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Trigoniophthalmus alternatus “Jumping bristletail” Figures 8 (lateral), 9 (dorsal, ventral) Plates 1 (lateral), 2 (dorsal), 3 (ventral) Trigoniophthalmus, as the representative of the most basal order in Class Insecta, is notable in its tracheal architecture by a complete lack of longitudinal connections between spiracles. This taxon is critical to understanding the apparent insect ground-plan tracheal structure. The thoracic tracheae of most taxa are supplied by both T 2- S and T 3- S, with T 2- DLT connecting longitudinally. In Trigoniophthalmus, however, T 3- S appears to only supply T 3- L and its associated coxa. Branches from T 2- S extend posteriorly into the metathorax, nominally supplied by T 3- S, including an apparent T 3- DLT. The naming of T 3- DLT here is an instance of using the positional criterion in homology (sensu Remane, 1952), in that the connectivity of this trachea is not consistent with other taxa; however the position suggests its identification as T 3- DLT (de Pinna, 1991). T 2-VB extends posterior, almost in the opposite direction of the anterior-reaching cephalic branches. While the position of T 2-VB may not appear to be “ventral” in nature, a comparison of its relative position in the hump-backed Trigoniophthalmus and in particular its connection to T 2- L with apterygote taxa from Zygentoma and Dermaptera demonstrates its homology as a ventral branch. Several tracheae range anteriorly or posteriorly beyond segment boundaries, most prominently A 5-DB-DVi, which extends posteriorly past A 6- S, reaching A 7- S; and A 7-Cr, which likewise extends posteriorly from A 7- S into both the cerci and terminal filament. However, tracheae for most segments, particularly in the abdomen, remain restricted to their individual segment, placing Trigoniophthalmus among the simplest tracheal body plans in this study and unique in its lack of longitudinal connections. This corroborates observations made by Palmén (1877) and reviewed by Dittrich and Wipfler (2021). DESCRIPTION: HEAD: Characteristically arched thorax, pronotum covering much of head, making boundary between thoracic tracheae and head tracheae rather indistinct. T 2- S at anteriormost margin of mesothorax, covered by overhanging tergum, shown in figure 10. H-DCT very thick, with three branches: H-Ic, extending anteriorly and dorsad; H-Oc-Ant anterior and medially before dividing into H-Oc toward midline and H-Ant laterally; H-Mx proceeds anteriad with H-MxPlp branching ventrally and anteriad. H-VCT very thick, with two branches: H-Lbm ventrad, extending into H-LbmPlp; H-Md anteriad, crossing over H-Md from opposite side of head but not connecting. THORAX: T2-S present, with five tracheae: H-DCT and H-VCT leading anteriad directly into head, beginning at T2-S and running parallel for length of prothorax; T2-DB leading dorsad; T2-VB and T2-L directly posteriad. H-DCT with no thoracic branches; H-VCT with T1-L branching ventrally; T1-VC present, extending from T1-L. T2-DB-Vi branches just dorsal of T2-S, extending anteriorly; T2-DB continuing dorsad, dividing into what appear to be anterior T1-DLT and posterior T2-DLT near midline; T1-DLT and T2-DLT extend into viscera with no connections to neighboring spiracles. T2-L running posteriad and ventrally, into midleg; large T2-L-Vi and smaller T2-VC divide from T2-L posterior from T2-S. T2-VC joins with anterior-arching T3-VC to form T2-VX intersection. T2-L-Vi extends posteriad into metathorax, with apparent dorsal T3-DLT, T3-Cx ventral, and numerous small visceral tracheae. T3-S ventral to and much smaller than T2-S, with two tracheae: T3-L and T3-VB. T3-L extending dorsad with small T3-L-Vi before arcing ventrad into hind leg; T3-VB directly toward midline with small T3-VC branch that arcs anteriorly to join T2-VC at T2-VX. ABDOMEN: A1..7-S present, all located ventrally. No longitudinal connections present; figure 11 with representative abdominal segment. A n -S with A n -DB running dorsad along arc of body wall, turning inward toward midline of body but not forming DC; A n -VB branching dorsally for a short distance before arcing ventrad toward midline. A n -VC absent. A n -DB with visceral tracheae A n -DB-MVi, branching anterior and medially halfway up body; A n -DB-DVi branches dorsally toward tergal wall. A5-DB-DVi extends posteriorly to 8th abdominal segment; A7-DB-DVi extends posteriorly past 8th abdominal segment, dividing into A-TF and A-Cr; neither A 5-DB-DVi nor A 7-DB-DVi connect to spiracles of other segments. A-TF with two tracheae per side (4 total); A-Cr single trachea per cercus., 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":["Remane, A. 1952. Die Grundlagen des naturlichen Systems, der vergleichenden Anatomie und der Phylogenetik. Leipzig: Geest & Portig.","de Pinna, M. C. C. 1991. Concepts and tests of homology in the cladistic paradigm. Cladistics 7: 367 - 394.","Palmen, J. A. 1877. Zur Morphologie des Tracheensystems. Leipzig: Wilhelm Engelmann.","Dittrich, K., and B. Wipfler. 2021. A review of the hexapod tracheal system with a focus on the apterygote groups. Arthropod Structure and Development 63: 101072."]}

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

Kaplin, V.G. (2021): A new species of the bristletail genus Petrobius Leach, 1809 (Microcoryphia: Machilidae) from Crimea. Far Eastern Entomologist 426: 22-28, DOI: 10.25221/fee.426.4, URL: http://dx.doi.org/10.25221/fee.426.4

Eight new species of the genus Coryphophthalmus are described from Abkhazia, Krasnodar Territory, and North Ossetia: Coryphophthalmus prosvirovisp. n.,C. brunioculussp. n.,C. messazhayisp. n.,C. silvestrissp. n.,C. lapidicolasp. n.,C. alanicussp. n.,C. viridioculussp. n., and C. bicolorioculussp. n. A review of the distribution, morphological characters, and a key to species of the genus Coryphophthalmus are given.

The paper continues the exploration of the morphological and functional diversity of male genitalia in Archaeognatha by the study of an undescribed Charimachilis species. Exoskeletal details are documented by drawings and SEM micrographs. The lack of tubular setae on gonapophyses and penis suggests sperm transfer by direct male-female contact (not via sperm deposited on a thread), while genitalic morphology is not suggestive of the male clasping the female. For a broader evolutionary interpretation of archaeognathan male genitalic characters and mating behaviour, we revisited the issue of the “paleoforms”: Charimachilis, Ditrigoniophthalmus, Mesomachilis, and Turquimachilis had been categorized as such, i.e. as the putatively basalmost offshoots of crown-group Archaeognatha (Machiloidea = Meinertellidae + potentially non-monophyletic Machilidae). A recent cladistic analysis, however, had found all four genera subordinate in a monophyletic Machilidae. After revision of characters and addition of zygentoman outgroup taxa, we re-analysed this dataset, finding that except for the monophyly of Meinertellidae the dataset provides no convincing resolution of machiloid relationships. The status of the paleoforms is thus unclear with currently available data. Yet, based on comparison with zygentomans, mating with use of a sperm-thread appears plesiomorphic for Archaeognatha. We also briefly discuss whether the distribution of elements of the vestiture (e.g. scales, setae, and non-articulated trichoid sensilla) can support our previously published tentative hypothesis of the archaeognathan (and insect) penis being formed by sternal, coxal, and gonapophyseal elements of abdominal segment 10.

Kaplin, V. G., Kiseleva, L. V., Kozhevnikova, O. P. (2020): NEW SPECIES OF BRISTRLETAILS OF THE GENUS CORYPHOPHTHALMUS VERHOEFF, 1910 (ARCHAEOGNATHA: MACHILIDAE) FROM NORTH AND SOUTH OSSETIA. Far Eastern Entomologist 406: 1-13, DOI: 10.25221/fee.406.1, URL: http://dx.doi.org/10.25221/fee.406.1

Petrobiellus sachalinensis sp. nov. from the northwest of Sakhalin Island (Russia) is described and illustrated. It is compared with the three other known species of the genus, P. takunagae Silvestri, 1943 from Honshu Island (Japan), P. curvistylis Uchida, 1954 from Hachijo-jima Island (Japan), and P. kusakini Kaplin, 1980 from Simushir Island (Russia). The new species can be distinguished from the congeners by the colour of body and scales, by distribution of pigment, by colour and shape of paired ocelli, and by structure of compound eyes, maxillary palps, legs, urites, and ovipositor.

