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GENUS SYMPHYLURINUS SILVESTRI, 1909 Type species: Symphylurinus grassi Silvestri, 1909., Published as part of Sánchez-García, Alba, Sendra, Alberto, Davis, Steven & Grimaldi, David A., 2023, Fossil diversity in ' dawn' hexapods (Diplura: Projapygoidea), with direct evidence for being chemically predaceous in the Cretaceous, pp. 847-870 in Zoological Journal of the Linnean Society 198 (3) on page 856, DOI: 10.1093/zoolinnean/zlac101, http://zenodo.org/record/8146868

SYMPHYLURINUS SP. 2 (FIGS 12, 13) Material: M-2233 (Figs 12, 13), Ettore Morone collection, Turin, Italy (available through the AMNH), probably female, adult; specimen virtually complete, with the right antenna and cercus cut off at the amber surface, observable laterally (both sides); the piece is a transparent, orange colour, but with internal fractures obscuring details, triangular in shape, 21 mm × 22 mm × 18 mm; syninclusions include a myriapod, some coprolites, fungal hyphae and other undetermined organic remains., Published as part of Sánchez-García, Alba, Sendra, Alberto, Davis, Steven & Grimaldi, David A., 2023, Fossil diversity in ' dawn' hexapods (Diplura: Projapygoidea), with direct evidence for being chemically predaceous in the Cretaceous, pp. 847-870 in Zoological Journal of the Linnean Society 198 (3) on page 857, DOI: 10.1093/zoolinnean/zlac101, http://zenodo.org/record/8146868

SYMPHYLURINUS SP. 1 (FIGS 6–11) Material: AMNH JZC-DR005 (Figs 6–8), Zigras collection (in AMNH), probably female, adult; specimen entirely preserved and observable ventrolaterally (left side), with the abdominal segments II and III swollen owing to taphonomic processes; the piece is a transparent, orange in colour, with a rounded shape, 32 mm × 20 mm in largest dimensions; syninclusions include seven oniscidean isopods, one acari, one ant, an undetermined number of Collembola hidden by bubbles, coprolites mainly associated with two of the isopods, fungal hyphae and other undetermined organic remains. AMNH KL-DR2022-1 (Figs 9–11) donated by K. Luzzi, probably female, adult; specimen entirely preserved and observable dorsally, ventrally and laterally (both sides); the right cercus (Fig. 10F) expelling the glandular substance probably as a stress behaviour in response to being trapped in resin; preserved in a rectangular chip of transparent yellow amber trimmed to 4mm × 8 mm × 1mm;syninclusions include two Coleoptera larvae, at least 11 nematodes, plant trichomes and other undetermined organic remains., Published as part of Sánchez-García, Alba, Sendra, Alberto, Davis, Steven & Grimaldi, David A., 2023, Fossil diversity in ' dawn' hexapods (Diplura: Projapygoidea), with direct evidence for being chemically predaceous in the Cretaceous, pp. 847-870 in Zoological Journal of the Linnean Society 198 (3) on page 856, DOI: 10.1093/zoolinnean/zlac101, http://zenodo.org/record/8146868

SYMPHYLURINOPSIS GEN.NOV. Zoobank registration: urn: lsid: zoobank. org:act: 5DD3AC90-782C-4455-91F0-8CF232C0951F Etymology: The generic name is based on the genus Symphylurinus Silvestri, 1909, with the Greek suffix - opsis (meaning, ‘sight, appearance’; thus ‘looking like’). The gender of the name is masculine. Type species: Symphylurinopsis punctatus sp. nov. by monotypy. Diagnosis: Male. The new genus is readily distinguished from all other genera in the family Projapygidae by the following combination of characters: cuticle dorsally and ventrally with distinct setigerous punctures; antenna short, 0.3× length of body, with 23 antennomeres, dolioform; antennomere II with secondary sexual characters, lacking secondary sexual characters on antennomere III and pyriform sensillum on antennomere VII; labial and maxillary palpi uniarticulate and elongate, the maxillary palpus longer than the labial palpus; legs short, slightly increasing in length from first to third pair; tibiae without comb of spatuliform setae and with calcars; claws subequal; cercus elongate, 0.3× length of body, with 15 segments plus the infundibuliform complex, basal segments I–V fused, without secondary sexual characters.

