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Oligotoma negra “Black webspinner” Figures 75 (lateral), 76 (dorsal, ventral) Plates 47 (lateral), 48 (dorsal, ventral) The respiratory system of Embioptera remains rather understudied, with the most recent detailed analysis of tracheal anatomy by Lacombe (1958, 1971). The specimen of Oligotoma shown here is a male and is notable because of the air-filled alimentary canal (fig. 77). Male webspinners do not feed, and like Ephemeroptera (fig. 21), the alimentary canal is coopted as a large air space, spanning the length of the body. In this specimen, the distention of the alimentary canal compressed tracheae against the inner body wall, making determination of morphology and assessment of homology challenging (particularly in the abdomen). Some notable differences were observed between Lacombe (1958) and the specimen here, particularly in the thorax, but her work was useful in mapping abdominal tracheae. The meso- and metathoracic wing base T 2,3- Wbr are likely present but partial in this scan, likely due to fluid infilling of this specimen as it was frozen to -20° C rather than -80° C. Although a tympanal hearing organ in the femur (of pro- and mesoleg, and occasionally hind leg) of webspinners was described by Szumik et al. (2019), there is no evidence of an air space in femur of any leg (as seen here in the foretibia of Gryllus, for example). DESCRIPTION: HEAD: H-DCT extending anteriad, curving laterally along head capsule; H-VCT slightly ventrad. H-DC present, slightly anterior of cervix. H-DCT with H-DCT-DVi running dorsad anterior of cervix, following head capsule. H-DCT runs directly anteriad, dividing into H-DVB ventrad and H-Ant. H-VCT with H-VCT-DVi extending dorsally, nearly in contact with H-DCTDVi and following head capsule laterally. H-VCT continues anteriorly, with dorsal H-DVB connection to H-DCT. After short extension, H-VCT divides into H-Lbm, and subsequently into H-MxPlp, H-Md, and H-Oc. H-Lbm branch connecting mediad as H-VC. Branch leading to H-Oc continues to H-Ft. THORAX: T 2- S with four branches: H-DCT, H-VCT, T 2-DB, and T 2-VB. T 2- CT absent. H-DCT runs directly anteriad, with T 1-DVi branching dorsad before continuing dorsally along prothoracic tergum. T 1-DC present, extending mediad near branching of T 1-DVi. H-VCT runs anteriad, with T 1- L branching ventrad just prior to a nearly 90° turn ventrad, subsequently turning anteriad along prothoracic sternite. T 1-VC present, branching off T 1- L. T 2-VB runs posteriad and slightly ventral briefly before curving dorsad; small T 2- AWL branching anteriad at apex of curve, dividing into T 2-AWba dorsad and T 2- AL continuing posteriorly toward midleg. T 2-DB continues, curving medially with several branches, likely for flight muscle, before extending posteriad as T 2-ADLT. Full DLT not visible but likely present, possibly fluid infilled; Lacombe (1958) indicates presence of dorsal connective. T 2-VB runs posteroventrad, with large T 2- FM branching dorsad and laterally. T 2-VB continues as T 2- VLT, connecting with T 3- S. T 2-VC present, branching mediad approximately halfway between T 2- S and T 3- S. T 3- S with four branches: T 3- AWL, T 2- PWL, T 3-DB, and T 3-VB. T 3- AWL running dorsad, curving posteroventrad where T 3-W-c-r splits dorsally and remaining branch continues as T 3- L. T 2- PWL connecting directly from anterior; T 2- PL branching ventrally, joining with T 2- AL after a short distance and continuing as T 2- L; T 2-Wbr continues anteriad from T 2- PWL; T 2-Wbr partial likely due to fluid infilling. T 3-DB runs mediad, dividing after a short distance into T 2-PDLT anteriorly and T 3- DLT posteriad. T 3-VB runs posteroventrad, with a connection to T 2- VLT very close to T 3- S; T 3-VB continues posteriorly as T 3- VLT, linking up with A 1- S. T 2-VLT-Vi on right side, extending past T 3- S into abdomen; unclear if T 2-VLT-Vi links with an abdominal spiracle. T 3- S on right side slightly different from left, possibly due to displacement of tracheae by distended alimentary canal air space; T 3- S on right side with T 3- VLT positioned posteriad as with right side, but with short, curving spiracular branch dorsad and anteriad, with T 2- VLT connecting from anterior; branch continues dorsad to T 2- AWL, T 2-DB, T 2- PWL split as with left side. ABDOMEN: A 1..8- S present. A 1- S modified from subsequent abdominal segments; A 1- S with five branches: A 1-DB, T 3-PWBa, T 3- VLT, and T 3- PL. A 1-DB short and running mediad, connecting with T 3- DLT from anteriad and continuing as A 1- DLT posteriad. T 3-WBr (also partial, likely due to fluid infilling) runs anteriad, continuing with small T 3-W-cu-a into trailing edge of hind wing. T 3- VLT from directly ventrad. T 3- PL runs ventrad, lateral from T 3- VLT, joining with T 3- AL and extending into hind leg as T 3- L. Segments A 2..7 likely similar but morphology difficult to determine due to distention of air-filled alimentary canal. A 2..7- DLT present, arcing slightly dorsad and usually sinuous. A n - VLT present. A 6-VC visible; other A n -VC likely present but displaced against body wall, see figure 78. Several visceral tracheae visible, but dif- ficult to determine morphology; A 3-Vi and A 4-Vi large and directly posteriad, spanning sev- eral segments. A-Cr visible at base of cerci., Published as part of Herhold, Hollister W, Davis, Steven R, Degrey, Samuel P & Grimaldi, David A, 2023, COMPARATIVE ANATOMY OF THE INSECT TRACHEAL SYSTEM PART 1: INTRODUCTION, APTERYGOTES, PALEOPTERA, POLYNEOPTERA, pp. 1-184 in Bulletin of the American Museum of Natural History 459 (1) on pages 1-184, DOI: 10.5531/sd.sp.55, http://zenodo.org/record/7730159, {"references":["Lacombe, D. 1958. Contribuicao ao estudo dos Embiidae. III. Aparelho respiratorio de Embolyntha batesi Mac Lachlan, 1877 (Embiidina). Studia Entomologica 1 (1): 177 - 195.","Lacombe, D. 1971. Anatomy and histology of Embolyntha batesi MacLahlan, 1877 (Embiidina). Memorias do Instituto Oswaldo Cruz 69 (3): 331 - 396.","Szumik, C., M. L. Juarez, M. J. Ramirez, P. Goloboff, and V. V. Pereyra. 2019. Implications of the tympanal hearing organ and ultrastructure of chaetotaxy for the higher classification of Embioptera. American Museum Novitates 3933: 1 - 32."]}