Genus Charimachilis Wygodzinsky, 1939 Type species: Praemachilis orientalis Silvestri, 1908., Published as part of Kaplin, V. G. & Martynov, V. V., 2022, A new species of the bristletail genus Charimachilis Wygodzinsky, 1939 (Microcorypha: Machilidae) from Eastern Ukraine, pp. 1-8 in Far Eastern Entomologist 449 on page 2, DOI: 10.25221/fee.449.1, http://zenodo.org/record/7166840

Kaplin, V.G., Martynov, V.V. (2022): A new species of the bristletail genus Charimachilis Wygodzinsky, 1939 (Microcorypha: Machilidae) from Eastern Ukraine. Far Eastern Entomologist 449: 1-8, DOI: 10.25221/fee.449.1, URL: http://dx.doi.org/10.25221/fee.449.1

Genus Silvestrichiloides Mendes, 1990 Silvestrichiloides Mendes, 1990: 91���92. Type species: Silvestrichilis beckeri Paclt, 1960., Published as part of Kaplin, Vladimir & Shakula, Georgiy, 2021, New species of bristletails of the family Machilidae (Archaeognatha) from Kazakhstan, pp. 435-445 in Acta Entomologica Musei Nationalis Pragae 61 (2) on page 436, DOI: 10.37520/aemnp.2021.024, http://zenodo.org/record/5821053, {"references":["MENDES L. F. 1990: An annotated list of generic and specific names of Machilidae (Microcoryphia, Insecta) with identification keys for the genera and geographical notes. Estudos, Ensaios e Documentos, Instituto de Investigacao Cientifica Tropical, Lisboa 155: 1 - 127.","PACLT J. 1960: Felsenspringer (Ins. Thysanura) des Senckenberg-Museums. Senckenbergiana Biologica 41 (5 / 6): 325 - 332."]}

The fauna of bristletails of the family Machilidae in Kazakhstan currently includes one species of the genus Silvestrichiloides Mendes, 1990 and 13 species of the genus Allopsontus Silvestri, 1911. The present study describes one new species of the genus Silvestrichiloides (S. berkarensis Kaplin, sp. nov. from South Kazakhstan) and two new species of the genus Allopsontus (A. (Kaplinilis) nigrostriatus Kaplin, sp. nov. and A. (Machilanus) perfectus Kaplin, sp. nov. from Southeastern Kazakhstan). Silvestrichiloides berkarensis sp. nov. differs from the other species of this genus in the structure of antennal flagellum, apical palpomere of labial palp and ovipositor. Among species of the subgenus Kaplinilis Mendes, 1990, A. nigrostriatus sp. nov. belongs to a group of species characterized by numerous short chaetae on the ventral surface of the 5–7th palpomeres of the male maxillary palp and by the absence on the labial palp. This group includes two species: A. volgensis Kaplin, 1999 from Samara Region and A. smelyanskii Kaplin, 1999 from Orenbourg Region (both Russia). The new species differs from A. volgensis and A. smelyanskii in the length of the body and antenna, color of scales on the upper surface of the body, shape of the compound eye and paired ocellus, structure of the flagellum and apical palpomere of the male labial palp. The subgenus Machilanus Silvestri, 1934 is represented only by A. bitschi Wygodzinsky, 1962 from Afghanistan and A. perfectus sp. nov., which are characterized by numerous short chaetae on the ventral surface of the 2nd–7th palpomeres of the male maxillary palp. Allopsontus perfectus sp. nov. differs from A. bitschi in the shape of compound eyes, paired ocellus, structure of male labial palp and genitalia.

The order Microcoryphia, commonly known as bristletails, is considered as the most primitive one among living insects. Within this order, two species, Coreamachilis coreanus and C. songi (Machilidae: Machilinae), display the following contrasting reproductive strategies: parthenogenesis occurs in C. coreanus, whereas sexual reproduction is found in C. songi. In the present study, the complete mitogenomes of C. coreanus and C. songi were sequenced to compare their mitogenome structure, analyze relationships within the Microcoryphia, and assess adaptive evolution. The length of the mitogenomes of C. coreanus and C. songi were 15,578 bp and 15,570 bp, respectively, and the gene orders were those of typical insects. A long hairpin structure was found between the ND1 and 16S rRNA genes of both species that seem to be characteristic of Machilinae and Petrobiinae species. Phylogenetic assessment of Coreamachilis was conducted using BI and ML analyses with concatenated nucleotide sequences of the 13 protein-coding genes. The results showed that the monophyly of Machilidae, Machilinae, and Petrobiinae was not supported. The genus Coreamachilis (C. coreanus and C. songi) was a sister clade to Allopsontus helanensis, and then the clade of ((C. coreanus + C. songi) + A. helanensis) was a sister clade to A. baii, which suggests that the monophyly of Allopsontus was not supported. Positive selection analysis of the 13 protein-coding genes failed to reveal any positive selection in C. coreanus or C. songi. The long hairpin structures found in Machilinae and Petrobiinae were highly consistent with the phylogenetic results and could potentially be used as an additional molecular characteristic to further discuss relationships within the Microcoryphia.

The complete mitochondrial genome of Pedetontus zhejiangensis (Microcoryphia: Machilidae) was successfully sequenced. The mitochondrial genome of P. zhejiangensis was a circular molecule of 15,602 bp in length, containing 13 protein-coding genes, 2 rRNA genes, 22 tRNA genes, and the control region, which showed the typical insect mitochondrial genome arrangement. The AT content of the whole genome was 73.8% and the length of the control region was 671 bp with 82.5% AT content. In BI and ML phylogenetic trees, P. zhejiangensis was a sister group to Pedetontus silvestrii, and the monophyly of Pedetontus was strongly supported. The genus Pedetontinus was a sister group to Pedetontus.

Two new bristletail species of the family Machilidae are described: Allopsontus tyrnyauzi sp. n. from the Caucasus and A. zinchenkoi sp. n. from the Pamir-Alai Mountain system. The former species belongs to the subgenus Kaplinilis Mendes, 1990. Among the species described within this subgenus, numerous short, pigmented, appressed setulae on the underside of the 5-7th maxillary palpomeres and the 3rd labial palpomere in males are found only in A. tyrnyauzi sp. n. and A. bifarius (Wygodzinsky, 1970), the latter known from western Mongolia and Tuva. Allopsontus tyrnyauzi sp. n. differs from A. bifarius in the relatively short eye contact line, more dilated paired ocelli, urosternites with a right apical angle, the narrower sensory fields on the fore femora in males, and fewer paramere segments. Allopsontus zinchenkoi sp. n. belongs to the subgenus Anisopsontus Mendes, 1990 and differs from the other consubgeners in the color of the compound eyes, the width to length ratio of the paired ocelli, longer abdominal styli, larger sensory fields on the fore femora in males, and the relatively short ovipositor.

A review of phylogenetic relationships of species of the genus Charimachilis based on comparative analysis of zoogeographic distribution and plesiomorphic and apomorphic states of their main morphological features is performed for the first time. The II and III instar larvae of Charimachilis caucasica are described. The genus Charimachilis belongs to the subfamily Machilinae (Machilidae) where, together with the genus Turkimachilis, it forms the Charimachilis group of genera occupying an isolated position within Machilidae close to the subfamily Petrobiinae. The genus Charimachilis probably originated in the mountainous areas of the east and northeast Mediterranean coasts in the zone of evergreen forests and shrubs. The patterns of dispersal of its species deep into the European continent along the south and west Black Sea coast, as far northwards as 50°N, are traced. Reduction of body size and transition to parthenogenesis in the genus Charimachilis were probably determined by the increasing aridity of climate and deterioration of living conditions; bisexual species remained only in the humid environments of relict communities in the Caucasus. The new species Charimachilis morozovi sp. n. most closely resembles Ch. palaestinensis in the absence of lateral digging teeth on the anterior gonapophyses but differs from the latter in the structure of the ovipositor and urosternites.

Genus Turquimachilis Bach de Roca, Fanciulli, Cicconardi, Molero-Baltanás et Gaju-Ricart, 2013 Type species: Turquimachilis mendesi Bach de Roca, Fanciulli, Cicconardi, Molero-Baltanás et Gaju-Ricart, 2013, Published as part of Kaplin, V. G., 2021, A new species of bristletails of the genus Turquimachilis (Microcoryphia: Machilidae) from the Crimea, pp. 40-45 in Zoosystematica Rossica (Zoosyst. Rossica) (Zoosyst. Rossica) 30 (1) on page 41, DOI: 10.31610/zsr/2021.30.1.40

