Ottmar Herchenröder
email: ottmar.herchenroeder@med.uni-rostock.de

Martin Löchelt
email: m.loechelt@dkfz.de

Florence Buseyne
email: florence.buseyne@pasteur.fr

Antoine Gessain
email: antoine.gessain@pasteur.fr

Marcelo A Soares
email: masoares@inca.gov.br

Arifa S Khan
email: arifa.khan@fda.hhs.gov

Dirk Lindemann
email: dirk.lindemann@tu-dresden.de

Twelfth International Foamy Virus Conference-Meeting Report

Keywords: foamy virus, spumaretrovirus, cross-species virus transmission, zoonosis, restriction factors, immune responses, FV vectors, virus replication, latent infection

The 12th

Introduction

The baroque city of Dresden, Germany and its modern biomedical research campus set the stage for the 12th International Foamy Virus Conference hosted and organized by Dirk Lindemann and his team. Based on a 24-year-old tradition since the first meeting, which was held in 1994 in London (Table 1), Dirk arranged an exciting scientific program on current aspects of foamy viruses (FVs), also known as spumaretroviruses. FVs are the only type of viruses in the subfamily of Spumaretrovirinae, while all other retroviruses belong to the Orthoretrovirinae. A biannual meeting brings together most of the FV researchers from different international institutions to present their progress and discuss new developments, and also provides a platform for collaborations. The presentations at the Dresden meeting reflected the dynamics of the field: New and evolving topics, "old" questions being addressed with new technologies, and identification of priority topics that need attention in the coming years. The scientific and generational changes in the field were noted by first-time participating scientists in the meeting. Meanwhile, some topics addressed at past gatherings have now apparently been solved, for instance, the acceptance of the uniqueness of the FV replication strategy, which has led to the establishment of Spumaretrovirinae as a separate subfamily of retroviruses, followed by an updated taxonomy and nomenclature [START_REF] Khan | Spumaretroviruses: Updated taxonomy and nomenclature[END_REF]. As another milestone, it had been the integrase protein of the prototype FV (PFV; designated as SFVpsc_huHSRV.13), which led to the first successful crystallization of a complete retroviral intasome complex and subsequent ultrastructural analyses of the active intermediates [START_REF] Hare | Retroviral intasome assembly and inhibition of DNA strand transfer[END_REF]. Thus, the FV integrase serves as the template structure for all retroviral/retroid element integrases. Due to their unique molecular biology, FVs may be considered relevant models to study yet unidentified principles of (retro-)virology. Ultimately, the taxonomic "upgrade" could draw other virologists' attention to these viruses and eventually get more scientists involved in FV research.

In addition to presentations on many aspects of basic, applied, and translational biology of a constantly growing number of new and molecularly characterized FV isolates from diverse hosts, there were keynote presentations from related fields as well as opportunities that were provided for informal discussions and scientific exchange.

Summary of Scientific Sessions

The session chairs in consultation with the speakers developed the summaries below, which provide an overview of the current status and future directions for advancing the topic.

Epidemiology of Natural and Zoonotic Infections (Session Chair: Ottmar Herchenröder)

FVs are infectious agents that persistently infect primate, feline, bovine, and equine species as well as chiropterans (bats). Generally, FVs co-evolve with their host species. Having been an issue of debate in the early decades of FV research, we nowadays know that humans are not natural hosts of FVs. However, transmissions of simian FVs (SFV) to man are not uncommon in natural habitats shared by humans and non-human primates (NHPs), or in settings where the latter are held in captivity, such as in zoos or primate research centers. In contrast to the most prominent example of interspecies transmissions of retroviruses to man that initiated the worldwide AIDS epidemic caused by human immunodeficiency viruses (HIV), FVs are considered apathogenic in both their natural hosts and after interspecies transmissions.

In this session, researchers from the U.S., Brazil, and France gave further insights on the epidemiology of FVs and transmissions between species. Sue VandeWoude (Colorado State University, Fort Collins, CO, USA) reported on her group's comparative epidemiologic surveys amongst mountain lions and domestic cats that share, in part, habitats in the Rocky Mountains foothills. Feline FVs (FFVs) are highly prevalent in these wild animals and, interestingly, interspecies transmissions from domestic cats to mountain lions were frequent as documented by sequence comparisons. Vice versa, FV from mountain lion (Puma concolor), was not seen in domestic cats, presumably, as the interaction between individuals of both species necessary for virus transmission may usually not allow the pet to walk away unscathed [START_REF] Kehl | Non-simian foamy viruses: Molecular virology, tropism and prevalence and zoonotic/interspecies transmission[END_REF]. Marcelo Soares' group (Universidade Federal do Rio de Janeiro, Rio de Janeiro, Brazil) had previously isolated and characterized FVs from a series of New World NHPs in Brazil. In Dresden, Marcelo's co-worker, Cláudia Muniz, added new data on interspecies transmissions of SFVs from the "invading species", Golden-headed lion tamarin (Leontopithecus chrysomelas), to the endangered local population of Golden lion tamarins (Leontopithecus rosalia) in the greater Rio de Janeiro area. This study also underlined the validity and robustness of oral swabs as a non-invasive method to achieve both species assignment and FV isolation and characterization [START_REF] Muniz | A non-invasive specimen collection method and a novel simian foamy virus (SFV) DNA quantification assay in New World primates reveal aspects of tissue tropism and improved SFV detection[END_REF]. Antoine Gessain (Institut Pasteur, Paris, France) presented a case-control study executed by French and Cameroonian scientists amongst hunters in Cameroon's south-eastern rain forests that gave pause for thought whether simian FV-infections after interspecies transmissions always remain benign over the lifetime of the human recipient. The cases presented with deviations in several blood parameters in comparison to controls matching in residence, age, and social status [START_REF] Buseyne | Clinical Signs and Blood Test Results Among Humans Infected With Zoonotic Simian Foamy Virus: A Case-Control Study[END_REF]. Since clinical assessments did not differ between the cases and the controls, the data are reassuring for infected people. However, this study and most previous reports suffer from a strong bias as only apparently healthy individuals were studied. The audience's attention was brought back to domestic animals by Magdalena Materniak-Kornas (National Veterinary Research Institute, Puławy, Poland), who analyzed co-infections with gammaherpesviruses and bovine FV (BFV) in cows suffering from post-partum metritis. By multiplex qPCR and Elisa techniques, the researchers found a good number of animals coinfected with both virus species [START_REF] Materniak-Kornas | Development of a Recombinant Protein-based ELISA for Detection of Antibodies Against Bovine Foamy Virus[END_REF]. Whether FVs have any influence on the clinical onset and burden of metritis in cattle remains speculative.

Altogether, some general beliefs hold true after this latest gathering of FV researchers. First, FV infections are common in numerous species, including livestock and many of those we humans conceive affection for, such as apes, monkeys, and of course cats. Second, FV infections remain in both the natural hosts or the foreign recipients after interspecies transmissions, apparently apathogenic as consented in the aftermath of the first FV conference (Table 1) [START_REF] Weiss | Reverse transcription. Foamy viruses bubble on [news[END_REF]. More recent data, however, imply an association with hematologic abnormalities [START_REF] Buseyne | Clinical Signs and Blood Test Results Among Humans Infected With Zoonotic Simian Foamy Virus: A Case-Control Study[END_REF] and demand further research. Third, unlike many other retroviruses, FV distribution is a worldwide affair and those viruses appear phylogenetically more ancient than their pathogenic counterparts. As such, FVs may be considered as models to study in depth retrovirus biology and evolution.

The author of this section had a long time ago propagated FV isolates in Cf2Th, a cell line derived from canine thymus, which eventually resulted in the first complete sequence of a chimpanzee FV [START_REF] Herchenröder | Isolation, cloning, and sequencing of simian foamy viruses from chimpanzees (SFVcpz): High homology to human foamy virus (HFV)[END_REF]. Cf2Th cells are very permissive for several FVs, although all canine species are considered free from non-endogenous retroviruses. With the surprises FV have entertained virologists over the last decades, he wonders, whether some day one will isolate a FV from man's best companion, the dog. Similarly, it may be worth to look again whether any rodents harbor FVs.

Interactions of Foamy Viruses with the Immune System (Session Chairs: Antoine Gessain, Marcelo A. Soares)

Immunological aspects of infection by FV are still poorly characterized, both in naturally infected animals and in human infections after cross-species transmission. This scarcity of data includes both innate and acquired immunity, and several issues are still to be addressed in these areas.

What do we know about the immune responses directed against FVs? Several findings suggest strongly an efficient control of SFVs by the immune system in NHPs. These include the apparent lack of pathogenicity of SFVs in vivo, restriction of SFV RNA expression mainly to the oral cavity, and seemingly the gastrointestinal tract in some NHP species, as well as the very limited genetic variability in vivo. Furthermore, recent data indicate that FVs are susceptible to several restriction factors (see virus-host interactions Section 2.4), demonstrating the role of innate immunity in SFV infection, as well as, in some instances, in humans infected by SFVs of zoonotic origin [START_REF] Pinto-Santini | Foamy virus zoonotic infections[END_REF][START_REF] Rua | evolution and innate immune control of simian foamy viruses in humans[END_REF]. In contrast, the control of SFV replication by innate immunity (antiviral molecules and cells), as well as adaptive immune responses (virus-specific antibodies and T lymphocytes), remain poorly characterized in NHP and even less in humans infected by an SFV.

The following questions remain poorly answered: (i) Can an immunodepression or a viral co-infection alter the pathogenicity of FVs in animal models and in natural infections? (ii) What are the molecular mechanisms by which FVs are efficiently sensed by innate immune cells? This will allow us to better understand how SFVs trigger an innate immune response, especially interferon (IFN) production, which has been demonstrated in blood plasmacytoid dendritic cells, a quite relevant model [START_REF] Rua | Innate sensing of foamy viruses by human hematopoietic cells[END_REF]. (iii) What is the respective part of each arm of the immune system in the modulation of FV infection and the apparent lack of pathogenicity of these viruses, especially in the context of zoonotic infection? (iv) What is the efficacy of antiviral activity of antibodies, especially the role and importance of neutralizing antibodies in vivo?