Herhold, Hollister W, Davis, Steven R, Degrey, Samuel P, Grimaldi, David A (2023): COMPARATIVE ANATOMY OF THE INSECT TRACHEAL SYSTEM PART 1: INTRODUCTION, APTERYGOTES, PALEOPTERA, POLYNEOPTERA. Bulletin of the American Museum of Natural History 459 (1): 1-184, DOI: 10.5531/sd.sp.55, URL: http://dx.doi.org/10.5531/sd.sp.55

Aposthonia guizhouensis sp. nov. Figs. 1–9 Type material. Holotype male, (ICJUST) China: Guizhou Province, Guiyang City, Guiyang Forest Park, 26.5499891° N, 106.766504° E, 1100 m, 6-VI-2019, Lu Qiu. Paratypes: four males and four females, same locality and data as holotype (ICJUST). Etymology. The species is named after the type locality, Guizhou Province. Distribution. This species is known only from Guiyang Forest Park, Guizhou Province, China. Description Alate male (n = 5). Head length 1.5–2.0 mm, width 0.9–1.2 mm; body length 10.0–11.0 mm, width 0.8–1.0 mm; forewing length 6.5–8.0 mm, hindwing length 6.5–7.0 mm. Head capsule dark brown (Figs. 1–3), near 1.5 times longer than broad with kidney-shaped small eyes, sides behind eyes near parallel, gradually converging posteriorly. Clypeus pale and near trapezoidal (Fig. 3A). Labrum membranous posteriorly and laterally, anteriorly with a dark brown, elliptical sclerite (Fig. 1). Maxillary palpi brown, five-segmented; labial palpi brown and short, threesegmented (Figs. 2, 3B). Submentum brown, near rectangular, with medial concave at both anterior and posterior margins, posterior corners slightly projected laterad (Fig. 3B). Mandible dark brown. Antennae brown throughout, at least 21-segmented (Figs. 1–2). Prothorax dark brown (Fig. 1), much shorter and narrower than head, near quadrate with subqual length and width, surface with deep transverse and longitudinal grooves; meso- and metathorax generally brown with dark margins. Wings brown with longitudinal pale stripes (Figs. 1–2); anterior medial vein (MA) not forked. All legs dark brown except the pale articulations. Hind leg with only one basitarsal papilla (Fig. 2). Abdomen brown throughout with terminalia darker. Terminalia with left (10L) and right (10R) hemitergites of segment 10 of relatively equal size (Figs. 4A, B). Process of 10L (10LP) sickle-shaped, subapically with a triangular spine pointing laterad (Figs. 4A–D); process of 10R (10RP) slightly wider and much longer than 10LP, inner margin dentate, apex pointed and curved laterad (Figs. 4A–D); epiproct (EP) sclerotized, length ca. two times longer than width, with concave lateral margins and rounded posterior margin (Figs. 4A–B). Hypandrium (H = sternite 9) broad and rounded (Figs. 4C–D). Process of H (HP) broad and near trapezoidal. Left paraproct (LPPT) broad, sclerotized, acutely pointed and hooked posterolaterally (Figs. 4A–D). Left cercus-basipodite (LCB) less developed, forming a brown plate at base of left cercus. Basal segment of left cercus (LC 1) dilated distally and slightly lobed without echinulation (Figs. 4A–D). Apterous female (n = 4). Head length 1.5–2.0 mm, width 1.0– 1.2 mm; body length 11.0–12.0 mm, width 1.2–1.5 mm. Head capsule dark (Fig. 5A–B), convex, slightly longer than wide. Eyes dark, smaller than in male. Submentum rectangular (Fig. 5B). Antennae with at least 22 segments, basal 14–17 segments brown, distal segments pale (Figs. 5A–B). Prothorax entirely pale; meso- and metathorax generally brown. Foreleg with pale femur, tibia and pale brown apical two tarsal segments; tibia of mid leg and hind leg with both ends conspicuously pale (Figs. 5A–B). Hind leg with only one basitarsal papilla (Figs. 5B). Abdomen brown throughout, similar to meso- and metathorax. Tenth tergum elongated posteriorly (Fig. 6A); tenth sternum symmetrically divided longitudinally into two lateral plates (Fig. 6B). Cerci entirely dark brown (Figs. 5A–B). DNA barcode. The obtained COI sequence of A. guizhouensis sp. nov. was 678 bp in length. The genetic distance between A. guizhouensis sp. nov. and all other sequenced congeners is a high value of over 0.2 (Fig. 7A), which excludes its identity with these congeners. The NJ and ML trees generated very similar tree topology (Figs. 7B–C), where A. guizhouensis sp. nov. clustered with an unidentified species from Mexico, Aposthonia sp. KBMEB165 (accession No. JQ907087). However, the high