Turquimachilis taurica sp. nov. (Figs 1–14) Holotype. Female (slide-mounted); Republic of Crimea, Grand Canyon of Crimea, 44°31 ′ 40 ″ N, 34°01 ′ 00 ″ E, 500–600 m, under stones, 7.IX.2020, V. Kaplin leg. Description. Body length: 9.6 mm; body width: 2.0 mm; length of antenna: about 7.5 mm; length of cerci: 3.4 mm; total width of eyes: 1.00 mm; length of eye: 0.44 mm; width of paired ocellus: 0.36 mm; length of paired ocellus: about 0.13 mm; length of coxal styli of legs: about 0.4 mm; length of ovipositor: 1.45 mm. General body colour (in ethanol) light brownish, with dark brown scales on upper and lower sides of body. Frons, clypeus, occiput, mandibles, maxillae, maxillary and labial palps, and legs with reddish brown or reddish violet pigment. Head, maxillary palps and coxae most pigmented. Scapus, pedicellus and flagellum of antennae without pigment. Antennae shorter than body. Distal chains of flagellum divided into 9–11 annuli (Fig. 1). Cercus approximately 0.36 times the body length. Apex of cercus (Fig. 2) with two lateral spikes. Articles of cerci, except for apical two, with colourless supporting macrochaetae on inner side. Compound eyes dark brown (in ethanol). Ratio of compound eye length to its width 0.89; ratio of contact line length to eye length 0.61 (Fig. 3). Paired ocelli black with narrow white rim, shoeshaped, subinferior to compound eyes. Frons slightly swollen between paired ocelli. Distance between inner margins of ocelli about 0.21 times and between their outer margins 0.92 times the total width of compound eyes. Maxillary palp (Fig. 4): apical palpomere 0.84 times as long as preceding one; ratio of lengths of 5th and 4th palpomeres about 1.64; dorsal surface of 7th, 6th and 5th palpomeres with 17–18, 16–18 and 5–6 hyaline spines, respectively. Labial palp (Fig. 5): apical palpomere triangular oval, 2.1–2.2 times as long as wide, with about 21–25 sensorial cones. Mandible with four well-developed distal teeth (Fig. 6). Fore and middle femur and tibia widened (Fig. 7). Fore and hind legs 1.09 and 1.16 times as long as middle legs, respectively. Ratios of length to width of femur, tibia and tarsus as in Table 1. Ratio of length of 3rd tarsomere of hind tarsus to its total length about 0.39. Ventral surfaces of coxae, femora, tibiae and tarsi without hyaline spine-like chaetae. Ratio of coxal stylus length to width of middle and hind coxa about 1.7 (Fig. 8). Urocoxites I and VI–VII with 1 + 1 eversible vesicles; urocoxites II–V with 2 + 2 eversible vesicles. Posterior angle of urosternites II–VI approximately 82–86°, that of urosternite VII about 74°. Ratios of lengths of urosternites and urocoxites II–VI 0.67–0.69. Ratios of lengths of urostyli (without apical spine) and urocoxites II–VII 0.50–0.54. Ratio of length of urostylus and urocoxite VIII 0.68, IX 0.45. Ratios of lengths of apical spines and urostyli II–VIII 0.55–0.62 (Figs 9–11). Urocoxite VII (Fig. 10) with well-developed lobes protruding between styli; ratio of length to width of one lobe about 0.75. Thoracic tergites, urosternites, urocoxites I–VIII, and urotergites I–VI and X without macrochaetae. Urocoxites IX (Fig. 12) with 3–5 inner sublateral spines (spiniform chaetae), without outer sublateral spines. Urotergites VII with 1 + 1, urotergites VIII and IX with 2 + 2 sublateral spines. Ovipositor weakly sclerotised, thickened, covered by urocoxites IX, stout, similar to that of T. mendesi. Gonapophysis VIII (Fig. 13) with 17 divisions; its apex with two small spine-like lobules and a subterminal chaeta between them; chaeta being a little longer than apical division; apical division with about 15 sensory spines and 2–3 relatively long thin chaetae in preapical part and with two curved chaetae in basal part. Remaining divisions of gonapophysis VIII with a row of setae, some of which, mainly those on inner and outer sides of gonapophysis, long and ciliary. Basal division without chaetae. Gonapophysis VIII without digging teeth. Gonapophysis IX (Fig. 14) with 19 divisions, a somewhat curved horn at apex and a subterminal chaeta being slightly longer than apical division. Apical division also with a group of 11–12 tiny sensory spines; these spines present also on preceding 5–6 divisions, but number of spines decreasing towards base. Each of these divisions also with about 2–4 chaetae; chaetae on outer side longer. Basal division without chaetae. Comparison. According to the structure of the female gonapophyses and 2 + 2 eversible vesicles on urocoxites II–V, the new species belongs to the genus Turquimachilis (Bach de Roca et al., 2013), which was monotypic, with the only species T. mendesi. The main morphological differences between T. taurica sp.nov. and T. mendesi are given in Table 2. The new species differs from T. mendesi in the structure of the compound eyes, maxillary palp, urocoxites, and the number of divisions of gonapophyses. The ratio of lengths of contact line and compound eye in female of T. taurica sp. nov. is about 0.6, while in female of T. mendesi it is about 0.5. The numbers of hyaline spines on the dorsal surface of the 6th and 7th palpomeres of maxillary palps are 16–18 and 9–12, respectively. The ratios of length to width of hind tibia are about 2.8 and about 2.6, respectively; the ratios of lengths of urostyli and urocoxites II–V are 0.50–0.52 and 0.38–0.43; the number of divisions of gonapophyses are 17–19 and 15–16, respectively. The urocoxites of T. mendesi have no spiniform chaetae, whereas urocoxites IX of T. taurica sp. nov. bear 3–5 inner sublateral spines. Etymology. The species name is a Latin adjective referring to Chersonesus Taurica, an ancient name of the Crimea, where the holotype was collected. Habitat. The holotype of T. taurica sp. nov. was collected in a forest (Carpinus, Fagus, Fraxinus excelsior, Quercus robur) among the stones., Published as part of Kaplin, V. G., 2021, A new species of bristletails of the genus Turquimachilis (Microcoryphia: Machilidae) from the Crimea, pp. 40-45 in Zoosystematica Rossica (Zoosyst. Rossica) (Zoosyst. Rossica) 30 (1) on pages 41-45, DOI: 10.31610/zsr/2021.30.1.40

Coryphophthalmus troglophilus Kaplin, sp. nov. Figs. 2–14 Diagnosis. Coryphophthalmus troglophilus sp. nov., with 2 + 2 eversible vesicles on urocoxites II–IV, belongs to the subgenus Coryphophthalmus s. str., a group of species with long thin chaetae on the legs, clypeus and maxillary palpomeres 2‒4, but lacking on male labial palp; without needle-like macrochaetae on legs. Only one species, C. abchasicus (Kaplin, 2017) is affiliated to this group other than the new species (Kaplin, 2017). Description. Body length of males 7.0– 7.3 mm, females 7.2–8.2 mm; width of males and females 1.8–2.1 mm; antennae length of males 8.5–8.7 mm, female 6.8 mm (slightly broken); cercus length of males 3.8 mm, females 3.6–4.2 mm; total eyes width of males 0.86–0.88 mm, females 0.86–1.00 mm; eye length of males and females 0.43–0.46 and 0.44–0.48 mm, respectively; paired ocelli width of males 0.26–0.27 mm, females 0.23–0.26 mm; paired ocellus length of males and females 0.15–0.16 and 0.13–0.14 mm, respectively; coxal styli length of males 0.65–0.70, females 0.60–0.65 mm; ovipositor length 2.3–2.6 mm. General body color (in ethanol) whitish or light yellow, almost without hypodermal pigment. Antennal base, frons, gena, lateral sides of clypeus, basal parts of mandibles, thoracic sterna with brown-violet hypodermal pigment of light to medium intensity. Scale color on surface of body grey, dark grey, almost white and black, light brown or brown, spotted on the upper side of body (Fig. 2). Antennae of males and females slightly longer than body. Distal chains of male and female flagella divided into 8–12 annuli. Аpical, 3rd, 5th and 7th annuli of distal chains with one or two basiconic sensilla of D form; each annulus of chains also with one “rosette-shaped” sensillum. Clypeus and labrum of male with long thin chaetae. Cerci of males and females about 0.50–0.55 times body length, divided into 22–23 annuli, with apically bifurcated spikes. Two distal annuli of cerci without lateral hyaline spines. Remaining cercal annuli with 1–5 inner lateral spines in both sexes. Compound eyes round, green or dark green (in ethanol). Length to width ratio of compound eye 1.00– 1.03 in both sexes; ratio of line of contact to length of male and female eyes 0.50–0.53 and 0.42–0.46, respectively. Paired ocelli submedian, pyriform, dark brown with narrow white borders. Ratios of width to length of male and female ocelli 1.7 and 1.8, respectively. Ratio of distance between inner and outer margins of ocelli to total width of compound eyes 0.16–0.18 and 0.64–0.66 eye in both sexes, respectively (Fig. 3). Apical maxillary palpomere 0.55–0.65 times (male) and 0.72–0.75 times (female) that of the preceding one, ratio of length of 5th and 4th palpomeres 1.3 in male and 1.4 in female. Dorsal surface of 7th, 6th and 5th palpomeres of maxillary palp with 10–11, 13–14 and 3–4 hyaline spines in male, and 12–14, 14–16, and 4–7 spines in female, respectively. Underside of palpomeres 2–4 of male maxillary palp with relatively numerous and long thin chaetae, missing on the dorsal surface of male labial palpomeres (Fig. 4). Apical labial palpomere triangularly oval, 2.3–2.5 times (male) and 2.6–2.7 times (female) longer than wide, with 30–35 sensorial cones (Fig. 5). Mandibles with three teeth in female and four in male (Figs. 6, 7). Fore and middle femora and tibiae of female widened. Hind tarsus 1.2–1.3 times longer than fore and middle tarsi in both sexes. Fore tibia longer than middle tibia 1.3 times in female and 1.2 times in male. Hind tibia longer than middle tibia, 1.5 times in female and 1.3 times in male. Fore femur of male without sensory field. Ratios of length to width of femur, tibia and tarsus given in Table 1. Ratio of length of apical hind tarsomere to total length of hind tarsus 0.32 in male and 0.35 in female (Fig. 8). Trochanter and femur of male with long, thin chaetae. Ventral surface of femora, tibiae and tarsi without spine-like macrochaetae in both sexes. All tibiae with 1–2 lateral, relatively long, colorless, thickened macrochaetae. Pretarsa with two well-developed claws and arolium with flexible pad between them (Fig. 9). Urocoxites I and V–VII with 1 + 1, II–IV with 2 + 2 eversible vesicles. Posterior angle of urosternites II–VI 74–80°, VII about 85° in both sexes. Length ratios of urosternites, urocoxites and urostyli given in Table 2. Inner posterior lobes of female urocoxites VII protruding. Ratio of length to total width of protruding lobes about 0.34. Thoracic tergites, urotergites I–IV, all urosternites, urocoxites I–V without macrochaetae in both sexes. Distribution of sublateral spines on urocoxites and urotergites given in Table 3. Urocoxites IX with 2–3 + 2–3 outer spines in both sexes and with 6–8 + 6–8 inner sublateral spines in male and 7–11 + 7–11 ones in female (Figs 10, 13). Ovipositor slender, elongate, visibly surpassing apex of styli IX (Fig. 10). Anterior and posterior gonapophyses with approximately 38–40 or 41–43 divisions, respectively. One or two basal divisions of anterior gonapophyses and 20–21 basal divisions of posterior gonapophyses glabrous. Distal spines of gonapophyses as long as 3–4 apical divisions combined. Apical divisions of anterior and posterior gonapophyses with 8–9 and 5–8 chaetae, respectively (not including apical spines) (Figs 11, 12). Male genitalia with one pair of parameres on urite IX. Parameres with 1 + 5 divisions (Fig. 12), surpassing apex of penis. Penis and parameres clearly not attaining level of apex of urocoxites IX, ratio of the distance between apexes of penis and of urocoxites IX to width of distal division of penis 5.0–6.5. Basal division of penis 1.3–1.4 times longer than distal division (Fig. 14). Material examined. Holotype: male (on slide) (ZIN); Abkhazia, Ochamchira District, Otap, Aymara, W Caucasus, 42°56′N, 41°31′E, elev. 440 m, Kotsha Cave, ˗20 to ˗ 25 m depth, on wet stoneflows, 20.09.2018 (R. S. Vargovitsh). Paratypes: 2 males, 7 females (one male and two females on slides); same locality and data (R. S. Vargovitsh) (ZIN). Habitats. All specimens of Coryphophthalmus troglophilus sp. nov. were collected on walls, stalagmites and among stones in a cave at depths of ˗20 to ˗ 25 m. The cave is narrow, almost all vertical, with a total depth of ˗ 33 m (Fig.14). The arthropod fauna in the cave is quite rich. The main adaptations of this new species to life in conditions of low light on the rocky surface substrate include green or dark green eyes; weak body pigmentation; pretarsa with well-developed arolium and claws; relatively long cerci, tarsi and tibiae of the hind legs; apical needles of the urostyli; large sublateral spines on urocoxites IX, especially in the male; reduced number of apical sensorial cones on the third labial palpomere and sensilla on the apical part of labium; and camouflaged by the mottled color of scales on the upper side of the body. These characteristics place it as a troglophile. Etymology. The species name troglophilus is Latin for “troglo = cave; philus = preference”. Discussion. Coryphophthalmus troglophilus sp. nov. resembles C. abchasicus in the presence of numerous long thin chaetae on the undersurface of the male maxillary palp, absence of such chaetae on the upper surface of the male labial palp and absence of spine-like chaetae on the undersurface of the tarsus. The main differences between C. troglophilus and C. abchasicus consist in the color of compound eyes, the line of eye contact, lengths of cercus and apical spines of the urostyli, ratio of length to width of the apical palpomere of the labial palp and number of divisions of the ovipositor and parameres (Table 4)., Published as part of Kaplin, Vladimir & Vargovitsh, Robert S., 2020, New species of bristletails of the family Machilidae (Microcoryphia) from caves in Abkhazia and Ukraine, pp. 530-540 in Zootaxa 4885 (4) on pages 532-536, DOI: 10.11646/zootaxa.4885.4.4, http://zenodo.org/record/4296928, {"references":["Kaplin, V. G. (2017) New species of the bristletail family Machilidae (Insecta, Microcoryphia) from Abkhazia. Entomological Review, 97 (2), 207 - 229. https: // doi. org / 10.1134 / S 0013873817020075"]}