Answers to some of these questions were given at this meeting. Concerning immunosuppression and FV infection, a previous study has demonstrated that SIV coinfection in a macaque model led to expanded tissue tropism of SFV [START_REF] Murray | Expanded tissue targets for foamy virus replication with simian immunodeficiency virus-induced immunosuppression[END_REF]. Conversely, SFV infection accelerated clinical progression, immunodeficiency, and viral load after SIV infection [START_REF] Choudhary | Influence of naturally occurring simian foamy viruses (SFVs) on SIV disease progression in the rhesus macaque (Macaca mulatta) model[END_REF]. Thus, Cláudia Muniz and colleagues (Universidade Federal do Rio de Janeiro, Rio de Janeiro, Brazil) performed experiments in order to study the kinetics of SFV viral loads during immunosuppressive therapy, i.e., with tacrolimus, a drug commonly used to prevent graft rejection after transplantation. SFV DNA load was stable without major variation during the immunosuppression period in the two studied groups. The authors concluded that the immune mechanisms affected by tacrolimus, mostly T lymphocytes, were unable to modulate SFV replication at least during the chronic phase of the infection. In a South American NHP (Brachyteles arachnoides) with symptoms indicative of clinical immunodepression, likely due to infection by a simian type-D retrovirus (SRV), the same team demonstrated the presence of very high SFV DNA levels in the saliva and identified a new isolate, SFVbar [START_REF] Muniz | First Complete Genome Sequence of a Simian Foamy Virus Infecting the Neotropical Primate Brachyteles arachnoides[END_REF]. Furthermore, transcriptome analysis from blood yielded SRV-related transcripts, but failed to identify SFVbar RNA, suggesting that SFVbar remained latent in PBMCs despite the immunocompromised status of the host.

Concerning antibodies, Florence Buseyne and colleagues (Institut Pasteur, Paris, France) demonstrated the presence of potent neutralizing antibodies in humans infected by zoonotic gorilla and chimpanzee FVs. Furthermore, Florence showed that the antibodies are directed against conserved epitopes located in the dimorphic domain of the surface envelope protein. The neutralization patterns provided evidence for persistent expression of viral proteins and a high prevalence of viral co-infection by two different viruses diverging in the env SU gene [START_REF] Lambert | Potent neutralizing antibodies in humans infected with zoonotic simian foamy viruses target conserved epitopes located in the dimorphic domain of the surface envelope protein[END_REF].

Mathilde Couteaudier and colleagues (Institut Pasteur, Paris, France) investigated whether plasma antibodies can inhibit SFV cell-to-cell transmission. Indeed, SFV transmission is considered highly cell-to-cell associated. Interestingly, she demonstrated that plasma samples from humans infected by zoonotic SFV, selected for their potent ability to neutralize cell-free virus, were unable to inhibit virus cell-to-cell spread.

In a second session devoted to the interaction between FV and the immune system, Florence Buseyne and colleagues (Institut Pasteur, Paris, France) characterized an immunodominant epitope located in the leader region of the SFV envelope protein (SFVpsc_huHSRV.13, aa 96-110) and recognized by the plasma antibodies of most SFV-infected hunters from Central Africa, similar to what has been recently described in FFV infections [START_REF] Mühle | Epitope Mapping of the Antibody Response Against the Envelope Proteins of the Feline Foamy Virus[END_REF]. Whereas plasma from subjects infected with SFV derived from gorillas strongly recognized peptides corresponding to that envelope region of SFV from apes and other Old World primates, recognition was poor or absent to the respective peptides from SFV infecting more distant New World primates or from a specific African green monkey strain, suggesting evolutionary constraints in the adaptive immune responses observed. Whether the magnitude of such antibody responses can serve as proxies of overall adaptive immunity against SFV still remains to be determined.

By using FFV, Cláudia Muniz and coworkers (Universidade Federal do Rio de Janeiro, Rio de Janeiro, Brazil) had determined the effect of coinfections between FV and other retroviruses, in this case, the feline leukemia virus (FeLV) in domestic cats. Since FeLV causes immunodeficiency in their natural hosts, the feline model can also be used to understand the kinetics of FFV infection in the context of immunosuppresion. The investigators found that among FeLV/FFV-coinfected cats, those displaying an FeLV regressive infection (that tends to spontaneous clearance of the virus in the infected host), FFV replication and potential transmissibility is also reduced in the oral cavity, suggesting a synergistic interaction of both retroviruses as recently confirmed by Powers et al. [START_REF] Powers | Feline Leukemia Virus (FeLV) Disease Outcomes in a Domestic Cat Breeding Colony: Relationship to Endogenous FeLV and Other Chronic Viral Infections[END_REF]. On the other hand, FFV proviral loads in the peripheral blood did not differ between cats with or without clinical signs after FeLV infection, again highlighting a lack of association between FV replication and the general immunosuppressive status of the host, as pointed out in the studies conducted in NHP described above.

The last two presentations of the session addressed the role of innate immunity on FV infection in a very complementary manner. Jakob Weber and coworkers (Technische Universität Dresden, Dresden, Germany) showed the utility of the Phorbol 12-myristate 13-acetate (PMA)-differentiated monocytic cell line, THP-1, as a model system to study innate sensing in macrophages and monocytes, the likely first immune cell types to contact FV particles in infected tissues. The researchers found that incubation of these cells with PFV resulted in the induction of interferon-stimulated genes (ISGs). Furthermore, they demonstrated that PFV sensing is independent of Tas expression and integration of the proviral genome, but rather occurs at cell entry and requires access to the cytoplasm of the infected cell, with reverse-transcribed DNA being a likely recognized molecule. Maïwenn Bergez and colleagues (Paul-Ehrlich-Institut, Langen, Germany) showed that transduction of PFV vector particles into both THP-1 cells and primary human monocyte-derived dendritic cells induces the expression of the interferon regulatory factor 3 (IRF3)-dependent ISGs. Such induction was highly attenuated when RT-deficient viruses were used or azidothymidine was added during transduction, confirming that the newly reverse-transcribed PFV DNA acts as a pathogen-associated molecular pattern that stimulates the host cell innate immune system. The cytosolic sensors involved in the innate immune response to FV infection are yet to be determined.

Molecular and Cellular Biology of Foamy Viruses (Session Chair: Arifa S. Khan)

The development of advanced molecular tools and new generation biochemical reagents have enabled us to re-visit important questions regarding the biology and replication of FVs. Our current knowledge regarding the FV life cycle is mainly based on studies with SFVpsc_huHSRV.13, since this was initially believed to be a human FV (until it was sequenced and identified as a chimpanzee FV, which had infected a human by cross-species transmission). The information has been expanded based on FV isolates from other NHP species and extended by studies on FFVs and BFVs. The majority of the early work is based on laboratory virus strains. Although a few efforts are currently directed at studying the biology of natural FV isolates, most are from NHPs and only recently have also included New World monkeys. This research gap is further being addressed by studies investigating natural infections of FVs in bovine and feline species. Moreover, the availability of new and more sensitive assays that can investigate the different steps of FV replication, from infection/entry through integration/expression, resulting in release/exit, are important for understanding natural virus transmission and cross-species infections in humans by NHP FVs. To aid these goals, there is a need to develop reagents and functional assays for studying the molecular and cellular biology of naturally-occurring FV isolates. This session highlighted applications of new molecular tools and assays for investigating critical steps in spumaretrovirus replication.

Stefanie Richter from the Lindemann laboratory (Technische Universität Dresden, Dresden, Germany) reported on the adaptation of the fluorescent-based β-lactamase (BlaM) fusion assay [START_REF] Jones | The beta-Lactamase Assay: Harnessing a FRET Biosensor to Analyse Viral Fusion Mechanisms[END_REF] for FVs to obtain new insights about the kinetics and temperature requirements of FV Env-mediated fusion. The study used BlaM-containing SFVpsc_huHSRV.13 single-round vector particles habouring either SFVpsc_huHSRV.13 or SFVmcy_FV21 Env proteins and the Pac2 zebrafish embryonic cell line, which was previously thought to be resistant to FV Env-mediated entry [START_REF] Stirnnagel | Analysis of prototype foamy virus particle-host cell interaction with autofluorescent retroviral particles[END_REF]. Using Pac2 cells loaded with the fluorescent BlaM substrate CCF4-AM, the researchers demonstrated that this cell line was permissive for FV Env-mediated fusion. These results highlighted the potential of this approach for providing insight into the differences seen in the susceptibility of cells to FV infection.

Ivo Glück from the Lamb laboratory (Ludwig-Maximilians-Universität, München, Germany) presented on the application of the TrIC method [START_REF] Dupont | Tracking image correlation: Combining single-particle tracking and image correlation[END_REF] to track steps involved in the fusion process of single FV particles in live HeLa cells. Virus particles used in the study contained the viral envelope protein (Env) tagged with mCherry and the capsid protein (Gag) tagged with eGFP. By tracking the eGFP signal and locally cross-correlating the eGFP and mCherry intensities, the individual viral fusion events could be visualized as the capsid separated from the envelope. The analysis revealed a previously undetected intermediate step in the FV fusion process, in which the capsid and envelope signals are separated by approximately 300 nm, but remain tethered for an average of 7.1 min before full separation. An important next step is to identify the linking component that tethers the viral capsid to the envelope.

Martin Löchelt (German Cancer Research Center, Heidelberg, Germany) extended the previous identification of the chromatin binding site (CBS) in PFV Gag by functionally characterizing a highly conserved motif of the CBS of the Gag protein of FFV. It was demonstrated that the RYG residues in the QPQRYG motif are, in addition, essential during the early stages of infection after entry into the cytosol since mutagenesis of RYG abrogated the accumulation of Gag and proviral DNA in the nucleus and, subsequently, DNA integration into the host genome, similar to mutations within the chromatin binding. The results confirmed that chromatin binding by foamy virus Gag is a shared feature among FVs belonging to diverse genera and mediated by a conserved protein region located at the C-terminus of Gag, which is of prime importance for the provirus integration site. This motif likely influences the incoming virus capsid or its disassembly intermediates, but not newly synthesized Gag or its assembly products. The study, which was recently published [START_REF] Wei | The chromatin binding domain, including the QPQRYG motif, of feline foamy virus Gag is required for viral DNA integration and nuclear accumulation of Gag and the viral genome[END_REF] provides a strategy for comparing different FVs for mechanism and host cell conditions that can influence the efficiency of integration.

FV research using new assays and methodologies can be further enhanced and extended by the development of advanced technologies, such as next generation or high-throughput sequencing (NGS, HTS), which has emerged as a powerful tool that can help to investigate various aspects of virus biology and replication. In particular, transcriptomics and genomics studies using NGS could help determine the virus-host interactions that result in the seemingly general FV latency and limited expression seen in natural infections. The results from such studies may aid in predicting potential outcomes of FV infections in humans.

Mechanisms of Virus-Host Interactions: Cellular Antiviral Factors and Viral Antagonists (Session Chair: Florence Buseyne)

The interactions of FVs with their hosts were shaped by millions of years of coevolution [START_REF] Rethwilm | Evolution of foamy viruses: The most ancient of all retroviruses[END_REF]. FV have a ubiquitous tropism in vitro and numerous cross-species transmission events occurred in their evolutionary history. Thus, these viruses escape most constitutively expressed restriction factors. On the other hand, they are sensitive to the action of type I and type II IFN in vitro and are apparently apathogenic in vivo. Among the ISG products, TRIM5α, N-myc-interactor, IFP35, TRIM19, APOBEC, and tetherin were described to inhibit FV, while MxB and SAMHD1 had no action, and some discordant results are thus far unresolved. The currently open questions in the field, summarized after the former FV conference in 2016 [START_REF] Buseyne | Eleventh International Foamy Virus Conference-Meeting Report[END_REF], were about the role of restriction factors and IFN-induced genes in controlling FV gene expression in vivo in their natural hosts and in humans after cross-species transmission. Lack of knowledge on the induction and action of non-IFN antiviral actors (inflammasome, cytokines, and miRNA) were striking.