sequence divergence (0.2) and the remarkable geographical isolation exclude conspecific status of the two species. Remarks. The new species is assigned to genus Aposthonia instead of Oligotoma or Eosembia on the basis of the following combination of characters: absence of mesal lobing of the left cercus-basipodite (LCB) in the male; hind basitarsus with single ventral papilla in both male and female (Ross 2007, Poolprasert et al. 2011a, b). The presence of an outer subapical spine on 10LP in combination with the apically pointed and out-curved 10RP can easily separate A. guizhouensis sp. nov. from all other species of Aposthonia (Poolprasert et al. 2011a). The new species can be distinguished from A. borneensis, the only known Aposthonia species from China, by the small eyes, different shapes of 10LP, 10RP, HP and LPPT in the male and by the apparently different body color in the female (see Figs. 2–3 in Poolprasert et al. 2011a). The DNA barcode of A. guizhouensis sp. nov. also exhibits a high ratio of sequence divergence from all included sequences of A. borneensis (Fig. 7A), strongly supporting A. guizhouensis sp. nov. as a distinct different species from A. borneensis. Aposthonia guizhouensis sp. nov. is common in Guiyang Forest Park, inhabiting various habitats from dirt road in wild jungle (Figs. 8A–C) to man-made stone railings and columns (Figs. 9A–C). There are also numerous examples of adults of A. guizhouensis sp. nov. in Guiyang City. Their abundance is evidence that Guiyang City is a warm area with a high humidity and plenty of vegetable matter such as dead leaves and lichens. The Embioptera fauna in China lacks systematic investigation and is apparently underestimated. More undescribed webspinners are expected to be discovered in this vast area., Published as part of Chen, Zhi-Teng, 2022, Aposthonia guizhouensis sp. nov., a new webspinner of Oligotomidae (Insecta: Embioptera) from China, pp. 190-198 in Zootaxa 5213 (2) on pages 191-194, DOI: 10.11646/zootaxa.5213.2.7, http://zenodo.org/record/7354432, {"references":["Ross, E. S. (2007) The Embiidina of Eastern Asia, Part I. Proceedings of the California Academy of Sciences, 58, 575 - 600.","Poolprasert, P., Sitthicharoenchai, D., Butcher, B. A. & Lekprayoon, C. (2011 a) Aposthonia Krauss, 1911 (Embioptera: Oligotomidae) from Thailand, with description of a new species. Zootaxa, 2937 (1), 37 - 48. https: // doi. org / 10.11646 / zootaxa. 2937.1.3"]}

Aposthonia guizhouensis sp. nov. from Guizhou Province of China is described and illustrated. The new species is characterized by the distinctive terminalia structure in males. Aposthonia guizhouensis sp. nov. represents the second species of Aposthonia known from China. DNA barcode of the new species is sequenced and compared with its related congeners.

Eight new species of Embioptera from Brazil are described, diagnosed, and illustrated. For Anisembiidae: Chelicerca achilata Szumik, Pereyra & Juárez, sp. nov.; Saussurembia borba Szumik, Pereyra & Juárez, sp. nov. For Archembiidae: Archembia oruma Szumik, sp. nov.; Embolyntha oriximina Szumik, Pereyra & Juárez, sp. nov.; Pararhagadochir bonita Szumik, Pereyra & Juárez, sp. nov., Pararhagadochir marielleae Szumik, Pereyra & Juárez, sp. nov.; Pararhagadochir para Szumik, Pereyra & Juárez, sp. nov. For Clothodidae: Chromatoclothoda langa Szumik, Pereyra & Juárez, sp. nov. To clarify the higher classification of the Order and to have an accurate taxonomy, a species catalog and introduction to the four families present in Brazil is also detailed, including phylogenetic relationships, taxonomic actions, composition, distributions, and records maps. Herein, several taxonomic acts are proposed: (1) the synonymy of Chelicerca Ross (= Dactylocerca Ross confirmed junior synonym; = Schizembia Ross syn nov.; = Pelorembia Ross, confirmed junior synonym; = Cryptembia Ross, syn. nov.) and Saussurembia Davis (= Stenembia Ross, syn. nov.). (2) new status and delimitation for family Archembiidae Ross, stat. rev.; subfamily Archembiinae Ross, stat. rev.; subfamily Pachylembiinaestat. rev.; subfamily Scelembiinaestat. rev., and their genera included. (3) Diradius unicolor (Ross) (Teratembiidae) comb. nov., and (4) new locality records for previously cited species in the region.