Trigoniophthalmus ukrainensis Kaplin, sp. nov. Figs. 15–21 Diagnosis. Trigoniophthalmus ukrainensis sp. nov., with 2 + 2 eversible vesicles on urocoxites II–V, belongs to the subgenus Trigoniophthalmus s. str., which has been composed of only one species, T. alternatus (Silvestri, 1904). The two species differ in the number of annuli in distal chains of flagellum, and the absence of numerous long, thin chaetae on the undersurface of the male maxillary palp and the dorsal surface of the labial palp. Description. Holotype, male: body length 7.7 mm; body width 1.8 mm; antenna length 4.0 mm (broken); cercal length 3.0 mm. Coxal styli length 0.65–0.68 mm. General body color (in ethanol) yellowish light brown, with brown-violet hypodermal pigment of light to medium intensity on antennal base, frons, gena, lateral sides of clypeus, mandibles, maxillae, hypopharynx, first palpomere of maxillary and labial palpi, labium, labrum and thoracic sterna. Scale color on surface of body light brown or brown. Antennae shorter than body (slightly broken). Distal chains of flagellum divided into 9–12 annuli (two apical chains broken). Clypeus and labrum with long thin chaetae. Cercal length about 0.4 times as long as body length, with two apical spikes, divided into 17 or 18 annuli. Each article of cercus with 2 or 4 rows of scales. Annuli of cerci, except for apical two, with 1 or 2 inner lateral hyaline spines. Compound eyes slightly expanded, black (in ethanol). Length to width ratio of compound eye 0.9; ratio of line of contact to length of eye 0.50. Paired ocelli submedian, subtriangular, dark brown, 1.5 times wider than long. Ratio of distance between inner and outer margins of ocelli to total width of compound eyes 0.10 and 0.51, respectively (Fig. 15). Apical palpomere of maxillary palp 0.86 times as long as preceding one; ratio of length of fifth to fourth palpomere 1.3. Dorsal surface of 7th, 6th and 5th palpomeres of maxillary palp with 11, 9 and 3 hyaline spines, respectively (Fig. 16). Undersurface of maxillary palpomeres and dorsal surface of labial palpomeres without numerous long thin chaetae. Apical labial palpomeres triangular-oval, 2.0 times longer than wide (Fig. 17). Mandibles with three distal teeth (Fig. 18). Fore and middle femora and tibiae widened (Fig. 19). Hind legs longer than forelegs, much longer than middle legs. Hind tarsus 1.15 and 1.25 times longer, respectively, than those of fore and middle legs. Hind tibia 1.17 and 1.34 times longer, respectively, than those of fore and middle legs. Fore femur with narrow sensorial field on the external distal part consisting of 4–5 groups of very small “rosette-shaped” sensilla (Figs. 19, 20), each group containing about 12–16 sensilla. Ratios of length to width of femur, tibia and tarsus of middle leg shorter and relatively wider than those of fore and hind leg (Table 5). Ratio of length of apical hind tarsomere to total length of hind tarsus 0.37. Trochanter, femur and tibia lacking long, thin chaetae. Ventral surface of femora, tibiae and tarsi without pigmented, spine-like macrochaetae. Pretarsi with arolium. Middle and hind legs with coxal styli. Ratio of length of styli to width of middle and hind coxae 1.45 and 1.77, respectively. Urocoxites I, VI and VII with 1+1, II–V with 2 + 2 eversible vesicles. Posterior angle of urosternites II–VI 78–81°, VII 86°, VIII 110°. Length ratios of urosternites, urocoxites and urostyli around 0.5 (Table 6). Thoracic tergites, urotergites I–IV, urosternites, urocoxites I–VIII without spines. Distribution of sublateral spines on urotergites V–X and urocoxites IX with sublateral spines (Table 6). Urocoxites IX with 3 + 3 inner sublateral spines, missing on their outer sides (Fig. 21). Male genitalia with one pair of parameres on urite IX. Parameres with 1 + 5 divisions (Fig. 21), surpassing apex of penis. Penis and parameres completely covered by coxites IX. Material examined. Holotype: male (on slide) (ZIN); Ukraine, Zakarpatska Oblast, Tyachiv District, near Pidplesha, E Carpathians, Krasna Mountain Range, 48°9’25”N, 23°49’1″E, elev. 698 m, Tsherlenyi Kamin Cave System, Kanyon Cave, ~ 5 m from the entrance, 23.01.2000 (R. S. Vargovitsh). Habitats. Trigoniophthalmus ukrainensis sp. nov. was collected on the wall ~ 5 m from the entrance. The present new species exhibits no morphological adaptations to life in caves. Etymology. The new species is named after Ukraine, where it was collected., Published as part of Kaplin, Vladimir & Vargovitsh, Robert S., 2020, New species of bristletails of the family Machilidae (Microcoryphia) from caves in Abkhazia and Ukraine, pp. 530-540 in Zootaxa 4885 (4) on pages 537-538, DOI: 10.11646/zootaxa.4885.4.4, http://zenodo.org/record/4296928

Two new species of bristletails of the family Machilidae are described from caves in the Western Caucasus (troglophile Coryphophthalmus troglophilus sp. nov.) and in the Eastern Carpathians (trogloxene Trigoniophthalmus ukrainensis sp. nov.). The main morphological adaptations of C. troglophilus sp. nov. to life in caves include green eye color; weak body pigmentation; well-developed arolium and claws; relatively long cerci, tarsae and tibiae of the hind legs; long apical needles of the urostyli; large sublateral spines on urocoxites IX. C. troglophilus sp. nov. resembles C. abchasicus (Kaplin, 2017), but they are distinguishable in color of eyes, ratio of eye contact to length, length of cerci, structures of the labial palps, legs, urocoxites IX, ovipositor, and parameres. Trigoniophthalmus ukrainensis sp. nov. has no recognized morphological adaptations to life in caves. This species resembles T. alternatus (Silvestri, 1910), but they are easily distinguished in the number of annuli in distal chains of flagellum, ratio of width to length of paired ocelli, chaetotaxy of male maxillary and labial palps.