This conference provided new insights on four topics. Carsten Münk (Heinrich-Heine-Universität Düsseldorf, Düsseldorf, Germany) presented on APOBEC3s, which are potent inhibitors of FV. The viral Bet protein is the single antiviral factor antagonist described for FV. Its mechanism of action is unique: Bet prevents APOBEC3s incorporation into viral particles and thereby, provides protection of FVs, as well as lentiviruses, if Bet is expressed in trans, against APOBEC3s' antiviral activity. Conversely, Carmen Ledesma-Feliciano reported in the FV vector session (see 2.6) that the feline immunodeficiency virus (FIV) Vif protein can replace the FFV Bet protein in vitro, but the chimeric virus was attenuated in vivo [START_REF] Ledesma-Feliciano | Replacement of feline foamy virus bet by feline immunodeficiency virus vif yields replicative virus with novel vaccine candidate potential[END_REF]. These in vivo attenuated phenotypes of Bet-deficient FV highlighted that additional Bet functions impacting FVs interaction with their hosts are waiting to be described.

Wenhu Cao from the Löchelt laboratory (German Cancer Research Center, Heidelberg, Germany) presented new results on bovine FV (BFV) miRNAs. BFV encodes miRNAs with a noncanonical biogenesis involving RNA polymerase III, generation of a single pri-miRNA, and its processing into three functional miRNAs. Cellular proteins were identified whose expression were suppressed by BFV miRNA and this was shown to result in repression of IFN-β and NF-κB pathways. Importantly, deletion of miRNA led to in vivo attenuation of BFV, confirming the major role of miRNAs in virus-host interactions.

Melissa Kane from the Bieniasz laboratory (The Rockefeller University, New York, NY, USA) presented new data based on results from the screening of nearly 500 human and macaque antiviral genes against 11 retroviruses, including the PFV [START_REF] Kane | Identification of Interferon-Stimulated Genes with Antiretroviral Activity[END_REF]. The screen identified the ISG PHD finger protein 11 (PHF11) as a PFV-specific inhibitor. In vitro, PFV was unable to escape PHF11 restriction. PHF11 is involved in DNA damage signaling and DNA repair, and inhibits an early step in FV replication.

Arifa Khan (U.S. Food and Drug Administration, Silver Spring, MD, USA) presented the careful characterization of SFV-infected human A549 cell clones with various viral expression patterns, defined by viral DNA and RNA expression and production of infectious virions. Clones with fully latent and actively replicating SFV were analyzed for gene expression using next generation sequencing. Cells infected with different macaque SFV strains with widely different kinetics were included in the screen. Data overview showed diversity across strains in their ability to modulate IFN-signaling and other immune activation pathways.

Important questions and future directions were highlighted at the meeting. Data pointed to the uniqueness of FV (Bet, miRNA, and the PHF11) with elegant studies starting with molecular understanding of the activity of an FV component to in vivo assessment of its action. Such specificities open new molecular medicine applications, such as expression of heterologous regulatory RNA by BFV miRNA. The interest of high-throughput assays associated with well-validated in vitro models was evident and indeed, the PHF11 will not be identified by a hypothesis-driven strategy as early as with the comprehensive approach used [START_REF] Kane | Identification of Interferon-Stimulated Genes with Antiretroviral Activity[END_REF]. In the future, new input is expected from modern biology approaches to identify cellular factors regulating viral replication and latency.

Structural Studies of Foamy Virus Proteins (Session Chair: Martin Löchelt)

Structural biology provides challenging insights into the molecular and atomic organization and underlying functions of proteins and molecular assemblies of high complexity. In addition, structural biology is a major tool in drug discovery and optimization and has enormous implications for translational research, medical care, and disease prevention. Under these perspectives, the current state-of-the-art of FV structural biology, focused in particular on integrase (IN) and the Gag structural proteins, was summarized in excellent reviews by Paul Lesbats (CNRS UMR5234, Bordeaux, France) and Jonathan Stoye (The Francis Crick Institute, London, United Kingdom).

FV Pol proteins share most of the conserved sequence motifs with the other retroviruses. However, their expression mode via a spliced transcript, resulting in the lack of a Gag-Pol fusion protein, most probably determines the uniqueness of FV particle assembly, genome incorporation, protein processing, capsid maturation, and reverse transcription. Another unique feature is the lack of processing of FV Pol precursor molecules to release the protease [START_REF] Kehl | Non-simian foamy viruses: Molecular virology, tropism and prevalence and zoonotic/interspecies transmission[END_REF][START_REF] Rethwilm | Evolution of foamy viruses: The most ancient of all retroviruses[END_REF][START_REF] Lindemann | Foamy virus biology and its application for vector development[END_REF].

After a lot of unsuccessful attempts using other retroviruses, PFV IN was the first retroviral IN to be crystallized in its active form and as a high-molecular mass complex together with its target DNA [START_REF] Hare | Retroviral intasome assembly and inhibition of DNA strand transfer[END_REF]. Based on this structure and subsequent new data, highly divergent degrees of IN oligomerization in intasomes has been described for different retroviruses [START_REF] Engelman | Retroviral intasomes arising[END_REF]. Here, PFV IN tetramers contrast other retroviral octo-and hexadecameric arrangements that make up the active intasomes. It is possible that the comparably low degree of PFV IN oligomerisation favored or even allowed its crystallization and thus, its structural analysis. Based on his experience in retroviral IN interactions with chromatin, Paul Lesbats described the interaction of the PFV chromatin-binding site with histones confirming and extending knowledge on C-terminal Gag residues involved in this process that contributes to FV integration site selection [START_REF] Lesbats | Structural basis for spumavirus GAG tethering to chromatin[END_REF].

FV Gag and that of other retroviruses largely perform equivalent functions. However, several features, including the absence of genome-binding Cys-His motifs and a major homology region, the limited Gag processing at a very C-terminal site, and the essential requirement of N-terminal Gag and Env LP domains for particle release, are fundamentally different from that of other retroviruses [START_REF] Müllers | The foamy virus Gag proteins: What makes them different?[END_REF]. The N-terminal and central domains of the unique FV Gag proteins are the focus of Jonathan Stoye's past and current activities. Here, structural studies strongly target evolutionary issues related to retroviral capsid structures and their interaction with host-encoded restriction factors. Jonathan described structural analyses that identified two independent domains in a central part of FV Gag, which most probably derived from sequence duplications [START_REF] Ball | Structure of a Spumaretrovirus Gag Central Domain Reveals an Ancient Retroviral Capsid[END_REF]. Surprisingly, these structures reveal a greater and unforeseen similarity to the Ty3 retrotransposon-derived mammalian protein ARC, than to conventional retroviral capsid proteins [START_REF] Taylor | A comparative analysis of the foamy and ortho virus capsid structures reveals an ancient domain duplication[END_REF].

In summary, structural studies of FV Pol proteins have shown that they are highly related to those of other retroviruses and in the case of IN, even serve as model structures for the whole virus family. By contrast, although the amino-terminal and central domains of FV-Gag have been structurally characterized, the evolutionary origin and relationship with orthoretroviral Gag and exapted proteins found in mammalian genomes is the subject of current FV structural studies. Other FV proteins, most of them with unique features, have at large not been targets of high-resolution studies. Structural studies of FV Env are limited to a low-resolution (9 Å). Cryo-EM structure revealed the trimeric nature of the PFV Env gp80 SU spike [START_REF] Effantin | Cryo-electron Microscopy Structure of the Native Prototype Foamy Virus Glycoprotein and Virus Architecture[END_REF] and the crystal structure an Env gp18 LP peptide bound to the N-terminal domain of PFV Gag [START_REF] Goldstone | A Unique Spumavirus Gag N-terminal Domain with Functional Properties of Orthoretroviral Matrix and Capsid[END_REF]. Further high-resolution structural analyses of Env from two distinct serotypes of primate and feline FV SU could greatly foster the understanding of virus neutralization and the overlap with receptor binding, and may even support identification of the FV receptor(s). Finally, structural data on the non-canonical FV Tas transcriptional transactivator and the enigmatic Bet protein that counteracts APOBEC3-mediated restriction, but may also fulfil many additional functions, would fertilize further research on these essential and FV-specific proteins [START_REF] Lukic | Identification of the feline foamy virus Bet domain essential for APOBEC3 counteraction[END_REF].

2.6. Development and Application of Foamy Virus Vector Systems (Session Chair: Dirk Lindemann)

Over the last 50 years human gene therapy has developed from fiction to become reality. After approval of the first gene therapy drugs several years ago, the last 24 months have seen a good number of new gene therapy drugs being approved and entering the market. The majority of them are based on viral gene ferries, which is also true for drugs currently being examined in gene therapy trials worldwide. This most likely reflects the fact that viruses may be considered as evolutionarily optimized nucleic acid delivery entities, although it took researchers decades to develop viral vector systems with optimized features of transduction efficiency, balanced expression control, and safety. Retroviral vectors are still the most frequent gene ferries used in gene therapy clinical trials and approved drugs. Gammaretroviral vectors based on murine leukemia virus were the first gene ferries being used in clinical trials. They were also employed in the SCID-X1 clinical trials around the turn of the century and were the first gene transfer tools with unequivocal therapeutic benefits for the patients. Lentiviral vectors based on HIV-1 were developed only later. Due to their ability to also efficiently transduce non-mitotic cells they have become more popular, which can be seen by the rising number of clinical trials using them as gene delivery tools.

The first vector systems based on FV were developed about 20 years ago [START_REF] Lindemann | Foamy virus biology and its application for vector development[END_REF][START_REF] Trobridge | Large animal models for foamy virus vector gene therapy[END_REF]. FV have several natural features of a good candidate gene transfer vector, such as one of the largest retroviral genomes, an infectious DNA genome, a favorable integration site profile, and an extremely broad tropism. Most important is their apparent apathogenicity in both natural and zoonotic infections that renders them attractive for gene delivery. Furthermore, they have shown promising therapeutic results in preclinical studies using animal model systems [START_REF] Lindemann | Foamy virus biology and its application for vector development[END_REF][START_REF] Trobridge | Large animal models for foamy virus vector gene therapy[END_REF]. However, so far, FV vectors or individual components of FV, such as the FV Env protein, have not been used in human gene therapy clinical trials.