A cladistic analysis of embiopterans, based on 157 species (representing 70% of the known genera) and 186 morphological characters, is presented, as well as a molecular analysis for 22 taxa using genes encoding 16S, 18S and 28S rDNA and COI. Species of all known families are included, except Andesembiidae Ross (specimens of which are in a private collection). The evidence presented supports the monophyly of four of the families (Australembiidae, Oligotomidae, Teratembiidae, and Anisembiidae). Notoligotomidae is paraphyletic and included within the Afro‐neotropical family Archembiidae (which is also paraphyletic). The genera Embia, Cleomia, Macrembia, and Dihybocercus (Embiidae) form, together with Australembiidae, a group strongly supported by morphology; the position of the remaining genera of Embiidae has two quite different resolutions. Almost 80% of the genera of Anisembiidae recently described appear as either paraphyletic or polyphyletic. Contrary to the opinion of other specialists, the major groups as well as the monophyly of some families are supported by features which have been ignored in classical approaches to the systematics of Embioptera, such as the ovipositor and cephalic and leg structures, characters with an almost perfect fit. Fil: Szumik, Claudia Adriana. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico - Tucumán. Unidad Ejecutora Lillo; Argentina

Genus Madygembia Aristov & Storozhenko, gen. nov. Type species. Madygembia rasnitsyni sp. nov., here designated. Etymology. The new genus is named after the Madygen locality and genus Embia. Gender feminine. Diagnosis. Forewings characters only. Sc reaching distal quarter of wing. RS arising near proximal third of wing and forming a long fusion (anastomosis) with MA. MP simple, desclerotized near middle. CuA with four branches directed to posterior margin of wing. Cross-veins numerous, simple. Wing membrane with wide hyaline intervenal lines. Remarks. New genus is most similar to Nestorembia but differs from latter in characters given above in the key., Published as part of ARISTOV, DANIIL S. & STOROZHENKO, SERGEY YU., 2021, A new genus of the family Alexarasniidae (Insecta: Embiida) from the Triassic of Kyrgyzstan, pp. 433-435 in Palaeoentomology 4 (5) on page 434, DOI: 10.11646/palaeoentomology.4.5.7, http://zenodo.org/record/5530302

The Recent webspinners (Embiida = Embioptera) comprise a small order of insects; there are about 400 species distributed mainly in tropical and subtropical regions (Miller et al., 2012). They are gregarious, living subsocially in galleries of fine silk which they spin from glands on their forelegs (Edgerly et al., 2002). The webspinner lineage is traceable back via Middle Cretaceous Sorellembiidae, Jurassic Sinembiidae and Late Permian–Triassic Alexarasniidae to the Early–Middle Permian Atactophlebiidae (Engel & Grimaldi, 2006; Huang & Nel, 2009; Shcherbakov, 2015).

Madygembia rasnitsyni Aristov & Storozhenko, sp. nov. (Fig. 1) Holotype. PIN 2785/2162, counterpart, stored at the Borisyak Paleontological Institute of Russian Academy of Science. Etymology. The new species is named in honor of the famous Russian paleoentomologist and hymenopterologist Prof. Alexandr P. Rasnitsyn. Diagnosis. As for the genus. Type locality and horizon. Madygen locality, Dzhailoucho near village of Madygen, Batken Region, Kyrgyzstan; Madygen Formation, Middle or Upper Triassic, Ladinian or Carnian (sensu Shcherbakov, 2015). Description. Length of imprint 20 mm. Anterior margin of forewing slightly concave near middle. Costal area crossed by numerous simple veins, as wide as subcostal area in basal quarter of wing. RS with four branches bur posterior branch fused with simple MA. Intercubital area narrow, crossed by simple cross-veins., Published as part of ARISTOV, DANIIL S. & STOROZHENKO, SERGEY YU., 2021, A new genus of the family Alexarasniidae (Insecta: Embiida) from the Triassic of Kyrgyzstan, pp. 433-435 in Palaeoentomology 4 (5) on page 434, DOI: 10.11646/palaeoentomology.4.5.7, http://zenodo.org/record/5530302, {"references":["Shcherbakov, D. E. (2015) Permian and Triassic ancestors of webspinners (Embioidea). Russian Entomological Journal, 24 (3), 187 - 200. https: // doi. org / 10.15298 / rusentj. 24.3.01"]}