Charimachilis rostoviensis Kaplin, sp. nov. (Figs 6, 7) Type material. HƟΓƟΤΥΡΕ: ♀ (slide-mounted, ZIN), RUSSIA: Rostov-on-Don, Botanical garden of the Southern Federal University, 47°14′13″N, 39°39′12″E, 50 m a.s.l., under a dead pine trunk, June 19, 2019, V. Martynov leg. PΑ*©ΑΤΥΡΕඌ: ♀♀ 3 (one on slide), 6 juv., the same locality, V. Martynov leg. (ZIN). Description. Female. Body length 9.6–11.0 mm; body width 2.8–3.1 mm; antennal length 5.8–6.8 mm; cercal length 4.1–4.4 mm; total eyes width 1.00– 1.03 mm, eye length 0.42–0.44 mm; paired ocelli width 0.50–0.51 mm, length 0.18–0.19 mm. Coxal styli length 0.62–0.65 mm. Ovipositor length 1.4–1.6 mm. General body color whitish, with brown hypodermal pigment of faint or medium intensity only on antennal base, frons, gena, lateral sides of clypeus, mandible, galea of maxilla. Frons between eyes convex. Color of scales on upper and lower surface of body brown. Antennae shorter than body. Distal chains of flagellum divided into 8–12 annuli. Cercus 0.42–0.46 times as long as body length, with about 21 divisions. Apex of cercus with one large lateral spike (Fig. 6A). Divisions of cerci, except for apical three, with 1–3 colorless supporting macrochaetae on inner side. Compound eye color from dark brown to almost black with light gray tint in central part and near eye contact line. Ratio of length to width of compound eye about 0.86; ratio of contact line length to eye length 0.33–0.35. Paired ocelli shoe-shaped, black with narrow white rim, located in front of eyes. Distance between inner margins of ocelli 0.08–0.10 and between their outer margins 0.96–0.98 total width of compound eyes (Fig. 6B). Apical article of maxillary palpus 1.12–1.14 times as long as preceding one. Dorsal surface of 7 th, 6 th, and 5 th articles of maxillary palpus with 17, 13 or 14, and 8 or 9 hyaline spines, respectively (Figs 6C, D). Apical article of labial palpus triangularly oval, 2.3 or 2.4 times as long as wide (Fig. 6E). Mandibles with four distal teeth (Fig. 6F). Fore and middle femur and tibia widened (Fig. 6G). Ratios of lengths to widths of femur, tibia and tarsus as shown in Table 10. Ratio of length of 3 rd tarsomere to total length of tarsus 0.30–0.31. Ventral surface of femora, tibiae and tarsi with spine-like chaetae as shown in Table 11. Middle and hind legs with coxal styli. Ratio of styli length to width of middle and hind coxae about 1.7. Abdominal segments I–VII with 1 + 1 eversible vesicles (Figs 7A, B). Posterior angle of urosternites II–IV, V, VI, and VII approximately 84–86°, 79°, 67°, and 77°, respectively. Ratios of lengths of urosternite, urocoxite and urostylus (without apical spine) II–IX as shown in Table 12. Inner posterior lobes of urocoxites VII protruding (Fig. 7B); ratio of length to width of one lobe about 0.54. Thoracic tergites, urotergites I–IV, urosternites, urocoxites I–VI without macrochaetae. Distribution of sublateral macrochaetae on other urocoxites and urotergites as shown in Table 12. Urocoxites IX with 1 + 1 outer and 6 + 6 inner sublateral spines. Urotergite X with small thin chaetae (Fig. 7C). Ovipositor weakly sclerotized, thickened, completely concealed by urocoxites IX, typical of genus Charimachilis. Anterior and posterior gonapophyses with 17 and 15 divisions, respectively (Figs 7D, E). Ultimate division of anterior gonapophyses with preapical spine and 2 apical lateral projections, one of which is pointed and sclerotized. Apical spines as long as 2 or 3 apical divisions combined. Anterior gonapophyses with 4 or 5 lateral digging spikes. Posterior gonapophyses with well developed, sclerotized curved apical horn and preapical spine, as long as 2 apical divisions combined. Distribution of sensory and simple chaetae on divisions of anterior and posterior gonapophyses as in Figs 7D, E. Male. Unknown. The species is probably parthenogenetic, similar to most its congeners. Differential diagnosis. Genus Charimachilis Wygodzinsky, 1939 includes 13 described species (KΑΡΓΙΝ 2019). Only females are known in 11 species. This makes it difficult to identify them. Charimachilis rostoviensis sp. nov. most closely resembles C. ukrainensis Stach, 1958 in ovipositor morphology. The main morphological differences between them are shown in Table 13. The posterior angle of urosternites is clearly acute (less than 75°) in C. rostoviensis sp. nov. and almost right (79–86°) in C. ukrainensis from Ukraine. Ratio of cercus length to body length is about 0.3 in C. ukrainensis and more than 0.4 in C. rostoviensis sp. nov. Ratio of stylus length (apical spines excluded) to urocoxite II–VII is 0.6–0.7 in C. ukrainensis and about 0.5 in C. rostoviensis sp. nov. C. rostoviensis sp. nov. is also similar to C. wahrmani Wygodzinsky, 1939 from Turkey in the structure to compound eye, maxillary palpus, urocoxite IX (WΥǤƟĐƶΙΝඌΚΥ 1959). C. wahrmani differs from the new species in ratio of distance between the inner margins of ocelli to the total width of compound eyes, ratio of stylus length to urocoxite VIII, number of sublateral spines on urocoxite IX, and the structure of the anterior and posterior gonapophyses (Table 13). Etymology. The species is named after Rostov-on-Don where it was collected; adjective. Habitats. Old artificial stands of pine Pinus sylvestris L. with undergrowth of Acer tataricum L. in the valley of the Temnik River. Under a trunk of a fallen dead pine.

Two new species of Machilinus Silvestri, 1905 (M. petrophilus Kaplin, sp. nov., M. obscurus Kaplin, sp. nov.) from Ukraine and one new species of Charimachilis Wygodzinsky, 1939 (C. rostoviensis Kaplin sp. nov.) from Southern Russia are described. The new species of Machilinus belong to the subgenus Machilinus s. str. with 1 + 1 eversible vesicles on urocoxites II–VII, and urostyli with apical spines; to the group “rupestris” with 2nd and 3rd articles of male maxillary palpus without ventral spines (M. obscurus sp. nov.), and with spines on these articles (M. petrophilus sp. nov.). Machilinus petrophilus sp. nov. differs from M. rocai Bach, 1975 in the color and ratio of length to width of compound eyes, the presence of numerous short chaetae on the clypeus, the structure of the lateral apophysis on the 2nd article of the male maxillary palpus, and the number of ventral spines on the legs. Machilinus obscurus sp. nov. differs from the other species of the group “rupestris” in the color of compound eyes, the presence of numerous short chaetae on the male frons and clypeus, and in other features. Charimachilis rostoviensis sp. nov. (Machilidae) most closely resembles C. ukrainensis Stach, 1958. Charimachilis rostoviensis sp. nov. differs from C. ukrainensis in the shape of posterior angle of urosternites, and ratios of lengths of urostyli and urocoxites.

