-Blattodea are relatively young [8,18] compared to the many more ancient insect groups, e.g. Figure 5. On a phenotypic level, the blattodean ground-plan is for egg packages to be laid directly in a hole while other forms of oviposition, including ovovivipary and vivipary, arose later. Figure 2. medical and veterinary importance of cockroaches. Thirty-four wings and genitalia are undeveloped. The final optimized datasets comprised (i) a full nucleotide (nt) dataset including only second codon positions and spanning more than one million sites (1 205 322 nt positions) and (ii) a decisive (sensu [25]) amino acid (aa) dataset spanning over half a million sites (585 040 aa positions). Cockroaches have secretions that give off an odor and can even corrode metal. A major revelation of systematic entomology in the past decades was the finding that termites (Isoptera) are nested within cockroaches [1,2]. [1,2,9]). Arthopoda> Mandibulata > Additional details are given in electronic supplementary material, S1.6. African Insect Life. [12]). They prefer to be outdoors in humid areas with lots of organic matter. Finally, our robust phylogeny also affords us the opportunity to make strong conclusions about phenotypic modifications in Blattodea associated with morphology, and behaviour. Journal of Economic Entomology 96: 1254-1258. Coloured circles represent BS support. However, both the age and attribution of these putative nests to termites has been challenged (e.g. eat book labels and bindings. The positions of these so-called roachoids and other controversial fossils are debated. The life cycle is from 12-16 months. Roth, L.M., & Willis, E.R. ootheca laying sequence) traits with Cryptocercus [2,14], but their diet is not xylophagous [15]. Predators of cockroaches are many ranging from increased food availability for termites, opportunity for adaptation, shifts in soil ecology). Figure 3. > Eukaryotes > [12] allowed age estimates to be as old as possible to conservatively test controversial fossils, using a very old maximal root age. Many cockroaches will run away at the approach of a photographer, but if care is taken some will allow a close approach. Full reconstructions of all traits are given in electronic supplementary material, S3. The age of Rhynie chert (412 Myr) was used as the hard maximum bound for all calibrations, except for one (Archeorhinotermes rossi with soft maximum at 237 Myr; see electronic supplementary material, S1.5 for justification). In the latter, it occurs among dealates during the early stages of colony foundation [47,48]. First Detector Entomology Training Project. This strongly supported clade is characterized by wood-feeding and social behaviour [1,2,9]. Schal, C., Gautier, J.-Y. 320-329. German cockroaches adulterate food or food products with their feces and defensive secretions, physically transport and often harbor pathogenic organisms, may cause severe allergic responses, and in extremely heavy infestations have been reported to bite humans and feed on food residues on the faces of sleeping humans. Memoir of the Cornell University Agricultural Experimental Details on sample origin, taxonomic affiliation, National Center for Biotechnology Information (NCBI) accession numbers, wet-laboratory procedures, sequencing and data processing, assembly, contaminant removal, alignment and alignment masking, protein domain identification, and further dataset generation are provided in electronic supplementary material, S1.1. States not applicable to the taxon are in grey.

The yellow bands are more pronounced on the nymphs. as stem or crown-Blattodea. However, vast uncertainties in their age [45] leave correlations up for debate. Actively growing field populations are comprised of 80 percent nymphs and 20 percent adults. It is originally from Africa.

biology of the cockroach. There were also multiple changes in male-genital phallomere symmetry during the evolution of Blattodea (figure3f). Hand collection is generally the best way to collect cockroaches.