Insects of the order Phasmatodea are mainly distributed in the tropics and subtropics and are best known for their remarkable camouflage as plants. In this study, we sequenced three complete mitochondrial genomes from three different families: Orestes guangxiensis, Peruphasma schultei, and Phryganistria guangxiensis. The lengths of the three mitochondrial genomes were 15,896 bp, 16,869 bp, and 17,005 bp, respectively, and the gene composition and structure of the three stick insects were identical to those of the most recent common ancestor of insects. The phylogenetic relationships among stick insects have been chaotic for a long time. In order to discuss the intra- and inter-ordinal relationship of Phasmatodea, we used the 13 protein-coding genes (PCGs) of 85 species for maximum likelihood (ML) and Bayesian inference (BI) analyses. Results showed that the internal topological structure of Phasmatodea had a few differences in both ML and BI trees and long-branch attraction (LBA) appeared between Embioptera and Zoraptera, which led to a non-monophyletic Phasmatodea. Consequently, after removal of the Embioptera and Zoraptera species, we re-performed ML and BI analyses with the remaining 81 species, which showed identical topology except for the position of Tectarchus ovobessus (Phasmatodea). We recovered the monophyly of Phasmatodea and the sister-group relationship between Phasmatodea and Mantophasmatodea. Our analyses also recovered the monophyly of Heteropterygidae and the paraphyly of Diapheromeridae, Phasmatidae, Lonchodidae, Lonchodinae, and Clitumninae. In this study, Peruphasma schultei (Pseudophasmatidae), Phraortes sp. YW-2014 (Lonchodidae), and species of Diapheromeridae clustered into the clade of Phasmatidae. Within Heteropterygidae, O. guangxiensis was the sister clade to O. mouhotii belonging to Dataminae, and the relationship of (Heteropteryginae + (Dataminae + Obriminae)) was recovered.

The articulated appendages of arthropods are highly adaptable and potentially multifunctional, used for walking, swimming, feeding, prey capture, or other functions. Webspinners (Order Embioptera) are a paragon in this context. In contrast to other arthropods producing silk, they utilize their front feet for silk production. However, employing the same leg for alternative functions rather than for pure locomotion potentially imposes constraints and compromises. We here present morphological and experimental evidence for a "passive" pressure-induced silk spinning mechanism induced by external mechanical stimuli. Furthermore, we demonstrate that, as a consequence of the conflicting functions for their front feet, webspinners have evolved a unique style of walking that reduces the potentially problematic contact between silk ejectors and the substrate. Here we answer for the first time a long-term question within this enigmatic group of insects-how webspinners can use their front feet to spin their nanoscale silk. This knowledge may open the door for experimental studies on an artificial spinning process and for future utilization in applied fields of robotics or chemistry.

A summary is presented of papers published on minor insect orders (MIO) in Zootaxa’s first 20 years, as well as the number of new species described therein. The MIO orders currently covered by the editors and summarized here include Archaeognatha (Microcoryphia), Dermaptera, Embioptera, Siphonaptera, Zoraptera and Zygentoma, as well as the hexapod classes Protura and Diplura. Both fossil and extant taxa of these groups are included in the MIO purview. The MIO editors also have frequently served as interim editors for groups temporarily without a subject-matter editor, such as Carabidae, Blattodea and Mantodea; as a backup editor for Mecoptera; and as ad-hoc editors for papers written by the editors of a different taxonomic group or for papers without a single-taxon focus. In the period 2001‒2020, descriptions of 130 new species were published, compared with 816 species in all other journals. The greatest number of species were for Protura (49), while Zoraptera had the highest proportion of papers compared to all publications (50%).

Pararhagadochir is a South American genus of Archembiidae that can be identified by its anterior margin of submentum strongly concave, caudal process of the 10th left hemitergite forked, with internal tip seemingly a hook and the external tip a flat lobe, both tips always shorter than the width of 10th left hemitergite. Six species are recorded in Brazil, being distributed in all regions. In this work, Pararhagadochir noronhensis sp. nov. is described from the archipelago of Fernando de Noronha, Brazil.