Genus Coryphophthalmus Verhoeff, 1910 Type species: Coryphophthalmus banaticus Verhoeff, 1910., Published as part of Kaplin, V. G., Kiseleva, L. V. & Kozhevnikova, O. P., 2020, NEW SPECIES OF BRISTRLETAILS OF THE GENUS CORYPHOPHTHALMUS VERHOEFF, 1910 (ARCHAEOGNATHA: MACHILIDAE) FROM NORTH AND SOUTH OSSETIA, pp. 1-13 in Far Eastern Entomologist 406 on page 2, DOI: 10.25221/fee.406.1, http://zenodo.org/record/7165475

Petrobiellus sachalinensis sp. nov. (Figs 1–13) Holotype. Female (slide-mounted), Russia, Sakhalin Prov., western shore of northern Sakhalin I., near Aleksandrovsk-Sakhalinsky, 50°54′N 142°09′E, shoreline, under stones, 8.VIII.2019, V. Kaplin leg. Paratypes. 11 females (1 female on slide), same data as for holotype. Description. Female. Body length (not including antennae): 8.4–11.6 mm; body width: 2.9–3.1 mm; cercal length: 5.5–5.7 mm; total width of compound eyes: 0.94–1.00 mm; eye length: 0.54– 0.57 mm; paired ocelli width: 0.49–0.51 mm, length: 0.17–0.18 mm. Middle and hind coxal styli length: 0.60–0.65 mm. Ovipositor length: 3.35– 3.50 mm. General body colour whitish or slightly yellowish, almost without pigment. Antennal base, frons, lateral parts of clypeus, basal part of first and third articles of maxillary palps, mandibulae, hypopharynx, and first, third and basal half of second tarsomeres with brown or brown-violet pigment of weak or medium intensity. Maxillary and labial palps, flagellum and pedicellus of antennae, legs, coxal and abdominal styli without scales. Scapus with very sparse scales. Colour of scales whitish or light gray. Metanotum also with one pair, urotergites with three pairs of large spots of brown scales (Fig. 1). Antennae very long, 1.5– 1.9 times as long as body. Distal chains of flagellum divided into 14–23 annuli (Fig. 2). Clypeus without long thin bristles. Cercus approximately 0.48– 0.55 times as long as body, with about 21–24 divisions. Apex of cercus with one apical spike (Fig. 3). All divisions of cerci, except for apical one or two, with 1–3 colorless spines on inner side supporting cerci above substrate. Divisions in middle part of cerci also with one outer lateral spine (Fig. 4). Compound eyes dark, with bluish tinge (in ethanol). Ratio of length to width of compound eye about 1.13–1.15; ratio of contact line to length of eyes about 0.45. Paired ocelli sole-shaped, brown, with narrow white bordering, 2.8–3.0 times as wide as long. Distance between inner margins of paired ocelli 0.13–0.15 and between their outer margins 0.96–0.98 times total width of compound eyes (Fig. 5). Frons clearly convex between paired ocelli. Note. Length of the stylus does not include apical spines. Note. Petrobiellus takunagae Silvestri, 1943 is not included in the table due to insufficient data in its original description; the characters distinguishing this species from P. sachalinensis sp. nov. are given in the Comparison section after the description of the new species. . Apical article of maxillary palp 0.60–0.62 times as long as preceding one. Fifth article 1.04– 1.06 times as long as 4th article (Fig. 6). Dorsal surface of 7th, 6th and 5th articles of maxillary palp with 13–16, 10–13 and 3 hyaline spines, respectively. Apical article of labial palp fingerlike, 3.3–3.4 times as long as wide (Fig. 7). Mandibles with 2–3 distal teeth (Fig 8). Femora slightly widened. Ratios of length to width of femur, tibia and tarsus as shown in Table 1. Fore and middle legs shorter than hind legs. Fore and middle tarsus and tibia 0.77 and 0.83 times as long as those of hind legs, respectively. Ratio of length of apical hind tarsomere to total length of hind tarsus about 0.32. Ventral surface of legs without long thin bristles. Fore, middle and hind tarsi and hind tibia with slightly pigmented spines (Fig. 9). Number of spines as shown in Table 2. Middle and hind legs with coxal styli. Ratio of length of styli to width of middle and hind coxae 1.03–1.06 and 1.09, respectively. Urites I, VI and VII with 1 + 1, urites II–V with 2 + 2 eversible vesicles (Fig. 10). Posterior angle of urosternites II, III–V, VI, and VII approximately 118°, 110–112°, 107°, and 120°, respectively. Inner posterior lobes of urocoxites VII protruding. Ratio width to length of protruding lobe about 1.0. Ratios of lengths of stylus (without apical spine), urosternites and urocoxites II–IX as shown in Table 3. Ratio of lengths of urosternite and urocoxites I about 0.35. Urocoxites IX with 1 + 1 outer sublateral spines. Ovipositor slender, elongate, significantly protruding beyond apices of styli IX. Anterior and posterior gonapophyses with 52 or 53 divisions (Fig. 11). Four basal divisions of anterior gonapophyses and about 11 basal divisions of posterior gonapophyses glabrous. Apical spines of gonapophyses as long as three apical divisions combined. Distal divisions of anterior and posterior gonapophyses with seven or eight setae (not counting sensory setae and apical spines) (Figs 12, 13). Males are unknown; probably the species is parthenogenetic. Comparison. Petrobiellus sachalinensis sp.nov. can be distinguished from its congeners by the following characters: the general body colour whitish or slightly yellowish (vs yellowish or rufous); colour of scales whitish or light gray (vs brownish or dark brownish); body (excluding head and tarsi) almost without pigment (vs significantly pigmented); and paired ocelli almost sole-shaped (vs pistiliform, strongly narrowed in middle parts). The new species also differs from all other species of the genus in the most elongated compound eyes; relatively short coxal styli; long 4th article of maxillary palp; small number of short supporting spines in ventral part of tarsi; ratios of lengths of urosternite and urocoxite, stylus and urocoxite; posterior angle of urosternite; and number of divisions of ovipositor. The main morphological differences of females of P. sachalinensis sp. nov. from those of the other species of Petrobiellus are given in Table 4. According to the brief description of P. takunagae by Silvestri (1943), the distance between the paired ocelli in this species is about half the length of ocellus, and in P. sachalinensis sp. nov., this distance is almost equal to the length of ocellus. The new species differs from P. takunagae also in femora without pigment and tibiae poorly pigmented (vs femora and tibiae with intensive blackish-violet pigment); and four basal divisions of anterior gonapophyses glabrous (vs about 12 basal divisions of anterior gonapophyses glabrous). Etymology. The species name is derived from the toponym of Sakhalin. Habitats. Western shores of the northern part of Sakhalin Island. Under stones near the line of seashore (Fig. 14; see Addenda section)., Published as part of Kaplin, V. G., 2020, A new species of bristletails of the genus Petrobiellus (Microcoryphia: Machilidae) from Sakhalin, pp. 17-22 in Zoosystematica Rossica (Zoosyst. Rossica) (Zoosyst. Rossica) 29 (1) on pages 18-22, DOI: 10.31610/zsr/2020.29.1.17, {"references":["Kaplin V. G. 1980. New species of bristletails (Microcoryphia, Machilidae) from the Kuril Islands and Primorsky Krai. In: Lehr P. A. (Ed.). Taksonomiya nasekomykh Dal'nego Vostoka [Taxonomy of insects of the Far East]: 3 - 9. Vladivostok. (In Russian).","Uchida H. 1954. Apterygota of the Hachijo-Jima and its adjacent islands. Science Reports of the Faculty of Literature and Science, Hirosaki University, 1 (1): 1 - 17.","Silvestri F. 1943. Contributto alla conoscenza dei Machilidae (Insecta, Thysanura) del Giappone. Bollettino del Laboratorio di Zoologia, generale e agraria della R. Scuola superiore d'Agricoltura in Portici, 32: 283 - 306."]}

Pedetontus sp. These specimens were stored in isopropyl alcohol, and the exsertile vesicles are completely decomposed on almost all specimens. They are likely P. saltator, but it is impossible to tell how many vesicles existed on each of the abdominal segments, which would be diagnostic. As noted above, 3 males and 5 females of Pedetontoides atlanticus were collected at the same site by the same collector on 24 June 1986, but those specimens were in good condition (despite the dilute isopropyl alcohol preservative) and the shape of the urosternites and the presence of the parameres on the 8 th abdominal segment in males distinguished that genus. These specimens recorded here are not Pedetontoides. MISSISSIPPI: Hancock County: 6♂ + 5♀, Point Clear Island, pitfall human dung, 29 June 1986, P. K. Lago; 1♂, Point Clear Island, 15 August 1986, S. Testa; 2♂ + 2♀, Point Clear Island, pitfall, 17 August 1986, S. Testa; 1♂ + 1♀, Point Clear Island, pitfall human dung, 17 August 1986, S. Testa., Published as part of De Jong, Grant D., 2020, Jumping bristletail (Insecta: Apterygota: Microcoryphia) records in the southeastern United States, pp. 1-8 in Insecta Mundi 755 on page 5, DOI: 10.5281/zenodo.3703027