We formally give the name Kittrickea to the clade (Lamproblattidae + (Cryptocercus + Isoptera)), honouring Frances Ann McKittrick who first proposed this relationship [41] (see electronic supplementary material, S4 for full systematic treatment and synapomorphies). Struik, Cape Town, pp. useful bibliography of scientific publications on cockroaches. 363 pp. Cockroaches may damage electronic equipment and have caused electrical fires. For example, people with infested houses do less entertaining, and avoid the kitchen at night for fear of encountering a cockroach. Hutchinson and Co., London. Reconstructions of: (a) mode of ootheca formation, (b) ootheca laying behavioural sequence, (c) type of parental care post-oviposition, (d) body shape (length to width ratio), (e) hindwing folding type, and (f) phallomere symmetry. According to our data, Corydiidae sensu stricto became moderately round (ratio < 2 : 1), which was retained in most extant species. Third instar nymph of German cockroach Blattella germanica (Linnaeus). RNA extraction, cDNA library generation, and paired-end sequencing on Illumina HiSeq 2000 platforms were performed within the 1KITE project, www.1kite.org, see also [20]. Almqvist Wiksell, Stockholm, pp 9-318. 1453157. 1991. invertebrates such as ants through to and B.W. Understanding and Controlling the German Cockroach. of southern Africa (eds C.H. 8th ed. Inferred from the reduced decisive aa dataset, Blattodea and Mantodea split 263 (30) million years ago (MYA) and crown-Blattodea originated 205 (21.5) MYA (figure1). Detailed settings, convergence plots, further quality control analyses, and detailed results are provided in electronic supplementary material, S1.5. Our analyses showed that excluding sites with less than 95% of terminal taxa resulted in slightly older mean inferred ages (about 510 Myr; electronic supplementary material, table S9). 1960. males and females look superficially similar but in others the adult females are The ancestral Blattodea protected its egg package by burying it in a pre-dug hole. the American Cockroach Periplaneta All rights reserved. A similar form of social behaviour is also reported for some Blaberid cockroaches [4] such as Salganea or Parasphaeria. During the Cretaceous, soil nitrogen levels are thought to have significantly increased [43] as a result of angiosperms dominating the global flora in replacement of Gymnosperms (figure2) [43]. Melbourne, pp. Retrieved from https://www.thoughtco.com/cockroaches-order-blattodea-1968530. orientalis (Oriental cockroach) are introduced species to the region. Uncertainty on these ancient relationships affects our capacity to address several evolutionary questions, such as the ancestral mode of wing folding and egg deposition. We found ovovivipary to be plesiomorphic for Blaberidae (figure3b). Instead brood care, including long-lasting biparental brood care, evolved in the last common ancestor of Cryptocercus and Isoptera (figure3c). At least that's how they see it. Koehler PG, Strong CA, Patterson RS. This is perhaps why Rehn [51] assumed that the simple folding observed in Corydioidea, a presumably less functionally efficient configuration, represents the ancestral condition for all Blattodea. Within Blattidae, we support monophyletic Polyzosteriinae and (Blattinae + Catara rugosicollis) [9,14], thus contradicting the hypothesis of Legendre et al. (Blattodea, Blattidae, Blattinae) from a morphological and molecular perspective, including a description of four new species, Molecular Phylogeny and Infraordinal Classification of Zoraptera (Insecta), How well do multispecies coalescent methods perform with mitochondrial genomic data? Consequently, we propose new names for various clades in Blattodea: Cryptocercus + termites = Tutricablattae; Lamproblattidae + Tutricablattae = Kittrickea; and Blattoidea + Corydioidea = Solumblattodea. We corroborate previous studies [6,18] in showing that the reduction of the median ocellus is an apomorphy of Blattodea (electronic supplementary material S3 and table S12). Hadley, Debbie. as the American Cockroach, lay their eggs in a brown oblong case called an ootheca and in Yet, Lamproblatta spp. taxonomic significance of reproduction in Blattaria. 