Karol Budzik from the Russell laboratory (Mayo Clinic, Rochester, NY, USA) summarized his efforts to develop an FV platform for oncolytic virotherapy. SFVpve virus produced from a chimeric infectious molecular clone, which was derived from two chimpanzee FV strains (PAN1, PAN2), showed higher oncolytic activity in vitro than either parental strain or the PFV isolate in a variety of tumor cell lines. Furthermore, a U251 glioblastoma-derived reporter cell line was established and used to demonstrate in vivo replication of FV in a mouse model. This provides the basis for further examination of the tumoricidal potential of this new chimeric SFV strain or armed versions thereof in animal model systems.

Carmen Ledesma-Feliciano from the VandeWoude laboratory (Colorado State University, Fort Collins, CO, USA) presented a recently published study on the experimental infection in domestic cats with a replication-competent wild type and a novel FFV vaccine vector candidate expressing a truncated FFV-Bet/FIV-Vif fusion protein [START_REF] Ledesma-Feliciano | Replacement of feline foamy virus bet by feline immunodeficiency virus vif yields replicative virus with novel vaccine candidate potential[END_REF]. In vitro analysis in feline cells showed a requirement of FIV Vif expression without an FFV Bet N-terminal tag for efficient viral replication. In contrast, inoculation of immunocompetent domestic cats revealed a poor replication capacity of the vaccine vector in comparison to wild-type FFV. This suggests a yet uncharacterized role of FFV Bet for in vivo replication besides its anti-APOBEC activity.

Fabian Lindel from the Lindemann laboratory (Technische Universität Dresden, Dresden, Germany) reported on a novel PFV based vector system for largely transient expression of CRISPR/Cas9 genome editing tools in various target tissues. The system exploits the previously described natural feature of FV to encapsidate and efficiently transmit non-viral RNAs, which was exploited to express Cas9 fully transient in transduced cells. When combined with integration-deficient retroviral vectors harboring a U6 promoter-driven sgRNA, efficient gene inactivation was achieved in different target cell types. Additionally, the inclusion of a donor DNA template enabled efficient gene editing of reporter or cellular genes by homology directed repair mechanisms.

Finally, Jennifer Donau from the Valtink laboratory (Technische Universität Dresden, Dresden, Germany) summarized her work towards the establishment of a therapeutic proliferation stimulation strategy for primary human ocular tissues. She reported on the identification of viral and cellular proliferation promoting factors for immortalization of primary human corneal endothelial (CEC) or retinal pigment epithelial (RPE) cells upon stable transduction by lentiviral vectors pseudotyped by FV glycoproteins. Furthermore, she demonstrated that integration-deficient lentiviral vectors are not suited for transient growth stimulation in a therapeutic setting due to their residual non-viral mediated integration potential, which results in stable immortalization of primary tissues. At the end, she referred to her first attempts to achieve growth stimulation in these primary target tissues by transient expression of proliferation promoting factors using FV-mediated non-viral RNA gene transfer.

Keynote Lectures

The first keynote lecture was given by Welkin E. Johnson (Boston College, Chestnut Hill, MA, USA). Welkin presented an overview of a comparative study of ancient env genes of endogenous retroviral elements that provides insight into the co-evolution of retroviruses and their vertebrate hosts. He reviewed examples of potential cases of exaptations, whereby former endogenous retrovirus (ERV) env genes are preserved by purifying evolutionary selection and now provide cellular functions, for example, in the development (syncytins-mediated trophoblast fusion) or antiviral defense against extant viruses (ERV-mediated superinfection resistance). Furthermore, he summarized his and other people's work on the origins and exaptive evolution of the ERV-Fc locus in mammals.

Frank Buchholz (Technische Universität Dresden, Dresden, Germany) delivered the second keynote lecture reviewing his and other researchers' work on programmable nucleases and designer recombinases for genome surgery. He presented examples from his lab using CRISPR/Cas9 library screens to dissect driver mutations from passenger mutations in human cancer cell lines as an approach towards a personalized cancer therapy. Furthermore, he summarized the pioneering work of his and Joachim Hauber's laboratory (Heinrich-Pette-Institut, Hamburg, Germany) on the broad-range anti-HIV-1 recombinase Brec1 and presented the latest results from experiments using humanized mouse models.

Conclusions

Two major goals were achieved by the 12th International Foamy Virus Meeting. First, bringing together most of the senior and junior scientists in the field with expertise in different disciplines for a scientific exchange and providing the opportunity for discussing ongoing, and initiating new collaborations. Second, attracting new people to the field, which was reflected by several first-time attendees, whose presentations strongly underlined the interdisciplinary character of the meeting and demonstrated the continued interest in this unique type of retrovirus. We enjoyed lively discussions after the individual presentations and in the breaks. We hope that the session summaries and highlights provided in this report illustrate that there is so much to be discovered and learned from FVs and will encourage interested researchers to join us at the 13th International Spumaretrovirus Conference, which is being planned for 2020 in Rio de Janeiro, Brazil. More information can be obtained by the conference host: Marcelo Soares masoares@biologia.ufrj.br.

Table 1 .

 1 Foamy virus conference history.

	Year	Location		Key Findings Reported
			•	Identification of "human foamy virus (HFV)" being the result of a zoonotic transmission of a chimpanzee FV
	1994	London, UK	•	Involvement of a defective HFV genome in viral latency
			•	Details of Tas-dependent HFV transcriptional control involving internal and LTR promoter
	1997	Herzogenhorn, Germany	• • •	Full-length sequence of a feline FV isolate Reverse transcription during virus morphogenesis results in infectious FVs with a DNA genome First generation replication-deficient PFV vectors
	1999	Gif-sur-Yvette, France	• • •	Characterization of an equine FV isolate FV glycoprotein-mediated subviral particle formation FV glycoprotein leader peptide-dependent viral particle release mechanism
	2002	Atlanta, GA, USA	• •	Application of 2nd generation replication-deficient PFV vector for hematopoietic stem cell gene transfer Viral genome-dependent FV Pol encapsidation mechanism
	2004	Würzburg, Germany	• • •	Identification of FV Bet as inhibitor of the APOBEC restriction factors ESCRT-complex-dependent FV release First generation replication-deficient feline FV vector systems
	2006	Seattle, WA, USA	• • •	Successful treatment of canine leukocyte adhesion deficiency by FV vector-mediated HSC gene transfer Centrosomal latency of incoming FV capsids in resting cells Identification of in vivo sites of FV replication in infected primates
	2008	Heidelberg, Germany	• •	CRM1-mediated nuclear export of unspliced FV RNAs PFV protease structure
	2010	Argos, Greece	• •	Host cells DDX6 protein involvement in PFV genome encapsidation Heparan sulfate as FV Env-dependent attachment factor
	2012	Bethesda, MD, USA	• • •	Structure of the PFV Gag N-terminus in complex with FV Env LP peptides Innate sensing of FV by plasmacytoid dendritic cells Pseudotyping of FV vectors by heterologous viral glycoproteins
	2014	Puławy, Poland	• • •	Discovery of BFV LTR encoded microRNAs Host cell Polo-like kinase interaction with PFV Gag involved in proviral integration Update of FV taxonomy and new nomenclature proposed
	2016	Paris, France	• •	Structure of the central domain of PFV Gag Genetic diversity of the env gene in zoonotic gorilla and chimpanzee FV strains
	2018	Dresden, Germany	• • •	Identification of ISG PHF11 as cellular restriction factor of FVs Cross species dynamics of feline FV A FV vector system for transient expression of CRISPR/Cas genome engineering tools

Acknowledgments:

The meeting was supported by grants from the Deutsche Forschungsgemeinschaft (DFG, German Research Foundation, Projektnummer 406942900); from the Boehringer Ingelheim Stiftung; and the TU Dresden's Institutional Strategy (F-003661-533-Ü5D-1250000), funded by the Excellence Initiative of the German Federal and State Governments. No funding was received for covering the costs to publish this meeting report in open access.

Author Contributions: All authors participated in the writing and editing of the manuscript. All meeting participants approved their presentation summaries and consented to mentioning of their names and affiliations.

Conflicts of Interest:

The authors declare no conflict of interest. The funding agencies had no role in the decision to publish this report.


Vitor Dias Tarli

Philippe Grandcolas

Roseli Pellens

(Blattodea: Blaberidae, Blaberinae) 1 from the South American Atlantic forest, with the descriptions of five new species 2 3

Keywords: Neotropical Atlantic Forest, endemic cockroach, male genitalia, juvenile stages, 30 distribution range, review

The genus Monastria Saussure, 1864 includes medium to large sized (40-55 mm) dark brown or black 13 cockroaches found in the understory of the Neotropical Atlantic Forest from the Northeast of Brazil to 14 Paraguay and Argentina. The genus shows evident sexual dimorphism: males are elongated with fully 15 developed wings extending beyond the apex of cerci and females are oval and brachypterous. This 16 study is a revision of the genus with redescription of the three already known species, Monastria 17 biguttata (Thunberg, 1826), Monastria similis (Serville, 1838) and Monastria angulata Saussure, 18 1864, and description of five new ones, Monastria itubera sp. n. and Monastria itabuna sp. n. from 19 state of Bahia, Monastria cabocla sp. n. from state of Sergipe, Monastria kaingangue sp. n. from state 20 of São Paulo and Monastria sagittata sp. n. from state of Minas Gerais. The morphology of the genus 21 and all species is described in details, including male genitalia. Based on this revision, we proposed a 22 new combination for Hiereoblatta papillosa (Thunberg, 1826) comb. n., excluding it from the genus 23 Monastria. For the first time, the juvenile stages of Monastria are characterized and compared to other 24 genera of Blaberinae of the Atlantic forest. Three determination keys are provided. The two first are 25 aimed at identifying the adults and juvenile stages of the five genera of the Blaberinae radiation 26 endemic to the Atlantic forest, respectively. The third concerns the identification of the species of the 27 genus Monastria. A map indicating the localities where species were sampled is also provided.

Introduction

54

Cockroaches have evolved with diverse continental radiations. One of the most conspicuous 55 corresponds to the subfamilies Zetoborinae (Grandcolas, 1993a;Grandcolas & Pellens, 2002; 6 H), L2d (Figs. 5F,I) and R (right phallomere) (Figs. 5D,E). Sclerite L1 long and thin with the apical 186 part distinct, quite sclerotized and the left branch tooth-shaped; central portion with a small posteriror 187 projection and a branch on the right side directed upwards; apical region with a crown of sclerotized

192

R3v with a flattened and inverted v shape in ventral view; rounded anterior lateral apex, and short 193 caudal branch (Fig. 5E).