Pedetontus (Pedetontus) saltator Wygodzinsky and Schmidt, 1980 With two exsertile vesicles on the 6 th abdominal segment, this species is in the nominal subgenus with P. californicus and P. superior, two species found in California and Idaho (Silvestri 1911, Mendes 1990, De Jong 2014). It is distinguished from P. californicus by the shorter line of contact between the compound eyes and from P. superior by the shorter ovipositor in mature specimens, extending only to the tip of the styli of the 9 th abdominal segment (Wygodzinsky and Schmidt 1980). This species was previously reported only from the northeastern United States in Connecticut, Massachusetts, New Jersey, New York, and Pennsylvania. Online (Bartlett and Sellers 2018a, 2018b, 2018c, iNaturalist.org 2019) records of observations in GBIF of P. saltator include reports from Alaska, California, Massachusetts, Ontario, and Washington, DC, suggesting either a broad transcontinental distribution or a disjunct bi-coastal distribution. The Barcode of Life Database (BOLD) Barcode Index Number for P. saltator, based on a specimen from Alaska, is MOBIL6483-17 (IBLC 2017). The records detailed herein extend the distribution southward to Mississippi and North Carolina as new state records for the genus and species. Wygodzinsky and Schmidt (1980) suggested that the species might be parthenogenetic, because the 54 specimens they examined, including juveniles, were all females. These 32 specimens from the MEM corroborate that hypothesis. MISSISSIPPI: Issaqueena County: 1♀, 12 mi. SW Mayersville, Shipland Wildlife Management Area, 20 June 1992, M. S. Caterino; 1♀, 2 mi. SW Shipland, at light trap, 20 June 1992, P. K. Lago; Lafayette County: 1♀, 4 mi. W Oxford, cotton, 24 June 1977, A. E. Zuccaro; 1♀, Oxford, 31 October 1977, M. O. Mann; 1♀, Oxford, 18 October 1982, P. K. Lago; 1♀, Oxford, 19 June 2004, M. E. Pearson; Marshall County: 1♀, Holly Springs, 18 October 1980, H. H. Rather; Noxubee County: 1♀, Noxubee Wildlife Refuge, 8 December 1989, P. K. Lago; Oktibbeha County: 3♀, 6 mi. SW Starkville, 3 August 1984, J. Pooaitti; 1♀, Dorman Lake, 3 March 1985, J. Minr; 1♀, Dorman Lake, 3 March 1985, A. Schuster; 1♀, Starkville, 12 June 1985, C. M. Felland; Panola County: 3♀, 6 mi. SW Como, 20 June 1979, pitfall peripheral to cotton field, W. H. Cross, 4519 SA-1; 3♀, 3 mi. WSW Sardis, 1 August 1979, pitfall periph- eral to cotton field, W. H. Cross, 4622 W-1; 2♀, 14 mi. ESE Batesville, 15 September 1992, P. K. Lago; d ♀, 14 mi. ESE Batesville, 22 September 1992, P. K. Lago; Pontotoc County: 1♀, 1 mi. SE Ecru, 3 July 1980, pitfall peripheral to cotton field, W. H. Cross, 4750-1; 1♀, 1 mi. SE Ecru, 3 July 1980, pitfall in woods, W. H. Cross, 4750-6; 1♀, 1 mi. SE Ecru, 28 August 1980, pitfall in cotton field, W. H. Cross, 4760; 1♀, 1 mi. SE Ecru, 11 September 1980, pitfall in woods, W. H. Cross, 4761-3; 1♀, 1 mi. SE Ecru, 12 September 1980, pitfall in swamp, W. H. Cross, 4762-3; 1♀, 1 mi. SE Ecru, 25 September 1980, pit- fall in woods, W. H. Cross, 4763-5; NORTH CAROLINA: Edgecombe County: 1♀, 3 mi. W Tarboro, pitfall in cotton field, 13 June 1979, W. H. Cross, 4556; 1♀, 3 mi. W Tarboro, pitfall peripheral to cotton field, 13 June 1979, W. H. Cross, 4548 S1; 1♀, 3 mi. W Tarboro, pitfall in cotton field, 13 June 1979, W. H. Cross, 4549; 1♀, 2 mi. NW Tarboro, pitfall in cotton field, 24 July 1979, W. H. Cross, 4588; 1♀, 2 mi. NW Tarboro, pitfall in cotton field, 26 July 1979, W. H. Cross, 4591., Published as part of De Jong, Grant D., 2020, Jumping bristletail (Insecta: Apterygota: Microcoryphia) records in the southeastern United States, pp. 1-8 in Insecta Mundi 755 on pages 4-5, DOI: 10.5281/zenodo.3703027, {"references":["Wygodzinsky, P., and K. Schmidt, 1980. Survey of the Microcoryphia (Insecta) of the northeastern United States and adjacent provinces of Canada. American Museum Novitates 2701: 1 - 17.","Silvestri, F. 1911. Contributo alla conoscenza dei Machilidae dell'America settentrionale. Bolletino del Laboratorio de Zoologia Generale e Agraria della R. Scuola Superiore d'Agricoltura in Portici 5: 324 - 350.","Mendes, L. F. 1990. An annotated list of generic and specific names of Machilidae (Microcoryphia, Insecta) with identification keys for the genera and geographic notes. Estudos, Ensayos e Docu- mentos 155: 1 - 127.","De Jong, G. D. 2014. New species and records of jumping bristletails from the Rocky Mountains (Microcoryphia: Meinertellidae, Machilidae). Proceedings of the Entomological Society of Washington 116: 255 - 272.","Bartlett, T., and E. Sellers. 2018 a. BugGuide-Identification, Images, & Information For Insects, Spiders & Their Kin For the United States & Canada. Version 1.3. United States Geological Survey. Occurrence dataset https: // doi. org / 10.15468 / sk 2 lxk available at https: // www. gbif. org / occurrence / 1846836932. (Last accessed August 21, 2019.)","Bartlett, T., and E. Sellers. 2018 b. BugGuide-Identification, Images, & Information For Insects, Spiders & Their Kin For the United States & Canada. Version 1.3. United States Geological Survey. Occurrence dataset https: // doi. org / 10.15468 / sk 2 lxk available at https: // www. gbif. org / occurrence / 1846853357. (Last accessed August 21, 2019.)","Bartlett, T., and E. Sellers. 2018 c. BugGuide-Identification, Images, & Information For Insects, Spiders & Their Kin For the United States & Canada. Version 1.3. United States Geological Survey. Occurrence dataset https: // doi. org / 10.15468 / sk 2 lxk available at https: // www. gbif. org / occurrence / 1846853576. (Last accessed August 21, 2019.)"]}

Pedetontinus cf. atlanticus This specimen agrees in all aspects with the other specimens of P. atlanticus except that the first tarsal segment is darkened on all three pairs of legs. Bowser (2019a) reported specimens from the Western Interior Basin of British Columbia that are nearly identical to the figures in Mendes (1981a) but differ by the more conical nature of the distal segments of the male labial palpi, and he considered them to be potentially an undescribed species. Likewise, these Kentucky specimens may represent a separate species; however, the lack of multiple specimens in hand causes me to be reluctant to designate them as such. This record represents a new state record for this genus, family, and order. KENTUCKY: Carter County: 1 ♀, Carter���s Cave State Park, cliff face, 23 June 1983, G. T. Baker., Published as part of De Jong, Grant D., 2020, Jumping bristletail (Insecta: Apterygota: Microcoryphia) records in the southeastern United States, pp. 1-8 in Insecta Mundi 755 on page 4, DOI: 10.5281/zenodo.3703027, {"references":["Bowser, M. 2019 a. Archaeognatha of Canada. ZooKeys 819: 205 - 209.","Mendes, L. F. 1981 a. Notes et descriptions de Thysanoures de Nouveau Monde (Apterygota: Microcoryphia et Zygentoma). Nouvelle R e vue d'Entomologie 11: 221 - 231."]}

Pedetontus (Verhoeffilis) gershneri Allen, 1995 In this species, the 6 th abdominal segment has only one pair of exsertile vesicles, placing it in the subgenus Verhoeffilis Paclt with P. submutans Silvestri, P. persquamosus Silvestri, and P. calcaratus Silvestri, and distinguishing it from P. californicus Silvestri, P. superior Silvestri, and P. saltator Wygodzinsky and Schmidt in the nominal subgenus, which have two pairs of exsertile vesicles. The apical segments of the maxillary and labial palpi in this species lack strong spines, and it is known only from Arkansas, separating it from the other species in the subgenus Verhoeffilis, which are also mostly found in the Pacific Coast states and Canadian provinces (Allen 1995). This species was previously known only from the type series collected on Magazine Mountain, Arkansas (Allen 1995), and the specimens from the MEM were also collected at the type locality. While this species was described from moist forest floor among leaf and pine litter, Allen (1995) also reported Machiloides banksi Silvestri from xeric rock ledge habitat elsewhere on Magazine Mountain. It is therefore interesting that these species with apparently differing habitat preferences were both collected in the same sample on 17 May 1989. Perhaps pitfall traps were placed in intermediate habitats or multiple pitfall traps from different habitats were composited. ARKANSAS: Logan County: 1♂ + 3♀, Magazine Mt., 1350���, T6N R25W �� 16, 17 May 1989, J. MacGown & Q. Fang., Published as part of De Jong, Grant D., 2020, Jumping bristletail (Insecta: Apterygota: Microcoryphia) records in the southeastern United States, pp. 1-8 in Insecta Mundi 755 on page 4, DOI: 10.5281/zenodo.3703027, {"references":["Allen, R. T. 1995. Pedetontus gershneri, a new species of Machilidae from the interior highlands of North America (Insecta: Microcoryphia). Entomological News 106: 195 - 198."]}