Pararhagadochir noronhensis sp. nov. (Figs. 2–6) urn:lsid:zoobank.org:act: C168BF4F-1FF9-4103-8E0D-80FED1C2D67F Pararhagadochir sp., Rafael et al. 2020: 16. Type-series. Holotype Ƌ: Brasil, PE, Fernando de Noronha, Trilha Sancho, 3°51'30''S – 32°25'50''W, 20-27.ii.2020, hand-collected, J.A. Rafael, P.C. Grossi, F. Limeira-de-Oliveira. Paratypes: 4 subadult ♀, 1 subadult Ƌ, 28 nymphs, same data as holotype [INPA]. Etymology. The specific epithet refers to the type locality: Fernando de Noronha, Pernambuco, Brazil. Diagnosis. Pararhagadochir noronhensis sp. nov. is closest to P. balteata. The male of P. noronhensis sp. nov. is 4.5 mm long, apterous; with antennal setae of uniform size over the entire length of the antennae; submentum quadrate; epiproct large, triangular, entirely sclerotized, extending the full length of the flap; the female is 7 mm long, and has 17 antennal segments. On the other hand, the male of P. balteata is 11 mm long and alate; with very long antennal setae on the basal half of segments and shorter setae on the apical half; submentum with sides caudally divergent to the basal third; epiproct small, not triangular, partially unsclerotized, not extending the full length of the flap; the female is 14 mm long and has 23 antennal segments. Description. Male (holotype). Color details (based on specimen in alcohol): predominantly chestnut brown, with the exception of two intersomital white bands on thorax (Fig. 2A, B). Body length 4.5 mm. Apterous. Head subhexagonal, as long as wide, anterior region wider than the posterior region due to the small and slightly protruding eyes, occipital margin uniformly rounded (Fig. 2C), length = 0.68 mm, width = 0.70 mm, OR = 0.73; two depigmented areas present, slightly visible (Fig. 2C); occipital region with maculae indistinct in dorsal view. Antenna (incomplete) with 11–8 segments (Figs. 2A, B). Mm visible, sclerotic, but less pigmented than Sm and well separated; submentum sclerotic, quadrangular, wider than long (2:1), anterior margin straight with the edges rounded (Fig. 2D). Hind basitarsus length = 0.16 mm, width = 0.08 mm; setae rows: 2 inner, 2 proximal, 3 outer; medial bladder present, circular, in the middle of the hind basitarsus, length = 0.02 mm, width = 0.02 mm (Fig. 4A). Terminalia. 10L subquadrate, 10R wider basally, tapering distally to form the right process (Fig. 3A). 10Rp1 prominent, sclerotic talon. MF narrow and arched (Fig. 3A). Outer flange of 10Lp entirely translucent, the inner process curved outward and with same length as the outer flange (Fig. 3A); Ep present, visible, in the cleft membrane, seemingly attached at the base of the MF. H sclerotic; Hp well developed, central, right curved, with many transverse lines on its anterior margin (Fig. 3B); Lpp present, sclerotic (Fig. 3B); LC1dp well developed, quadrate, echinulate. Length of the left cercus: LC1 = 0.27 mm, LC2 = 0.29 mm; width of the left cercus: LC1 = 0.09 mm, LC2 = 0.07 mm; length ratio of LC1/LC2 = 0.93. Female subadult (Paratype). Color details (based on specimens in alcohol): same color pattern as male, but lighter (Fig. 5A). Body length 7 mm. Head circular, as long as wide (Fig. 5C), length = 0.85 mm, width = 0.85 mm; eyes small, not protruding, OR = 0.88; three depigmented areas present, sometimes the central one barely discernible (Fig. 5C); occipital region with four pairs of maculae visible in dorsal view, two larger lateral maculae reaching half of the head length, median macula smaller, and one contiguous central cordiform macula (Fig. 5C); two more pairs of maculae visible in ventral view (Fig. 5D). Antennae with 17–17 segments (Figs. 5A, B). Mm and Sm slightly sclerotic; Sm quadrangular, wider than long (1.6:1) (Fig. 5D). Hind basitarsus slightly more robust than male, length = 0.19 mm, width = 0.10 mm; setal rows: 1 inner, 2 proximal, 3 outer; medial bladder present, ellipsoid, same position as in male, length = 0.07 mm, width = 0.04 mm (Fig. 4B). Terminalia with 1st valvula slightly differentiated from the 8th sternum; 8th sternum with posterior margin uniformly rounded (Figs. 6A, B). Distribution. Brazil: Pernambuco, archipelago of Fernando de Noronha (Fig. 1). Types condition. Holotype: Good, lacking right hind leg; both antennae missing their distal halves (Figs. 2A, B). The four female paratype specimens apparently subadult (5th instar) based on morphological characteristics as some depigmented areas appear reabsorbed such as in the paraproct. Intraspecific variation. Female paratypes exhibit lighter or darker shades of chestnut brown color, though never so dark as male. Biology. The holotype male of P. noronhensis sp. nov. is apterous, an unusual characteristic for male webspinners. The reduction or absence of wings is probably induced by the secretion of the juvenile hormone, which is one of the factors responsible for the neotenization (Iwanaga & Tojo 1986; Ross 2000). The amount and timing of secretion of juvenile hormone during nymphal instars influences directly the development of male wings: complete, when macropterous; partial, when subapterous or micropterous; or none, when apterous. All of these conditions can occur within the same species population. In arid environments, apterous males tend to remain inside the colony, mate with sisters, and consequently increase the probability of new males also being apterous. On the other hand, macropterous males living in more favorable environments can fly from colony to colony and perform random matings, thus decreasing the tendency to apterism (Ross 2000). For Pararhagadochir the trend towards apterism has been observed at the population level in P. trachelia Navás, 1915 (Szumik 2004)., Published as part of Costa-Pinto, Paula J., Olivier, Renan S. & Rafael, José A., 2021, The first species of Embioptera (Insecta) from the archipelago of Fernando de Noronha (Pernambuco: Brazil), pp. 142-150 in Zootaxa 4941 (1) on pages 143-147, DOI: 10.11646/zootaxa.4941.1.9, http://zenodo.org/record/4594870, {"references":["Rafael, J. A., Limeira-de-Oliveira, F., Hutchings, R. W., Miranda, G. F. G., Neto A. M. S., Somavilla, A., Camargo, A., Asenjo, A., Pinto, A. P., Bello, A. M., Dalmorra, C., Mello-Patiu, C. A., Carvalho, C. J. B., Takiya, D. M., Parizotto, D. R., Marques, D. W. A, Cavalheiro, D. O., Mendes, D. M. M., Zeppelini, D., Carneiro, E., Lima, E. F. B., Lima, E. C. A., Godoi, F. S. P., Pessoa, F. A. C., Vaz-de-Mello, F. Z., Sosa-Duque, F. J., Flores, H. F., Fernandes, I. O., Silva-Junior, J. O., Gomes, L. R. P., Monne, M. L., Castro, M. C. M, Silva, M. P. G., Couri, M. S., Gottschalk, M. S., Soares, M. M. M., Monne, M. A., Rafael, M. S., Casagrande, M. M., Mielke, O. H. H., Grossi, P. C., Pinto, P. J. C., Bartholomay, P. R., Sobral, R., Heleodoro, R. A., Machado, R. J. P., Correa, R. C., Hutchings, R. S. G., Ale-Rocha, R., Santos, S. D., Lima, S. P, Mahlmann, T., Silva, V. S. & Fernandes, D. R. R. (2020) Insect (Hexapoda) diversity in the oceanic archipelago of Fernando de Noronha, Brazil: updated taxonomic checklist and new records. Revista Brasileira de Entomologia, 64 (3), e 20200052. https: // doi. org / 10.1590 / 1806 - 9665 - RBENT- 2020 - 0052","Iwanaga, K. & Tojo, S. (1986) Effects of juvenile hormone and rearing density on wing dimorphism and o ˆ cyte development in the brown planthopper, Nilaparvata lugens. Journal of Insect Physiology, 32 (6), 585 - 590. https: // doi. org / 10.1016 / 0022 - 1910 (86) 90076 - 4","Ross, E. S. (2000) Embia: contributions to the biosystematics of the insect order Embiidina. Part 1. Origin, relationships and integumental anatomy of the insect order Embiidina. Occasional Papers of the California Academy of Sciences, 149, 1 - 53.","Szumik, C. A. (2004) Phylogenetic systematics of Archembiidae (Embiidina, Insecta). Systematic Entomology, 29 (2), 215 - 237. https: // doi. org / 10.1111 / j. 0307 - 6970.2004.00239. x"]}