195

Female (Figs. 4,. Species of medium size (30-45 mm). Head rounded, with wide interocular 196 space measuring 1/2 the distance between the antennal sockets (2.0-3.1 mm). Eyes reniform with 197 straight interocular margin. Antennae reaching the apex of the tegmina, filiform and setose from the 198 eighth flagellar segment. Ocelli developed and deflected. Front broad and frontal suture with a 199 cuticular invagination (Fig. 6D). Pronotum subtriangular with anterior region rounded and slightly 200 concave near the margins, dorsal region rough with striae, lateral angles rounded ending in a corner,

201

posterior margin slightly curved in the median region (Fig. 6B). Brachypterous. Tegmina truncated 202 with a marked curvature towards the interior and not extending further than the second abdominal 203 tergite; wings much shorter and undeveloped (Fig. 6A). Legs short and robust. Fore femora ventro-204 anterior margins with 13 spines of the same size, the last one close to the apex; ventro-posterior 205 margins with 4 spines, the last one close to the apex. Middle legs ventro-anterior margins with 4 206 spines, the last one close to the apex. Supra-anal plate bilobed with a small median incision, each lobe 207 with slightly rounded lateral and straight posterior margin (Fig. 6C). Tergites with slightly rounded 208 lateral angles (Fig. 6A).

210

Juvenile stages. Juveniles of both sexes are oval, almost rectangular with a body length and width 211 ratio ≈ 1,7:1. Beige to light brown, uniformly colored, the dorsal surface of pronotum and tergites is 212 very rough (Fig. 7), covered by thorn-like cuticular microsculptures, often more abundant at their hind 213 margins. (Fig. 3A). Head, eyes, antennae, legs, supra-anal and subgenital plate very similar to those in 214 adults (Fig. 7E).

216

Habitat and Behavior. Specimens of Monastria were observed in a large array of forest ecosystems, 217 ranging from semi-deciduous forests at the Northeast to the humid montane forests in the central 218 region and the Araucaria forests in the South. All specimens were observed and collected in the forest 219 understory on the underside of dead trunks of various sizes, as it was described in details for M. 220 biguttata (Pellens & Grandcolas, 2003). Very often found grasping on the underside of the bark or the

271

Monastria biguttata Saussure, 1864a: 256;Saussure, 1864b: 348;Walker, 1868: 11;Kirby, 1904: 161; 272 Shelford, 1907Shelford, -1908: 469;: 469;[START_REF] Rehn | Notes on Paraguayan Orthoptera, with descriptions of a new genus and four new 964 species[END_REF]: 248, Rehn, 1913: 282, Rehn, 1915: 275, Rehn, 1920: 217; 273 Hebard, 1921: 246;Princis, 1949: 66;Princis, 1958: 75, Princis, 1963: 141.

274

Blabera nigripennis Walker, 1868: 6;Finot, 1897: 210;Princis, 1963: 142 (as syn. of M. biguttata).

275

Tarraga guttiventris Walker, 1868: 16;Finot, 1897: 213;Princis, 1963: 142 (as syn. of M. biguttata).

276

Blabera monstrosa Stâl, 1855: 351;Kirby, 1904: 165;Princis, 1963: 142 (as syn. of M. biguttata).

277

Blabera mostruosa Sjostedt, 1933: 10;Princis, 1963: 142 (as syn. of M. biguttata, lapsus calami).

295

Diagnosis. This species is characterized by the presence of two additional cuticular depressions in the 296 frons near the clypeus. L1 sclerite with lateral branch slightly curved forward, with the aspect of a big 297 sharp tooth, and with spines in the ventral region near the crown of spines; a pointy medial region with 298 a projection in the right side turned forward. R2 sclerite cleft very sclerotized, rounded at the side of N 299 sclerite. N sclerite tiny in the dorsal region, and large and globular in the ventral one. R3d slightly 300 curved dorsally and wider and straight in ventral view. R3v sclerite with a wide and rounded latero-301 distal region and a short and slightly narrow caudal branch.

303

Redescription. Male. Head subtriangular, with interocular space measuring approximately 1/5 of the 304 distance between the antennal sockets. Ocelli developed and slightly deflected. Frontal suture with a 305 cuticular invagination. Two additional cuticular depressions in the frons near the clypeus (Fig. 5C).

306

Pronotum transverse, pentagonal, dorsal surface rough with striae, fore margin rounded, lateral 307 margins with sharp angles and round ends, hind margin nearly straight (Fig. 5B). Legs short and 308 robust. Fore-femora ventro-anterior margins with 16 spines of the same size; ventro-posterior margins 309 with 3 spines. Middle-legs ventro-anterior margins with 3 or 4 spines. Supra-anal plate quadrangular 310 with setae on the surface, hind margin straight and slightly rounded laterally, bilobed with a strong 311 invagination in the median portion. Subgenital plate asymmetrical with long styles, funneled in the 312 apical region.

313

L1. With the left side of the apical region with a lateral branch (Lb) slightly curved forward, with the 314 aspect of a big sharp tooth with spines near the crown of spines. The right side of the apical region 315 with a pointy medial protuberance and a projection turned forward (Fig. 5G,H). L2d sclerite hook 316 with a soft subconical and slightly narrow anterior region, a short apical sclerotized region, and a wide 317 space connecting them. Apex internal cavity concave, short lateral-external margin and with a typical 318 sub-apical notch (Fig. 5F,I). R2 sclerite cleft very sclerotized, rounded and curved inward with the 319 apex directed upward (Fig. 5D). Sclerite N tiny in the dorsal region, and large and globular in the 320 ventral one. R3d slightly curved dorsally and wider and straight in ventral view. Sclerite R3v with a 321 flattened and inverted v shape in ventral view; rounded anterior lateral apex, and short caudal branch.

322

(Fig. 5D,E).

324

Female. Head rounded, with wide interocular space measuring 1/2 the distance between the antennal 325 sockets. Eyes with slightly inclined interocular margins. Ocelli developed and deflected. Front broad 326 and frontal suture with a cuticular invagination (Fig. 6D). Pronotum subtriangular with anterior region 327 rounded and slightly concave near the margins, dorsal region rough with striae, lateral angles rounded 328 ending in a corner, hind margin slightly curved in the median region (Fig. 6B). Tegmina with anterior 329 margin slightly triangular; hind margin truncated with a marked curvature inside; very short and not 330 extending further than the second abdominal tergite (Fig. 6A). Legs short and robust. Fore femora 331 ventro-anterior margins with 13 spines of the same size, ventro-posterior margin with 4 spines. Middle 10 legs ventro-anterior margins with 4 spines. Supra-anal plate bilobed with a small median incision, each 333 lobe with a slightly rounded lateral margin and a straight posterior margin (Fig. 6C). Tergites with 334 straight lateral angles (Fig. 6A

341

Coloration. m#: General coloration brown (Fig. 5A). Pronotum dark brown with anterior margin buff 342 brown; central disk pale brown with scattered black marks (Fig. 5B). Head dark brown-black; clypeus 343 and labrum amber. Antennae with basal segments brown pigmented and apical segments pale brown.

344

Ocelli pale brown (Fig. 5C). Legs and spines dark brown; tarsal claws, pulvilli and arolia amber.

345

Tegmina and abdomen following general coloration of body (Fig. 5A) but with dark brown tergites 346 and sternites hind margins.

348

Distribution. Brazil (Rio de Janeiro, Espirito Santo). See Fig. 17

355

Monastria similis Saussure, 1864a: 256;Walker, 1868: 11;Scudder, 1868: 54;Kirby, 1904: 161; 356 Princis, 1958: 75, Princis, 1963: 142.

357

Monachoda similis Brunner v. W, 1865: 367;Saussure, 1870: 120;Finot, 1897: 207.

358

Monastria flavomarginata [START_REF] Princis | Colombianische Blattodeen, gesammelt von Herrn G. Dahl und Frau M. Althin-951 Dahl in den Jahren 1936-1939[END_REF]: 162, Princis, 1963: 142 (as

374

São Paulo: 1m# Poa, 13 IV 1963, Rabello col. (MZUSP). 1m# Salesópolis, "Estação biológica de 375 Boracéia, Trilha do poço verde", 23°38'56.9"S, 45°52'50"W, Malaise, pt1, 17 -28 IV 2003, coll. A.P.

376

Aguiar & F.M. Rodrigues (MZUSP). 1m# Salesópolis, "Est. biológica de Boracéia", 850m, (MZUSP).

390

Redescription. Male. Head subtriangular, with interocular space measuring nearly 1/3 of the distance 391 between the antennal sockets. Ocelli developed and slightly deflected. Frontal suture with a slight 392 cuticular invagination (Fig. 8C). Pronotum transverse, pentagonal, dorsal surface rough with deep 393 striae, anterior margin rounded, lateral margins with sharp angles and round ends, hind margin nearly 394 straight (Fig. 8B). Tegmina with orange lateral margins in the first half of its length (Fig. 8A). Legs 395 short and robust. Fore-femora ventro-anterior margins with 21 or 22 spines decreasing in size from 396 basal to apical; ventro-posterior margins with 3 long and 2 slightly smaller spines. Middle legs ventro-397 anterior margins with 10 spines. Supra-anal plate quadrangular with setae on the surface, hind margin 398 straight and slightly rounded laterally, bilobed with an invagination in the median portion. Subgenital 399 plate asymmetrical with long styles, funneled in the apical region. L1 sclerite with apical region with 400 two distinct parts: a short, rounded lateral branch turned downwards, with strong spines near the 401 crown of spines of the posterior region; a wide region slightly concave with a slight curvature ending 402 in a small projection towards the front on the right (Fig. 8H,I). L2d sclerite hook with anterior region 403 subconical and slightly wide and a short apical region; internal cavity strongly concave, external 404 lateral margin very near the very narrow sub-apical notch (Fig. 8F,G). R2 sclerite cleft curved inwards 405 with the apex very sclerotized and directed upwards; shorter and forming a right angle (Fig. 8D).

406

Sclerite N dorsal region reduced, ventral surface wide and globular. R3d slightly curved dorsally and 407 wider and straight in ventral view. R3v small, triangular, with a short and slightly quadrangular latero-408 distal region; caudal branch short with a finger shaped apex (Fig. 8D,E).

410

Female. Head rounded, with wide interocular space measuring 1/2 of the distance between the 411 antennal sockets. Eyes with straight interocular margin. Ocelli small and located on the area slightly 412 deflected above the antennal socket. Frons broad and flattened, frontal suture with a small cuticular 413 depression. Clypeus distal half larger and yellow transparent (Fig. 6H). Pronotum subtriangular with 414 anterior region rounded and slightly concave near the margins, dorsal region rough with deep striae, 415 lateral angles rounded ending in a corner, hind margin slightly curved in the median region (Fig. 6F).

416

Tegmina with a rounded anterior margin, orange lateral margin, and rounded and curved hind margin.

417

Long, reaching the fifth abdominal tergite (Fig. 6E). Legs short and robust. Fore-femora ventro-

418

anterior margins with 21 or 19 long spines of the same size, ventro-posterior margins with 5 spines.

419

Middle legs ventro-anterior margins with 5 spines. Supra-anal plate bilobed with a small median 420 notch, each lobe with a slightly rounded lateral and a straight hind margin (Fig. 6G). Tergites with 421 rounded lateral angles (Fig. 6E).