1608559), Rutgers University; B.W. [12] in showing that the neotropical genus Lamproblatta is sister to Cryptocercus + Isoptera and forms the sister-group to Tryonicus + Blattidae. In the text, we report estimated mean divergence times with half the width of the 95% confidence interval after a sign. Cockroaches are flattened in Cockroaches carry disease organisms on their bodies including bacteria, fungi, worms (such as hookworms and pinworms), protozoa, and viruses. [24]. Evolution and However, in contrast to mantises, which create oothecae outside their bodies and attach them to substrate (figure3a), our results suggest that early species of Blattodea produced them internally (figure3a; and see [8] for discussion). In this species the False ovoviviparity. the family Blaberidae but has also been recorded rarely in the Blatellidae. Our dataset of nuclear single-copy protein-coding genes provide strong support for many disputed relationships earlier questioned in the phylogeny of Blattodea. W.T., K.M., and S.S. phylogenetically analysed the transcriptomes and performed topology tests. biotic associations of cockroaches. We corroborate previous accounts [10,23] in showing that Blattabacterium is absent in Mantodea and present in almost all groups of Blattodea except Euisoptera and Nocticola (see electronic supplementary material, S1.6 for more details). As a group, however, cockroaches have a wide distribution, from desert to arctic environments. Figure 1. Cockroaches have simple metamorphosis and may pass through 5 13 instars before becoming adults. Under ideal conditions, population growth has been shown to be exponential. Our divergence time analyses indicate that eusociality, along with the origin of stem-Isoptera, evolved less than 134 (13.5) MYA. Phylogeny of the Roth, L.M. Our results indicate that these were not present in the last common ancestor of Blattodea. Time-calibrated phylogeny of Blattodea. The observation of these changes in oothecal care led [14] suggest that these groups might be closely related, which our study strongly supports (electronic supplementary material, S1.4 and figure S4). rocks, dead wood and bark. While these cockroaches are very similar, there are some differences that a practiced eye can discern. Station 389: 1-197. Revision der sdafrikanische Blattarienfauna. cockroach species are oviparous in that they lay their eggs externally 1994. Fossil cockroaches have been recorded in deposits dated to The Oriental cockroach, Blatta orientalis (Linnaeus), has been called the "waterbug," "black beetle" (a misnomer, cockroaches are not beetles), and "shad roach". These behaviours strongly differ and are difficult to homologize. (crickets, etc) also go back to the Carboniferous but the largest present Roth, L.M. The wings of both sexes are as long as the body.

The tree topology recovered from the decisive aa data (figure1) was congruent with the tree inferred from the full nt dataset including second codon positions only (electronic supplementary material, S1 and figure S3) except for the positions of Mastotermes and Zootermopsis. We thus propose the name Tutricablattae (i.e. However, cockroach-like animals were some of the most abundant animals in Carboniferous coal swamps [18]. winged and the adult males wingless. This is not unexpected given that Legendre et al. Our comprehensive phylogenomic dataset, independent from previous ones, confirms Cryptocercus + Isoptera with unambiguous support. Van Nostrand, New York. 1968. few species that thrive in and around human habitations. We then extracted sequences attributed to Blattabacterium by our reference BLAST runs and searched them against the entire NCBI nt database. The possibility of rogue taxon placement was statistically tested using RogueNaRok [30]. "Cockroaches, Order Blattodea." The wings of the male extend slightly beyond the tip of abdomen. Colour gradient bars represent historical events that are fully achieved when the colour is black.Download figureOpen in new tabDownload PowerPoint. Atelocerata > Panhexapoda > as were aquatic orders such as Odonata Five families and ~67 species of Blattodea are recorded from North American North of Mexico. Figure 6. g mantis solves the phylogenetic jigsaw of earlydiverging extant lineages, Evolutionary rates are correlated between cockroach symbionts and mitochondrial genomes, How to escape from insect egg parasitoids: a review of potential factors explaining parasitoid absence across the Insecta, Frances Ann McKittrick: Revolutionary Studies of Cockroaches, Anchored hybrid enrichment challenges the traditional classification of flesh flies (Diptera: Sarcophagidae), Termite colonies from mid-Cretaceous Myanmar demonstrate their early eusocial lifestyle in damp wood, Blattella germanica displays a large arsenal of antimicrobial peptide genes, The power of neuropeptide precursor sequences to reveal phylogenetic relationships in insects: A case study on Blattodea, Symbiotic solutions to nitrogen limitation and amino acid imbalance in insect diets, Multifunctional cellulase enzymes are ancestral in Polyneoptera, Phylogenomics changes our understanding about earwig evolution, Fossil and phylogenetic analyses reveal recurrent periods of diversification and extinction in dictyopteran insects, Optimizing the widely used nuclear proteincoding gene primers in beetle phylogenies and their application in the genus