Pedetontoides atlanticus Mendes, 1981 The distinctive males of this monospecific genus are easily recognized and separated from other southeastern United States machilid males due to the presence of parameres on both the eighth and ninth abdominal segments. Among the Nearctic Microcoryphia, this character is otherwise present only in the genus Meximachilis Wygodzinsky, represented in the United States by M. cokendolpheri Kaplin from New Mexico. In both males and females of P. atlanticus, the posterior angle of the fourth urosternite is acutely pointed, although it can approach a right angle, which is characteristic of Pedetontus species. Previously, this species was known only from the type series of 12 adults and 5 juveniles from Durham, North Carolina (Mendes 1981a). These records add Alabama, Arkansas, Georgia, and Mississippi as new state records to the known distribution of this species. ALABAMA: Cherokee County: 1♂ + 1♀, Little River Wildlife Management Area, 1300���1380���, T7S R10E ��15SW-16SE, Crest Road, 22 May 1990, D. Hidebrandt & T. Schiefer; DeKalb County: 2♂, DeSoto State Park, 1600���1700���, T6 S R10 E ��19W, Trail O, 18 May 1990, R. Brown & D. Pollock; 3♂, DeSoto State Park, 1400���1500���, T6 S R10 E ��29NW, pitfall trap in deciduous woods, 18���24 May 1990, D. Hildebrant & T. Schiefer; 3♀, DeSoto State Park, 1600���1700���, T6 S R10 E ��19W, carrion-baited pitfalls, 19-24 May 1990, J. Hildebrandt & J. MacGown; 1♂ + 2♀, DeSoto State Park, 1600���1700���, T6 S R10 E ��19W, pitfalls, 19���24 May 1990, J. Hildebrandt & T. Schiefer; 1♀, DeSoto State Park, 1360���1460���, T6 S R10 E ��19SE��� 20SW, 19 May 1990, Azalea Trail, R. Brown & D. Pollock; Jackson County: 1♂, Bingham Mountain, Davis Cove, 1,300���, 14 May 2004, P. K. Lago; Monroe County: 1♂, Haines Island Park, black light, N31��43���23��� W87��28���10���, 26 May 1995, R. L. Brown; ARKANSAS: Logan County: 1♂, Cove Lake, 1020���, T7 N R25 W ��35SE, blacklight trap, 14���20 May 1989, R. L., and B. B. Brown; 1♂, Magazine Mt, 2500���, T6N R25W ��23NW, 15���16 May 1989, R, L. Brown & J. MacGown; 1♂, Magazine Mt., 16 May 1989, P. R. Miller; 2♂, Magazine Mt., 1350���, T6 N R25 W �� 16, 17 May 1989, J. MacGown & Q. Fang; 1♂, Cove Lake Campground, pitfall, 18 May 1989, P. & G. Miller; GEORGIA: Bartow County: 1♂ + 3♀, Red Top Mountain State Park, pit trap ��� human dung, 3 May 1992, M. S. Caterino; MISSISSIPPI: Adams County: 1♂, Natchez State Park, 23 March 1985, R. L Brown & S. Cho; Franklin County: 2♂ + 1♀ + 2 Juveniles, Trib. of McGehee Cr., T6N R4E ��26SW, 27 July 1992, on trunk of tree, T. L. Schiefer; Grenada County: 2♂ + 2♀, T21N R2E ��12���13N & R3E ��7S���18N, 7���14 August 1991, pitfall traps in woods, M. W. MacGown; Hancock County: 3♂ + 5♀, Point Clear Island, 24 June 1987, S. Testa; Jones County: 1♂, Laurel, under rotten log, 4 September 1981, T. Ishee; Lafayette County: 1♀, 9 mi. NE Oxford, 19 October 1982, R. Upchurch; 1♀, 8 mi. NE Oxford, 10 October 1993, R. Weens; Oktibbeha County: 1♂, Starkville, 2 September 1981, white pan trap on ground, W. L. Cross; 1♀, 6 mi. SW Starkville, 3 September 1984, R. L. Brown; 4♂ + 2♀, Dorman Lake, pitfall trap, 11-15 August 1989, T. Schiefer; Pontotoc County: 1♀, 1 mi. SE Ecru, 2 July 1980, pitfall in woods, W. H. Cross, 4784-4; 1♀, 1 mi. SE Ecru, 27 August 1980, pitfall in woods, W. H. Cross, 4759-6; 1♀, 1 mi. SE Ecru, 28 August 1980, pitfall peripheral to cotton field, W. H. Cross, 4760-1; 1♀, 1 mi. SE Ecru, 28 August 1980, pitfall in swamp, W. H. Cross, 4760-1; 3♂, 1 mi. SE Ecru, 28 August 1980, pitfall in swamp, W. H. Cross, 4760-4; 1♀, 1 mi. SE Ecru, 12 September 1980, pitfall in swamp, W. H. Cross, 4762-3; 1♀, 1 mi. SE Ecru, 26 September 1980, pitfall in woods, W. H. Cross, 4764-2; 3♀, 1 mi. SE Ecru, 26 September 1980, pitfall in woods, W. H. Cross, 4764-3; Tishomingo County: 1♂, Tishomingo State Park, boulder formation near water, 17 June 1986, S. Testa., Published as part of De Jong, Grant D., 2020, Jumping bristletail (Insecta: Apterygota: Microcoryphia) records in the southeastern United States, pp. 1-8 in Insecta Mundi 755 on page 3, DOI: 10.5281/zenodo.3703027, {"references":["Mendes, L. F. 1981 a. Notes et descriptions de Thysanoures de Nouveau Monde (Apterygota: Microcoryphia et Zygentoma). Nouvelle R e vue d'Entomologie 11: 221 - 231."]}

Archaeognatha (Machilida plus dagger Monura), with similar to 450 extant species worldwide, represent a key lineage for the understanding of insect evolution. In a recently published phylogenetic analysis on Archaeognatha, based on a total-evidence approach, a Machilidae key fossil (Gigamachilis triassicus) from the Middle Triassic Lagerstatte of Monte San Giorgio and a representative of dagger Monura (Dasyleptus triassicus) from the same fossiliferous site, were not considered. In the present study, the phylogenetic analyses performed by Zhang et al. (2018) using maximum parsimony and Bayesian inference were repeated including these two key taxa; in addition, Tricholepidion gertschi and Lepisma saccharina were included as representatives of Zygentoma. In the total-evidence Bayesian analysis, dagger Monura was found to be paraphyletic, whereas in the maximum parsimony analysis, performed only on morphological characters, they were monophyletic. Moreover, the attribution of G. triassicus to Machilidae is now confirmed by these analyses, indicating that members of the extant group of Machilidae were already present in the Middle Triassic, at least similar to 100 Myr before the previous estimate.

The fauna of bristletails of the genus Lepismachilis Verhoeff, 1910 in Montenegro and Serbia includes only one species L. (Berlesilis) targionii (Grassi, 1887) with 2 + 2 eversible vesicles on abdominal urocoxites II–VI. Three new species of this genus are described: L. (Lepismachilis) prijepolja sp. nov., L. (Lepismachilis) limensa sp. nov. from Serbia, and L. (Lepismachilis) alexandrae sp. nov. from Montenegro. All described new species belong to the species group of the subgenus Lepismachilis s. str. with 2 + 2 eversible vesicles on abdominal urocoxites II–V. Lepismachilis prijepolja sp. nov. differs from L. y-signata Kratochvíl, 1945 and L. notata Stach, 1919 by the color, drawings and ratios of the compound eyes; ratios of sensory field on fore femur of male, number of divisions of ovipositor. Lepismachilis limensa sp. nov. differs from L. hauseri Bitsch, 1974 and L. abchasica Kaplin, 2017 by ratios of paired ocelli, sensory field on fore femur of male; ratios and chaetotaxy of maxillary and labial palps. Lepismachilis alexandrae sp. nov. differs from L. abchasica by the drawings of the compound eyes; ratios of paired ocelli, sensory field on fore femur of male; number of divisions of the parameres and gonapophyses. A list of the Machilidae occurring in Balkan Peninsula is also provided.

A review of geographic distribution, phylogenetic relationships, and evolutionary trends of six species of the South Palaearctic genus Haslundichilis Wygodzinsky is performed for the first time. Bristletails of the genus Haslundichilis are common in temperate forest and steppe open landscapes in the foothills and mountains of the East Caucasus, Central Asia, Northwest and East China, and South Korea. The genus originated in the forest landscapes of the West Stenopean (mixed) province wherefrom its representatives spread as far eastwards as South Korea and as far westwards as the mountains of Central Asia and the East Caucasus. The new species Haslundichilis daghestanicasp. n. is the closest to H. afghani, from which it differs in a greater body size, narrower eyes, and more developed sensory fields on the fore femur of the male.

A new bristletail species, Trigoniophthalmus fiagdoni sp. nov., is described from the North Ossetia-Alania. Its distinctive characters are discussed.

Review of the six genera of the Silvestrichilis group (Dilta Strand, Haslundiella Janetschek, Silvestrichilis Wygodzinsky, Silvestrichiloides Mendes, Haslundichilis Wygodzinsky, and Himalayachilis Wygodzinsky) and of the 17 species of the southern Palaearctic genus Silvestrichilis was performed for the first time with a discussion of their phylogenetic relationships and evolutionary trends based on comparative analysis of the geographic distribution of the species and of the distribution of plesiomorphic and apomorphic states of their main morphological features. Haslundichilis quadrii Wygodzinsky, 1952 from northwestern India and H. lindbergi Wygodzinsky, 1962 from western Afghanistan are transferred to the genus Haslundiella, and new combinations Haslundiella quadrii (Wygodzinsky, 1952), comb. n. and Haslundiella lindbergi (Wygodzinsky, 1962), comb. n. are established. A new species, Silvestrichilis polinaesp. n., differs from all the congeners in the strongly dilated fore femur of the male and female, in the maxillary palpus chaetotaxy, and in the structure of the sensory field on the fore femur of the male.