423

428

Coloration. m#: General coloration brown (Fig. 8A). Pronotum dark brown with anterior margin buff 429 brown; central disk orange with large dark brown mark (Fig. 8B). Head dark burnt umber; clypeus 430 amber and labrum orange. Antennae with basal segments brown pigmented and apical segments pale 431 brown. Ocelli white (Fig. 8C). Legs brown and spines with brown base and dark brown apex; pulvilli 432 yellowish white; tarsal claws and arolia amber. Tegmina with orange lateral margins (Fig. 8A).

433

Abdomen following general coloration of body but with brown tergites and sternites hind margins.

441

Monastria angulata Saussure, 1864a: 257;Walker, 1868: 11;Kirby, 1904: 161;Princis, 1963: 142. deflected. Front broad and frontal suture with a small cuticular invagination (Fig. 9D). Pronotum 455 triangular, with anterior region rounded and slightly concave near the margins; dorsal roughness and 456 striae inconspicuous. Lateral angles pointy, without corners, hind margin slightly curved in the median 457 region (Fig. 9B). Tegmina very short and not extending further than the second abdominal tergite; 458 anterior angle nearly square, hind margin truncated with a marked curvature towards the medial 459 region. CuP vein ending at the half of the hind margin (Fig. 9A). Legs short and robust. Fore-femora 460 ventro-anterior margins with 16 or 17 spines of the same size, ventro-posterior margins with 4 spines.

461

Middle legs ventro-anterior margins with 4 spines. Supra-anal plate bilobed with a small median 462 notch, each lobe with a very rounded lateral margin and a slightly straight hind margin (Fig. 9C).

463

Tergites with quadrangular lateral angles (Fig. 9A).

471

Coloration. Syntype #f: General coloration burnt umber (Fig. 9A). Pronotum burnt umber with a 472 anterior margin ochre; central disk yellowish brown with scattered dark brown marks (Fig. 9B). Head 473 burnt umber and clypeus and labrum dark brown (Fig. 9D). Antennae with segments yellowish brown.

474

Ocelli ochre. Legs and spines dark brown; tarsal claws, pulvilli and arolia yellowish brown. Tegmina 475 with anal field burnt umber and marginal field brown; abdomen following general coloration of body 476 but with tergites and sternites hind margins dark brown (Fig. 9A). Supra-anal plate burnt umber (Fig.

513

Crown with a smaller number of sclerotized spines. (Fig. 10G,H). L2d sclerite hook with subconical 514 and very wide anterior region, a median apical region, and a narrow space connecting them. Apex 515 internal cavity concave with short lateral external margin and narrow subapical notch (Fig. 10F,I). R2 516 sclerite cleft sclerotized, curved inward with a conic opening at its base and an apex directed upwards.

517

Sclerite N triangular on the dorsal region (Fig. 10D) and with a small surface in the ventral one. R3d 518 with a slight depression in the dorsal median region and narrow in ventral view (Fig. 10D,E). R3v 519 sclerite with a long, rectangular and slightly wide latero-distal region, and quadrangular caudal branch 520 very near R3d (Fig. 10E).

522

Female (Allotype). Head rounded, with wide interocular space measuring 1/2 of the distance between 523 the antennal sockets. Eyes with curved interocular margin. Ocelli developed and deflected. Frons 524 broad, frontal suture with a large cuticular invagination (Fig. 9H). Pronotum subtriangular, dorsal 525 surface slightly rough and striated, with two quite evident lobes with plain surface, and a wider central 526 lobe covering the head; anterior margin rounded, lateral margins short with sharp angle and a 527 conspicuous small spine, hind margin nearly straight (Fig. 9F). Tegmina latero-anterior angle nearly 528 straight; lateral margin wider, and hind margin with a strong curvature near the radial vein; extends 529 further than the third abdominal tergite. CuP vein very marked (Fig. 9E). Legs short and robust. Fore-530 femora ventro-anterior margins with 19 small spines of the same size, ventro-posterior margins with 4 531 spines. Middle legs ventro-anterior margins with 9 spines. Supra-anal plate bilobed with a median 532 incision, and each lobe with straight posterolateral angles (Fig. 9G). Coloration. Holotype #m: General coloration sienna brown (Fig. 10A). Pronotum sienna brown with a 546 brown anterior margin; central disk seal brown with scattered dark marks (Fig. 10B). Head reddish 547 brown; clypeus and labrum amber. Antennae with basal segments dark brown pigmented and apical 548 segments brown. Ocelli pale brown (Fig. 10C). Legs and spines dark brown; tarsal claws, pulvilli and 549 arolia whitish brown. Tegmina sienna brown with marginal and scapular field with a buff brown part 550 (Fig. 10A). Abdomen following general coloration of body.

534

Measurements (mm

552

Etymology. A term from the Tupi meaning taken out of the forest. Here it refers to the habitat and the 553 color patterns of this species.

554

directed upwards (Fig. 11D). Sclerite N wider and near of R2 on the dorsal region, and a narrow and 592 small ventral surface. R3d narrow in the proximity of R2, and wider and rectangular in the distal 593 region when in ventral view (Fig. 11D,E). R3v sclerite small, subtriangular, with rounded latero-distal 594 end (Fig. 11E).

596

Female (Allotype). Head rounded, with interocular space measuring approximately 1/3 of the distance 597 between the antennal sockets. Eyes reniform with rounded interocular margin. Ocelli developed and 598 deflected. Concavity between eyes and ocelli reaching the frontal suture. Frons with a prominence 599 (Fig. 12D). Pronotum subtriangular, anterior region rounded with a depression near the margins; 600 dorsal surface rough with deep striae, presence of small spine at the end of lateral region, hind margin 601 nearly straight (Fig. 12B). Tegmina with rounded lateral anterior angle, hind margin truncated with a 602 slight curvature in the region of radial vein and merging to the CuP vein at the end, not extending 603 further than the second abdominal tergite (Fig. 12A). Legs short and robust. Fore-femora ventro-

604

anterior margins with 21 or 20 spines of the same size, ventro-posterior margins with 4 spines.

605

Middle-legs ventro-anterior margin with 8 spines, plus one near the apex. Supra-anal plate bilobed 606 with a median incision and each lobe slightly rounded (Fig. 12C).

628

rounded and slightly concave near the margins, dorsal region rough with deep striae, lateral margins 666 rounded ending in a small corner, hind margin slightly curved in the median region (Fig. 12F).

667

Tegmina with straight lateral anterior angle, hind margin truncated with a slight curvature in the region 668 of radial vein; not extending further than the second abdominal tergite and lateral margin orange all 669 along its extension (Fig. 12E). Legs short and robust. Fore-femora ventro-anterior margins with 19 670 small spines of the same size; ventro-posterior margins with 4 spines. Middle-legs ventro-anterior 671 margins with 9 spines. Supra-anal plate bilobed with a median incision and each lobe slightly rounded 672 (Fig. 12G). Tergites with nearly straight lateral angles (Fig. 12E).

674

679

× 13.00 width; interocular width -3.0; interantennal width between sockets 5.7.

681

Coloration. Holotype #m: General coloration dark brown (Fig. 13A). Pronotum dark brown with the 682 anterior margin yellowish brown; central disk yellowish brown with scattered black marks (Fig. 13B).

683

Head dark brown; clypeus and labrum yellowish brown. Antennae with basal segments dark 684 pigmented and apical segments whitish yellow. Ocelli yellowish/pale brown (Fig. 13C). Legs and 685 spines brown; tarsal claws, pulvilli and arolia yellowish brown. Tegmina and abdomen following 686 general coloration of body but with tergites and sternites with brown posterior margin (Fig. 13A).

688

Etymology. The specific name refers to the type locality Itabuna, state of Bahia, northeast Brazil.

690

Distribution. Brazil (Bahia). See Fig. 17

700

Paratypes -Santa Catarina: 1#m, Urubici, "Parque Nacional de São Joaquim, Vacas Gordas", 977m, 701 28°08'44.4"S, 49°37'09.3"W, 22 X 2015, coll. R. Pellens (MNHN). 2m#, 1f#, São Bonifácio, Santo 702 the margins; dorsal region rough with deep striae, lateral angles rounded ending in a corner and hind 740 margin slightly curved in the median region (Fig. 15B). Tegmina with little rounded lateral anterior 741 angles, orange lateral margins, and rounded and curved hind margins. Long, reaching the fifth 742 abdominal tergite (Fig. 15A). Legs short and robust. Fore-femora ventro-anterior margins with 31 or 743 32 spines of the same size; ventro-posterior margins with 3 spines. Middle legs ventro-anterior 744 margins with 5 spines. Supra-anal plate bilobed with a very small median incision. Lobe lateral 745 margins slightly rounded and posterior margin straight (Fig. 15C). Tergites with rounded lateral angles 746 (Fig. 15A).

780

Diagnosis. This species is characterized by having pronotum with shallow, smooth and slightly 781 striated dorsal surface; wide lateral margins with sharp angles. Tegmina anal field wide with the CuP 782 vein perpendicular to the lateral. Subgenital plate asymmetrical with one small style. L1 sclerite with a 783 distinct spear shaped apical region, hind margin with a large non-sclerotized projection, and fore 784 region with a small projection directed upwards. Lateral branch small with strong teeth and spines.

785

L2d sclerite, hook with subconical and narrow anterior region, very short and curved and internal 786 ventral margin strongly convex.

788

Description. Male. Head subtriangular, with interocular space measuring approximately 1/3 of the 789 distance between the antennal sockets. Ocelli developed and slightly deflected; frontal suture with a 790 cuticular invagination (Fig. 16C). Pronotum broad, transverse and pentagonal; dorsal surface shallow, 791 smooth and slightly striated; anterior margin rounded; lateral margins wider with sharp angles; hind 792 margin nearly straight (Fig. 16B). Tegmina characterized by a wide anal field with the CuP vein 793 perpendicular to the lateral (Fig. 16A). Legs short and robust. Fore-femora ventro-anterior margins 794 with 21 or 20 spines of the same size; ventro-posterior margins with 4 spines. Middle-legs ventro-795 anterior margins with 5 or 4 large spines. Supra-anal plate quadrangular with setae on the surface, hind 796 margin straight and slightly rounded laterally, bilobed with narrower lobes and strong invagination in 797 the median portion. Subgenital plate asymmetrical with one small style. L1 sclerite with a roughly 798 triangular spear-shaped apical region, hind margin with a large non-sclerotized projection, fore region 799 with a small lateral projection directed upwards, and lateral branch small with strong teeth and spines 800 (Fig. 16G,H). L2d sclerite hook with subconical and narrow anterior region, very short, curved and 801 internal ventral margin strongly convex (Fig. 16F,I). R2 sclerite cleft slightly curved inwards with the 802 apex slightly sclerotized (Fig. 16D,E). The other sclerites were partially destroyed with an unhappy

812

Coloration. Holotype m#: General coloration pale brown (Fig. 16A). Pronotum pale brown with a 813 buff anterior margin; central disk brown ochre with scattered dark brown marks (Fig. 16B). Head with 814 dark brown interocular space, orange frons and amber clypeus and labrum. Antennae with segments 815 yellowish brown. Ocelli brown (Fig. 16C). Legs and spines yellowish brown; pulvilli yellow whitish, 816 tarsal claws and arolia yellowish brown. Tegmina and abdomen following general coloration of body 817 (Fig. 16A).