Zoomorphology 40, 331 – 340 (2021). doi:10.1007/s00435-021-00526-4, The morphology of the antennal hearts in the head of Phasmatodea and Embioptera was investigated with particular reference to phylogenetically relevant key taxa. The antennal circulatory organs of all examined species have the same basic construction: they consist of antennal vessels that are connected to ampullae located in the head near the antenna base. The ampullae are pulsatile due to associated muscles, but the points of attachment differ between the species studied. All examined Phasmatodea species have a Musculus (M.) interampullaris which extends between the two ampullae plus a M. ampulloaorticus that runs from the ampullae to the anterior end of the aorta; upon contraction, all these muscles dilate the lumina of both ampullae at the same time. In Embioptera, only the australembiid Metoligotoma has an M. interampullaris. All other studied webspinners instead have a M. ampullofrontalis which extends between the ampullae and the frontal region of the head capsule; these species do not have M. ampulloaorticus. Outgroup comparison indicates that an antennal heart with a M. interampullaris is the plesiomorphic character state among Embioptera and the likely ground pattern of the taxon Eukinolabia. Antennal hearts with a M. ampullofrontalis represent a derived condition that occurs among insects only in some embiopterans. These findings help to further clarify the controversially discussed internal phylogeny of webspinners by supporting the view that Australembiidae are the sister group of the remaining Embioptera., Published by Springer, New York

Phasmatodea represents an order of hemimetabolous insects. This group includes species with extreme forms of masquerade crypsis, whereby they imitate twigs, bark, lichen, moss, and leaves. In this study, we sequenced and annotated three mitochondrial genomes (mitogenomes) from Phasmatodea. The lengths of the novel mitogenomes range from 14,162 bp to 15,879 bp. The gene content and organization correspond to those inferred for the ancestral insect. We conducted phylogenetic analyses together with the existing mitogenomes of polyneopterans and mayflies. In most cases, the Phasmatodea was non-monophyletic, with Embioptera and Zoraptera nested inside. The mitogenome sequences from Embioptera and Zoraptera suffered from high substitution rates and displayed very long branches in phylogenetic trees. The monophyletic Phasmatodea was recovered only when the analysis employed the site-heterogeneous CAT-GTR model in PhyloBayes and used the nucleotide dataset PCG_nt. The Euphasmatodea was well established by various data types and inference methods. In addition, the clade Heteropterygidae and the subfamilies Lonchodinae and Necrosciinae were strongly supported. The Australasian clade Lanceocercata was recovered across analyses. However, the Clitumninae was non-monophyletic.