1456 p. Triplehorn, C. A., and N. F. Johnson (eds). > Panarthropoda > Tritocerebra > Phylum: can encounter the large Indian Cockroach Blatta orientalis. ootheca for the entire embryonic development. Evolution of egg incubation strategies in Blattodea resulted in ovipary, ovovivipary, and full vivipary [4,14]. Most are nocturnal and spend their days resting deep within tight-fitting cracks or crevices. Figure 1. In addition, some scientists suggest that German cockroach infestations may cause human psychological stress and that the stigma associated with infestations alters human behavior. Some previous studies gave purported records of fossil termite nests from 181 to 235 MYA, see [44]. Photograph by James Castner, University of Florida. Eggs. Rearing improvements for the German cockroach (Dictyoptera: Blattellidae). Canadian 1988. Cockroaches use chewing mouthparts to scavenge on organic materials. Helfer, J. R. 1987. How to know the grasshoppers, crickets, cockroaches, and their allies. The cockroach families: a cladistic appraisal of morpho-anatomical data. Our study corroborates the hypotheses of [9] that Blattellinae is polyphyletic with respect to Anallacta. Resolving the phylogeny of Blattodea has specific implications for understanding evolution in sociality, morphology, diet, endosymbiosis, or egg laying strategiesaspects that remain controversial or poorly understood in these insects. taking the lead. Mimosilpha They can also Enter your email address below and we will send you your username, If the address matches an existing account you will receive an email with instructions to retrieve your username, Institut de Systmatique, Evolution, Biodiversit (ISYEB), Musum national d'Histoire naturelle, CNRS, Sorbonne Universit, EPHE, 57 rue Cuvier, CP50, 75005 Paris, France, Department of Ecology and Evolutionary Biology, The University of Tennessee, Dabney Hall, 1416 Circle Dr., Knoxville, TN 37996, USA, Institut fr Spezielle Zoologie und Evolutionsbiologie mit Phyletischem Museum, Friedrich-Schiller Universitt Jena, Vor dem Neutor 1, 07743 Jena, Germany, Center for Taxonomy and Evolutionary Research, Zoological Research Museum Alexander Koenig (ZFMK), Adenauerallee 160, 53113 Bonn, Germany, CR2P (Centre de Recherche en Palontologie Paris), MNHN CNRS Sorbonne Universit, UPMC Univ Paris 06, MNHN, CNRS, Paris, France, Musum national d'Histoire naturelle, 57 rue Cuvier, CP38, 75005 Paris, France, Center for Molecular Biodiversity Research (ZMB), Zoological Research Museum Alexander Koenig (ZFMK), Adenauerallee 160, 53113 Bonn, Germany, Sugadaira Research Station, Mountain Science Center, University of Tsukuba, 1278-294 Sugadaira Kogen, Ueda, Nagano 386-2204, Japan, Federated Department of Biological Sciences, Rutgers, The State University of New Jersey and NJIT, 195 University Ave, Newark, NJ 07102, USA, BGI-Shenzhen, Shenzhen 518083, People's Republic of China, Steinmann-Institute, Institute for Paleontology, University of Bonn, Nussallee 8, 53115 Bonn, Germany, Animal Ecology and Conservation, Zoological Institute, University of Hamburg, Martin-Luther-King-Platz 3, 20146 Hamburg, Germany, Biosystematics Group, Wageningen University and Research, Droevendaalsesteeg 1, 6708 PB Wageningen, The Netherlands, Hessisches Landesmuseum Darmstadt, Friedensplatz 1, 64283 Darmstadt, Germany, Beijing Advanced Innovation Center for Food Nutrition and Human Health, China Agricultural University, Beijing 100193, People's Republic of China, Department of Entomology, China Agricultural University, Beijing 100193, People's Republic of China, Australian National Insect Collection, CSIRO National Research Collections Australia (NRCA), Acton, Canberra, Australian Capital Territory, Australia, Evolutionary Biology and Ecology, Institute for Biology I, University of Freiburg, Hauptstr. Specimens may need to be captured and cooled/stunned/killed before they can be photographed. 1985. D.B., A.D., B.M, L.P., M.P., R.S.P., K.M., and S.S. developed scripts, datasets, and programs. Dictyoptera. Hexapoda the ootheca by swallowing air and inflating themselves, in this way splitting The cockroach. The ootheca can be seen protruding from the posterior end (genital chamber) of the female. Cockroaches enter buildings to escape cold, hot, dry, or wet conditions, and infest even the most sanitary buildings. Figure 7.