828

Monastria papillosa Shelford, 1907Shelford, -1908: 467;: 467;Kirby, 1910: 565;Princis, 1949: 66;[START_REF] Princis | Blattariae: Suborde Polyphagoidea: Fam.: Homoeogamiidae[END_REF] 829 142.

831

Among all the material examined to revise the genus Monastria was the type of Blatta papillosa 832 Thunberg, 1826. This species was assigned to Monastria very early and maintained until today after 833 being synonymized several times. As can be seen in the Fig. 18, the coloration, pronotum, head, 834 tegmina and general shape of this specimen does not correspond to those that characterize the genus 835 Monastria. In fact, the general characteristics of this species are in accordance with the genus 836 Hiereoblatta [START_REF] Rehn | New or little known Neotropical Blattidae (Orthoptera). Number four[END_REF] (see Fig. 1D) as described in our identification key. Here, for the sake of

  188 spines. Sclerite L2d hook-shaped, like in most species of Ectobiidae and Blaberidae. Distal area 189 elongated with subapical notch. Sclerite R (right phallomere) formed by sclerotized regions R2, R3d, 190 R3v and N (Figs. 5D). Sclerite R2 ("cleft") curved, deep and directed upward. Sclerite R3d wide and 191 elongated longitudinally with a dorsal part reduced and a large ventral one in the distal region. Sclerite

  221wood of dead trunks, immediately freezing when disturbed. The body of the juveniles are covered by

  -Rio de Janeiro: 7m# 3f# Niterói. "Parque da Cidade", 225m, 280 22°55'42"S, 43°05'10"W, 500m de la plage du saco de São Francisco, Forêt semp. humide, 21 IX 281 2009, R. Pellens & P. Grandcolas rec. (MNHN). 1m# Angra dos Reis, Ilha Grande, "Sentier 282 Abraão/Enseada das Palmas", 50m, 23°08'34''S, 44°08'91''W, 21km E Angra Dos Reis, Forêt semp. 283 humide, 07 VIII 2007, R. Pellens & P. Grandcolas rec. (MNHN). 1m#, 1f# Visconde de Mauá, "Apa 284 da Serrinha do Alambari", 528m, 22°23'16.4"S, 44°29'58.1"W, 26 XI 2010, R. Pellens & P.

285

  Grandcolas rec. (MNHN). 1m# Montagnes des Orgues, Prov. de Rio de Janeiro, Environs de la Tijuca, 286 E. R.Wagner -1902 (MNHN). 1m# Floresta da Tijuca, V 1966, M. Alvarenga col. (MZUSP). 1f# 287 "Tejuca", I 1857, coll. H. Clark (NHM). Espirito Santo: 2m#, 3f# Res de Linhares, "CVRD", 288 19°09'10.2"S, 40°11'07.8"W, 19 X 1999, 40km NNE Linhares, Forêt semi-décidue, "Mata Alta", R. 289 Pellens & P. Grandcolas rec. (MNHN). 1m# Linhares, Fragment "Sitio São Pedro", 19°09'14.2"S, 290 40°11'34.3"W, 11 VII 2005, 40km NNE Linhares, Forêt semi-décidue, R. Pellens & P. Grandcolas 291 rec. (MNHN). Santa Teresa, Est. Biol. Santa Lucia, 810m, 19°58'18.5"S, 40°32'07.6"W, 6 -9 IV 292 2001, Malaise, ponto 3 trilha, C. O. Azevedo & equipe col. (MZUSP). Without locality: 3f# 293 (America Meridionale) (MHNG). 2m#, 6f# (NHM).

  for details.

377

  Paraguay: 1f# Asunción, sept.1922-apl.1923, E.G.Kent, B.M. 1925 -262 (NHM). 1m#, Carlos 378 Pfanni (MHNG) Without locality: 1m# Brésilien, H. Fruhstorfer, 1903 -321(NHM). 2f# (NHM).

  3791m# (America Meridionale/Colombie?), D. Ruhl (MHNG). 1m# coll. Pantel, co-type (MNHN).

  Male -The apical left side of L1 with a short rounded lateral branch turned downwards 382 with strong spines near the crown of spines and a wide region slightly concave with a slight curvature 383 ending in a small projection towards the front on the right side. L2d sclerite hook with internal cavity 384 strongly concave, external lateral margin very near the very narrow sub-apical notch. R2 sclerite 385 shorter and forming a right angle. Sclerite N reduced in the dorsal region and a wide and globular 386 ventral surface. R3d slightly curved dorsally and wider and straight in ventral view. R3v small, 387 triangular, with a short and slightly quadrangular latero-distal region; caudal branch short with a finger 388 shaped apex.

  (São Paulo, Paraná, Santa Catarina, Rio Grande do Sul), Paraguay (Asunción).436See Fig.17for details.

  Figure 9A-D.

442

  Monachoda angulataSaussure, 1870: 120; Finot, 1897: 207. 443 13 444 Examined material. Syntype f#. Brazil, Bahia. (MHNG) 445 446 Diagnosis. Pronotum triangular, with rounded anterior region slightly concave near the margins, 447 inconspicuous dorsal roughness and striae. Lateral angles pointy, without corners, hind margin slightly 448 curved in the median region. Tegmina anterior angle nearly square, hind margin truncated with a 449 marked curvature towards the medial margin. CuP vein ending at the half of the hind margin. Supra-450 anal plate bilobed, lateral margins of each lobe very rounded. 451 452 Redescription. Female. Head subtriangular, with wide interocular space measuring 1/2 of the 453 distance between the antennal sockets. Eyes with straight interocular margin. Ocelli developed and 454

  ). Syntype #f: Body length 30.70; pronotum length 10.00 × 17.57 maximum 468 width ; tegmen length 9.80 × 12.20 width; interocular width 2.6; interantennal width between sockets 469 5.6.

  Holotype #m, Brazil, Sergipe, Santo Amaro das Brotas, "Gravata" 13 I 1979 (MZUSP).486Allotype #f and Paratypes 1#m, 2#f, same data as holotype (MZUSP). 1#f Sergipe, Santo Amaro das 487 Brotas, "871", 10°46'51.2"S, 37°03'22.8"W (coordinates assigned à posteriori) (MNRJ).

  Frons depressed below the antennal sockets and prominent above the ocelli. Ocelli 490 positioned laterally. Pronotum dorsal surface slightly rough and striated, with two quite evident lobes 491 with plain surface, and a wider central lobe covering the head. Short lateral margins with sharp angle 492 and a conspicuous small spine. Legs longer and less robust. L1 sclerite with a lateral branch little 493 sclerotized slightly curved downwards, smooth and with only a small spine and a region with a big 494 lateral dilatation and an edge in the posterior median region with a lateral projection turned forward. 495 Crown with a smaller number of sclerotized spines in the posterior region. L2d sclerite, hook with 496 subconical and very wide anterior region. Sclerite N triangular on the dorsal region. R3d with a slight 497 depression in the dorsal median region. 498 499 Description. Male. Head subtriangular, with interocular space measuring 1/4 of the distance between 500 the antennal sockets. Frons with a depression below the antennal sockets and prominent above the 501 ocelli. Frontal suture localized in a deep cuticular invagination, ocelli developed and positioned 502 laterally (Fig.10C). Pronotum transverse, pentagonal, dorsal surface slightly rough and striated, with 503 two quite evident lobes with plain surface and a wider central lobe covering the head, anterior margin 504 rounded, short lateral margins with sharp angle and a conspicuous small spine, hind margin nearly 505 straight (Fig.10B). Legs longer than in other species and less robust. Fore-femora ventro-anterior 506 margins with 24 or 22 spines slightly decreasing in size from basal to apical; ventro-posterior margins 507 with 5 spines. Middle legs ventro-anterior margins with 5 spines. Supra-anal plate quadrangular with 508 setae on the surface, hind margin straight and slightly rounded laterally, bilobed, lobes very narrow 509 and with a strong invagination in the median portion. Subgenital plate asymmetrical with long styles, 510 funneled in the apical region. L1 sclerite apical region with two distinct parts: a lateral branch little 511 sclerotized slightly curved downwards, smooth and with only a small spine, and a region with a big 512 lateral dilatation and an edge in the posterior median region with a lateral projection turned forward.

  ). Holotype #m: Body length 52.98; pronotum length 12.61 × 18.49 maximum 609 width; tegmen length 42.22 × 15.43 width; interocular width 0.4; interantennal width between sockets 610 2.6. Paratype #m: Body length 40.98; pronotum length 09.21 × 15.14 maximum width; tegmen length 611 31.22 × 14.31 width; interocular width 0.2; interantennal width between sockets 1.8. 612 613 Allotype #f: Body length 35.79; pronotum length 9.70 × 17.05 maximum width; tegmen length 8.90 × 614 11.15 width; interocular width 1.5; interantennal width between sockets 4.5.615 616Coloration. Holotype #m: General coloration brown (Fig.11A). Pronotum brown with the anterior 617 margin pale brown; central disk pale brown with scattered black marks (Fig.11 B). Head dark 618 brown/black; clypeus and labrum brown. Antennae with basal segments dark pigmented and apical 619 segments whitish yellow. Ocelli light brown (Fig.11C). Legs dark brown. Spines and tarsal claws 620 brown, pulvilli and arolia pale brown. Tegmina and abdomen following general coloration of body, 621 but with tergites and sternites dark hind margins (Fig.11A).

  The specific name refers to the type locality Itubera, state of Bahia, northeast Brazil 624 625 Distribution. Brazil (Bahia). See Fig. 17 for details. 626 627 Monastria itabuna Tarli, Grandcolas & Pellens sp. n.

  Holotype #m, Brazil, São Paulo, Campinas. "Pico das Cabras". 22°54'23.9"S 697 46°49'34.4"W. 14 october 2014, coll. V. M. Ghirotto (MZUSP). Allotype: 1#f , same data as holotype 698 (MZUSP).

  Holotype m#: Body length 52.80; pronotum length 12.05 × 17.35 maximum 749 width; tegmen length 40.75 × 14.70 width; interocular width 1.7; interantennal width between sockets 750 5.5. Paratypes m#: Body length 52.60 -64.30; pronotum length 11.15 -14.13 × 15.08 -19.07 751 maximum width; tegmen length 36.16 -42.34 × 13.81 -20.73 width; interocular width 0.9 -1.2; 752 interantennal width between sockets 2.5 -3.1. 753 Allotype f#: Body length 42.52; pronotum length 11.03 × 17.86 maximum width; tegmen length 17.05 754 × 12.55 width; interocular width 3.1; interantennal width between sockets 5.6. Paratypes f#: Body 755 length 39.15 -47.18; pronotum length 10.04 -12.22 × 16.12 -19.41 maximum width; tegmen length 756 14.86 -18.72 × 09.92 -13.34 width; interocular width 1.4 -1.6; interantennal width between sockets757 Holotype m#: General coloration dark brown (Fig.14A). Pronotum dark brown with the 761 anterior margin yellowish brown; central disk orange brown with scattered dark brown marks (Fig.14

  Figure 16.