Silks produced by webspinners (Order Embioptera) interact with water by transforming from fiber to film, which then becomes slippery and capable of shedding water. We chose to explore this mechanism by analyzing and comparing the silk protein transcripts of two species with overlapping distributions in Trinidad but from different taxonomic families. The transcript of one, Antipaluria urichi (Clothodidae), was partially characterized in 2009 providing a control for our methods to characterize a second species: Pararhagadochir trinitatis (Scelembiidae), a family that adds to the taxon sampling for this little known order of insects. Previous reports showed that embiopteran silk protein (dubbed Efibroin) consists of a protein core of repetitive motifs largely composed of glycine (Gly), serine (Ser), and alanine (Ala) and a highly conserved C-terminal region. Based on mRNA extracted from silk glands, Next Generation sequencing, and de novo assembly, P. trinitatis silk can be characterized by repetitive motifs of Gly-Ser followed periodically by Gly-Asparagine (Asn—an unusual amino acid for Efibroins) and by a lack of Ala which is otherwise common in Efibroins. The putative N-terminal domain, composed mostly of polar, charged and bulky amino acids, is ten amino acids long with cysteine in the 10th position—a feature likely related to stabilization of the silk fibers. The 29 amino acids of the C-terminus for P. trinitatis silk closely resemble that of other Efibroin sequences, which show 74% shared identity on average. Examination of hydropathicity of Efibroins of both P. trinitatis and An. urichi revealed that these proteins are largely hydrophilic despite having a thin lipid coating on each nano-fiber. We deduced that the hydrophilic quality differs for the two species: due to Ser and Asn for P. trinitatis silk and to previously undetected spacers in An. urichi silk. Spacers are known from some spider and silkworm silks but this is the first report of such for Embioptera. Analysis of hydropathicity revealed the largely hydrophilic quality of these silks and this feature likely explains why water causes the transformation from fiber to film. We compared spun silk to the transcript and detected not insignificant differences between the two measurements implying that as yet undetermined post-translational modifications of their silk may occur. In addition, we found evidence for codon bias in the nucleotides of the putative silk transcript for P. trinitatis, a feature also known for other embiopteran silk genes.

A new species of Pararhagadochir Davis, 1940 is described from the State of Tocantins, Brazil. Pararhagadochir castaneus sp. nov. differs from its congeners by relatively smaller size, elongated forewings, body uniformly chestnut-brown, submentum inflexed and strongly sclerotized, outer tip of 10Lp with both sclerotized and membranous areas and Cu vein unforked. An identification key to all species of Pararhagadochir and a short identification key to the Brazilian species are provided.

String sequence analysis revealed that silk spinning behavior of adult female Embioptera varies from species-specific to individualistic. This analysis included 26 species from ten taxonomic families with a total of 115 individuals. Spin-steps, 28 possible positions of the front feet during spinning, were scored from hour-long DVD recordings produced in the laboratory. Entire transcripts of hundreds to thousands of spin-steps per individual were compared by computing Levenshtein edit distances between all possible pairs of subsequences, with lengths ranging from 5 to 25—intraspecific similarity scores were then computed. Silk gallery characteristics and architecture, body size, climatic variables, and phylogenetic relationships were tested as possible drivers of intraspecific similarity in spinning behavior. Significant differences in intraspecific similarity aligned most strongly with climatic variables such that those species living in regions with high temperature seasonality, low annual precipitation, and high annual temperatures displayed more species-stereotypical spinning sequences than those from other regions, such as tropical forests. Phylogenetic signal was significant but weakly so, suggesting that environmental drivers play a stronger role in shaping the evolution of silk spinning. Body size also appears to play a role in that those of similar size are more like each other, even if not related.

Embioptera display the unique ability to spin silk with their front feet to create protective domiciles. Their body form is remarkably uniform throughout the order, perhaps because they all live within the tight confines of silken tubes. This study contributes to an understanding of the ecology of Embioptera, an order that is rarely studied in the field. We conducted a census to quantify the habitats of two species with overlapping distributions on the tropical island of Trinidad in a search for characteristics that might explain their distinct ecologies. One species, Antipaluria urichi (Saussure) (Embioptera: Clothodidae), lives in larger colonies with more expansive silk in habitats throughout the island, especially in the rainforest of the Northern Range Mountains. The other, Pararhagadochir trinitatis (Saussure) (Embioptera: Scelembiidae), was found only in lowland locations. We quantified silk-spinning behavior and productivity of the two species and found that A. urichi spins thicker silk sheets per individual and emphasizes spin-steps that function to create a domicile that is more expansive than that produced by P. trinitatis. Their silks also interact differently when exposed to water: the smaller-diameter silk fibers of P. trinitatis form more continuous films on the surface of the domicile after being wetted and dried than that seen in A. urichi silk. This tendency gives P. trinitatis silk a shiny appearance in the field compared to the more cloth-like silk of A. urichi. How these silks function in the field and if the differences are partially responsible for the distinct distributions of the two species remain to be determined.

A new genus and two new species of Embioptera are described from the upper Cretaceous of Burmese amber: Gnethoda odontophora sp. nov. is classified into Clothodidae Gnethoda due to its similar body characters and venation, but distinguished by the number of incisive teeth on left mandible and bladders on metabasitarsus from other species; Multivena curvivena gen. et sp. nov., is erected base on intensive long setae on antenna, MA forked, Cu not fork, multiple crossveins present between C and R, and assigned to Sorellembiidae. The new fossil species enrich the diversity of Embioptera in the Cretaceous. urn:lsid:zoobank.org:pub:4C5FD162-7E7E-42D4-A7BB-45561E012E9E.