The adult is 10 to 15 mm long, brown to dark brown in color with two distinct parallel bands running the length of the pronotum. It was introduced into the United States from Europe and is originally from Asia or Africa. the ootheca with debris so that it is difficult for it to be located by cockroaches. themselves from predators mainly by hiding away, but produce defensive Includes Aptera fusca Body length-width ratios have been greatly conserved within Blattodea (figure3d). Transcriptome sequences: Genbank Umbrella Bioproject ID PRJNA183205, see Supplementary Table S1 for single Bioprojects included in this study. Miscellaneous Collections 134 (10): 1-147 (reprinted 1967, Edwards

Photograph by James Castner, University of Florida. and We estimated divergence times with the software mcmctree as implemented in Paml v. 4.9 g [35]. Comparison of divergence times for selected Blattodea lineages retrieved in our study (black bars) with the fossil record (green bars), selected previous studies (blue bars: [9,11,12,16]) and important historical events as mentioned in the text. Entomology 15: 75-96. Superorder Dictyoptera, Roaches and Mantids, Habits and Traits of Centipedes, Class Chilopoda, The Insect Index - Sorted by Scientific Names, Insects: The Most Diverse Animal Group in the Planet. Abbreviations: Eukinola. = Eukinolabia and Xenono. = Xenonomia.Download figureOpen in new tabDownload PowerPoint. Within crown-Blattodea, our results more sharply contrast other divergence time studies [9,11,12,16] and show that most subgroups are much younger than previously thought. 304 MYA). The sister-group relationship between Cryptocercus and Isoptera is well supported (e.g. Members of the order Blattodea undergo incomplete or simple metamorphosis, with three stages of development: egg, nymph, and adult. Although many aspects of Blattodea biology such as morphology [2,6], palaeontology [7,8], biogeography [9], physiology [4,10], and other life-history aspects [4] have been studied, there still remain various open questions. Our divergence time estimates clearly show that modern cockroaches evolved after these roachoids with long ovipositors, thus further confirmation that these roachoids are not crown-Dictyoptera (figure2; further discussed in [7,8]). Within Blaberoidea, we retrieve a topology not previously recovered by any study, but all relationships are strongly supported by our data (figure1; electronic supplementary material, S1.4). Our ancestral state reconstruction suggests that laying an ootheca in a pre-dug hole is an ancestral trait in Blattodea (figure3b). The adult is 3.8 cm (1.5 in) long. We ensured all runs had an effective sample size of more than 200 and converged. Nymphs will often hatch from the ootheca while the female is still carrying it. Insecticide treatments are available in a wide variety of formulations including baits, sprays (emulsifiable concentrates, wettable powders, microencapsulated), dusts, and powders. It is dark brown to black in color. Journal of Medical Entomology 31: 704-710. Photograph by P.G. Appendix of The female is 1 inches long, and the male is 1 inch long. One aspect of maternal care in cockroaches relates to the egg package, i.e. harborage space and also negatively impact population size. Details of phylogenetic inference, bootstrap, bootstrap convergence, analyses for rogue taxa, and all topology tests are given in electronic supplementary material, S1.2 and S1.4. D.A.E. After depositing it, females of some species cover (dragonflies) and Ephemeroptera (Mayflies). We tabulated 20 morphological and behavioural characters from previous publications [2,11,14,37]. Our analyses clearly date the origin of Neoisoptera, which include the vast majority of termite species, to the beginning of the Palaeogene (53 13 MYA). Their life cycle may be completed in 2 months. Node