  Holotype m#: Body length 51.28; pronotum length 11.94 × 21.54 maximum 809 width; tegmen length 41.97 × 17.00 width; interocular width 1.8; interantennal width between sockets 810 5.4.

  The name "sagittata" derives from the Latin sagitta, meaning arrow, referring to arrow-820 shaped apex of the L1 sclerite.821 822Distribution. Brazil (Minas Gerais). See Fig.17for details.

  Figure 18.

Figure 1 . 1005 Figure 2 .

 110052 Figure 1. Five genera of Blaberinae endemic from the Neotropical Atlantic forest. Habitus, dorsal

Figure 3 .Figure 4 .Figure 5 .Figure 6 .

 3456 Figure 3. Scanning electron photographs showing dorsal cuticular microsculptures of juveniles: A)

Figure 7 .Figure 8 . 1035 Figure 9 . 1040 Figure 10 .Figure 11 . 1052 Figure 12 .Figure 13 .

 78103591040101110521213 Figure 7. Male juveniles of Monastria in different stages of development. Scales: Habitus=5mm. Note

Figure 14 . 1070 Figure 15 . 1074 Figure 16 .Figure 17 . 1084 Figure 18 .

 1410701510741617108418 Figure 14. Monastria kaingangue sp. n., Holotype ♂: A) Habitus, dorsal view; B) Pronotum, dorsal

  ).

	335
	336
	337	Measurements (mm). m#: Body length 51.70; pronotum length 12.36 × 17.35 maximum width;
	338	tegmen length 42.55 × 15.90 width; interocular width 1.0; interantennal width between sockets 5.5
	339	f#: Body length 38.50 ; pronotum length 11.20 × 17.55 maximum width ; tegmen length 10.00 × 11.25
		width; interocular width 2.3; interantennal width between sockets 5.0.

  syn. of M. similis). Florianopolis, "Chemin Mirante. route Lagoa da Conceiçao", 295m, 27°35'5.05"S, 48°28'6.08"W, 370 Forêt semp. humide, 01 XII 2008, R. Pellens & P. Grandcolas rec. (MNHN). 2f# Joinville, "Alto da 371 Serra Dona Franscisca", 755m, 26°11'882" S, 49°03'144" W, 35km NW Joinville, Forêt semp. 372 humide, III 2007, R. Pellens & P. Grandcolas rec. (MNHN). 1m# 4f# Lages, Brunner d. w. (MHNG). Rio Capivary, 1888, H. Fruhstorfer (MHNG). Rio Grande do Sul: 2m# Lhering, (MNHG).

	373
		1m# 1f#
	359
	360
	361	Examined material. Brazil -Paraná: 1f# Aurora do Iguaçu, "Fazenda Dona Iolanda", 260m,
	362	25°23'047"S, 54°07'048"W, 20km NE São Miguel do Iguaçu, Fragment=60ha, Forêt semp. Humide,
	363	18 XI 2008, Jour, coll. R. Pellens & P. Grandcolas (MNHN). 2m#, 1f# Foz do Iguaçu, "Fazenda John
	364	Keller", 220 m, 25°34'9.14"S, 54°26'7.41"W, 1km Parc National do Iguaçu, Forêt semp. humide, 20
	365	XI 2008, R. Pellens & P. Grandcolas rec. (MNHN). Santa Catarina: 1m# Blumenau, VI 1919, coll.
	366	Luderm? (MZUSP). 3m#, 2f# Campo Alegre, "Ilha, Fazenda Sr. Egon", 830m, 26°10'29''S,

49°16'22''W, 1km Campo Alegre, 14 VIII 2007, Forêt d

'Araucaria, R. Pellens & P. Grandcolas rec. 367 (MNHN). 1f# Campo Alegre, "Fazenda Sr. Gilson", 853m, 26°12'6.61"S, 49°18'4.58"W, 5km W 368 Campo Alegre, Forêt d'Araucaria, 15 VIII 2007, R. Pellens & P. Grandcolas rec. (MNHN). 1f#

  Body length 49.35; pronotum length 10.95 × 16.80 maximum width; 424 tegmen length 39.75 ×15.45 width; interocular width 1.6; interantennal width between sockets 4.6.

	425
	426	f#: Body length 36.65; pronotum length 11.35 × 16.90 maximum width ; tegmen length 16.50 × 11.00
		width; interocular width 2.6; interantennal width between sockets 5.6.

Measurements (mm). m#:

  ). Holotype #m: Body length 42.95; pronotum length 9.85 × 15.70 maximum

	535
	536	width; tegmen length 35.25× 13.45 width; interocular width 0.9; interantennal width between sockets
	537	3.6. Paratypes #m: Body length 41.54; pronotum length 09.94 × 15.20 maximum width; tegmen length
	538	31.36× 13.19 width; interocular width 0.5; interantennal width between sockets 1.8.
	539	Allotype #f: Body length 41.60 ; pronotum length 13.20 × 20.15 maximum width ; tegmen length
	540	16.05 × 14.50 width; interocular width 2.9; interantennal width between sockets 6.0. Paratype #f :
	541	Body length 40.82 -41.96; pronotum length 12.15 -12.86 × 20.12 -19.91 maximum width; tegmen
	542	length 14.53 -15.06 × 12.48 -13.76 width; interocular width 1.3 -1.4 ; interantennal width between
	543	sockets 2.4 -2.5.
	544
	545

  Body length 40.70; pronotum length 11.25 × 19.80 maximum width; tegmen length 12.30

	675	Measurements (mm). Holotype #m: Body length 53.05; pronotum length 12.70 × 19.37 maximum
	676	width; tegmen length 40.40 × 15.55 width; interocular width 0.9; interantennal width between sockets
	677	2.7 . Paratype #m: Body length 54.09; pronotum length 12.22 × 21.02 maximum width; tegmen length
	678	43.12 × 19.61 width; interocular width 0.5; interantennal width between sockets 2.8.
		Allotype #f:

asymmetric. Internally, attached to this plate, a membranous pouch with genital sclerites L1 (Figs.5G,

Spolia Zoologica Musei Hauniensis, 12, 1-72.

Princis, K. (1958) Revision der Walkerschen un Kirbyschen Blattarientypen im British Museum of

Natural History, London. II. OpusculaEntomologica, 23,[59][60][61][62][63][64][65][66][67][68][69][70][71][72][73][74][75]

Male -Figure 13.

646

13C). Pronotum transverse, pentagonal, dorsal surface rough with striae; anterior margin rounded; 647 lateral margins slightly wider with sharp angles, round ends and a conspicuous small protuberance; 648 hind margin nearly straight (Fig. 13B). Legs short and robust. Fore-femora ventro-anterior margins 649 with 20 or 18 spines of the same size; ventro-posterior margins with 4 small spines. Middle-legs 650 ventro-anterior margins with 5 or 4 large spines. Supra-anal plate quadrangular with setae on the 651 surface, hind margin straight and slightly rounded laterally, bilobed with a slight invagination in the 652 median portion. Subgenital plate asymmetrical with long styles, left funneled in the apical region, right 653 with same caliper all along. L1 sclerite with apical region with two distinct parts, a lateral branch 654 slightly curved with some spines that contact the crown in the posterior region (Fig. 13G,H). L2d 655 sclerite, hook with short apical region, internal margin concave, lateral external margin strongly 656 curved, with large sub-apical notch (Fig. 13F,I). R2 sclerite cleft curved inward with the apex directed 657 forward (Fig. 13D). Sclerite N smaller in the dorsal region and with a large rough ventral surface. R3d 658 narrow near R2 in dorsal view and wider, rectangular in the distal region when in ventral view (Fig.

659

13D, E). R3v sclerite large, with quadrangular and truncated latero-distal end. Caudal branch long, 660 wide in the center and narrow in the apex (Fig. 13E).

662

Female (Allotype). Head rounded, with wide interocular space measuring 1/2 the distance between 663 the antennal sockets. Eyes with straight interocular margin. Ocelli developed and deflected. Frons

707

Diagnosis. This species is characterized by having two intercalated rows of spines in the ventro-

708

anterior margins of fore-femora; one row with 5 or 6 spines, and another with 32 or 33 spines of the 709 same size. L1 sclerite with lateral branch curved, rounded and smooth with some small spines, and a 710 curved region with grooves in its interior and a projection with a rounded end directed upwards at its

717

antennae. Frons slightly elevated. Ocelli developed and deflected. Frontal suture with a cuticular 718 invagination (Fig. 14C). Pronotum pentagonal, fore margin rounded, with rounded lateral angles, hind 719 margin slightly curved in the medial portion (Fig. 14B). Legs short and robust. Fore-femora ventro-720 anterior margins with two intercalated rows of spines, one with 6 or 5, and another with 33 or 32 721 spines of the same size; ventro-posterior margins with one large and a thin spine out of the row, 6 722 small spines. Middle legs ventro-anterior margins with 11 or 10 spines. Tegmina with orange lateral 723 margins in the first half of its length (Fig. 14A). Supra-anal plate quadrangular with setae on the 724 surface, hind margin straight and slightly rounded laterally, bilobed with a slight invagination in the 725 median portion. Subgenital plate asymmetrical with long styles, left funneled in the apical region, right 726 with same caliper all along. L1 sclerite with apical region with two distinct parts: a lateral branch 727 curved, rounded and smooth with some small spines, and a curved region with grooves in its interior 728 and a projection with a rounded end directed upwards at its right side (Fig. 14G,H). L2d sclerite hook 729 with long apical region and rounded curvature of the margin near the notch (Fig. 14F,I). R2 sclerite 730 cleft curved inward with a wide and very sclerotized apex directed forward (Fig. 14D). Sclerite N tiny 731 in dorsal and large and wide in ventral view. R3d with a clear prominence like a tooth in ventral view 732 (Fig. 14D,E). R3v sclerite with quadrangular and truncated latero-distal region, and caudal branch 733 near the tooth from R3d sclerite (Fig. 14E).

735

Female (Paratype). Head rounded, with wide interocular space measuring approximately 1/2 the 736 distance between the antennal sockets. Eyes with straight interocular margin. Ocelli developed and 737 slightly deflected. Frontal suture with a cuticular invagination. Clypeus distal half transparent yellow 738 and larger (Fig. 15D). Pronotum subtriangular with anterior region rounded and slightly concave near Atlantic forest and the determination keys provided will significantly contribute to a rapid 850 identification of the species belonging to this group that are often found together in the field.