dates (posterior mean) were inferred using the decisive aa dataset reduced to sites containing 95% or more data completeness, and nine exhaustively vetted fossil calibrations. Improving sanitation by eliminating food and water sources and clutter can have a significant impact on reducing the chances of infestation population size. Non toxic and low toxic alternatives for German cockroach control are available. Furthermore, the position of Mastotermes as sister to all remaining termites (figure1) is widely accepted and supported by many lines of evidence (e.g. Together, they form the clade Blattodea with approximately 7500 described species [3]. Zoological Catalogue of Australia 23: 21-162. Duchailluiinae, Pelmatosilpha, Angusticonicus s.l., other Polyzosteriinae) would be required to finally resolve this question. It was introduced into the United States from Europe and is originally from Asia or Africa. Smithsonian Misc. RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies, Pruning rogue taxa improves phylogenetic accuracy: an efficient algorithm and Webservice, An approximately unbiased test of phylogenetic tree selection, Likelihood-mapping: a simple method to visualize phylogenetic content of a sequence alignment, Phylogenomic analysis of Apoidea sheds new light on the sister group of bees, Best practices for justifying fossil calibrations, PAML 4: phylogenetic analysis by maximum likelihood, Phylogeny of cockroaches (Insecta, Dictyoptera, Blattodea), with placement of aberrant taxa and exploration of out-group sampling, SIMMAP: stochastic character mapping of discrete traits on phylogenies, The phylogeny of cockroach families: a cladistic appraisal of morpho-anatomical data, A contribution to the understanding of cockroach-termite affinities. The vast majority of cockroach species (more than 99% of Since the German cockroach is considered an aesthetic pest, the action threshold for this insect depends upon the tolerance of the people living in the infested dwelling. Immatures grow by molting and may require from 2 months to 3 years to fully mature, depending on species and environment. The Blattabacterium, that provide their hosts with amino acids and recycle nitrogenous waste products into usable nitrogen [10]. The order Blattodea includes the cockroaches, insects unfairly reviled the world over. Most Blattodea are omnivorous detritus feeders [4], a diet with unpredictable and variable nutrient quality. Vandenberg (Coleoptera: Coccinellidae), Complex Evolution of Insect Insulin Receptors and Homologous Decoy Receptors, and Functional Significance of Their Multiplicity, A new mid-Cretaceous cockroach of stem Nocticolidae and reestimating the age of Corydioidea (Dictyoptera: Blattodea), Fossil amber reveals springtails longstanding dispersal by social insects, The Generalized Feeding Apparatus of Cockroaches: A Model for Biting and Chewing Insects, Evolution of Termite Symbiosis Informed by Transcriptome-Based Phylogenies, Revision of the genus Attaphila (Blattodea: Blaberoidea), myrmecophiles living in the mushroom gardens of leaf-cutting ants, Digging deep: a revised phylogeny of Australian burrowing cockroaches (Blaberidae: Panesthiinae, Geoscapheinae) confirms extensive nonmonophyly and provides insights into biogeography and evolution of burrowing, No Evidence for Single-Copy Immune-Gene Specific Signals of Selection in Termites, Old World and New World Phasmatodea: Phylogenomics Resolve the Evolutionary History of Stick and Leaf Insects, The diversification of termites: Inferences from a complete specieslevel phylogeny, Origin of symbiotic gut spirochetes as key players in the nutrition of termites, Combining molecular datasets with strongly heterogeneous taxon coverage enlightens the peculiar biogeographic history of stoneflies (Insecta: Plecoptera), Exploring the relationship of