SYMPHYLURINOPSIS GEN.NOV. Zoobank registration: urn: lsid: zoobank. org:act: 5DD3AC90-782C-4455-91F0-8CF232C0951F Etymology: The generic name is based on the genus Symphylurinus Silvestri, 1909, with the Greek suffix - opsis (meaning, ‘sight, appearance’; thus ‘looking like’). The gender of the name is masculine. Type species: Symphylurinopsis punctatus sp. nov. by monotypy. Diagnosis: Male. The new genus is readily distinguished from all other genera in the family Projapygidae by the following combination of characters: cuticle dorsally and ventrally with distinct setigerous punctures; antenna short, 0.3× length of body, with 23 antennomeres, dolioform; antennomere II with secondary sexual characters, lacking secondary sexual characters on antennomere III and pyriform sensillum on antennomere VII; labial and maxillary palpi uniarticulate and elongate, the maxillary palpus longer than the labial palpus; legs short, slightly increasing in length from first to third pair; tibiae without comb of spatuliform setae and with calcars; claws subequal; cercus elongate, 0.3× length of body, with 15 segments plus the infundibuliform complex, basal segments I–V fused, without secondary sexual characters., Published as part of Sánchez-García, Alba, Sendra, Alberto, Davis, Steven & Grimaldi, David A., 2023, Fossil diversity in ' dawn' hexapods (Diplura: Projapygoidea), with direct evidence for being chemically predaceous in the Cretaceous, pp. 847-870 in Zoological Journal of the Linnean Society 198 (3) on page 852, DOI: 10.1093/zoolinnean/zlac101, http://zenodo.org/record/8146868

ELECTROPROJAPYX GEN.NOV. Zoobank registration: urn: lsid: zoobank. org:act: 3496E6E3-6D13-4753-8808-BF7DA8CC3B2D Etymology: The generic name is a combination of the Latin electro (meaning, ‘amber’), and Projapyx Cook, 1899, type genus of the family. The gender of the name is masculine. Type species: Electroprojapyx alchemicus sp. nov. by monotypy. Diagnosis: The new genus is readily distinguished from all other genera of Projapygidae by the following combination of characters: body elongate; antenna elongate, 0.5× length of body, slightly tapered, with 27 antennomeres, moniliform, without secondary sexual characters on antennomeres II and III, lacking pyriform sensillum on antennomere VII; legs elongate and slender, metathoracic leg distinctly longer than others; tibiae with calcars; claws subequal; cercus short and slender (not compacted), 0.1× length of body, with at least seven segments plus the infundibuliform complex, without secondary sexual characters.

ELECTROPROJAPYX GEN.NOV. Zoobank registration: urn: lsid: zoobank. org:act: 3496E6E3-6D13-4753-8808-BF7DA8CC3B2D Etymology: The generic name is a combination of the Latin electro (meaning, ‘amber’), and Projapyx Cook, 1899, type genus of the family. The gender of the name is masculine. Type species: Electroprojapyx alchemicus sp. nov. by monotypy. Diagnosis: The new genus is readily distinguished from all other genera of Projapygidae by the following combination of characters: body elongate; antenna elongate, 0.5× length of body, slightly tapered, with 27 antennomeres, moniliform, without secondary sexual characters on antennomeres II and III, lacking pyriform sensillum on antennomere VII; legs elongate and slender, metathoracic leg distinctly longer than others; tibiae with calcars; claws subequal; cercus short and slender (not compacted), 0.1× length of body, with at least seven segments plus the infundibuliform complex, without secondary sexual characters., Published as part of Sánchez-García, Alba, Sendra, Alberto, Davis, Steven & Grimaldi, David A., 2023, Fossil diversity in ' dawn' hexapods (Diplura: Projapygoidea), with direct evidence for being chemically predaceous in the Cretaceous, pp. 847-870 in Zoological Journal of the Linnean Society 198 (3) on page 850, DOI: 10.1093/zoolinnean/zlac101, http://zenodo.org/record/8146868

ELECTROPROJAPYX ALCHEMICUS SP.NOV. (FIG. 2) Zoobank registration: urn: lsid: zoobank. org:act: B2A57148-0988-4316-9B09-99264D417E6F Etymology: The name is derived from the Latin Alchemia, an early, protoscientific practice, and the Latin suffix - icus. Holotype: AMNH JZCBu-1957 (Fig.2), Zigras collection (in AMNH), sex unknown, adult; specimen entirely preserved and observable laterally; the holotype is at the surface of a corner; the piece is a dark orange colour and turbid, irregular in shape, 28 mm × 14 mm in largest dimensions, with borings of Pholadidae; syninclusions include one heteropteran nymph, one partial wasp, at least four Collembola (three Symphypleona, one Entomobryomorpha), one acari, plant trichomes and other undetermined organic remains. Note on preserved behaviour: Specimen AMNH JZCBu-1957 (Fig. 2) preserves an example of predatory behaviour fossilized in Burmese amber. The specimen of E. alchemicus has between its antennae a Collembola Symphypleona (0.5 mm in length), while it is also releasing a glandular substance from the cercal tips. The antennae of E. alchemicus are in close contact with the Collembola (Fig. 2D); the elongated abdomen of the dipluran is elevated and curved above the thorax and head with the cerci pointing to the front (Fig. 2A, C); there is a line of droplets and short streams of secretion connected by long, fine filaments (presumably, strands of silk) issuing from the tips of both cerci (Fig. 2B); several of these strands are nearly the length of the abdomen. The position of the collembolan, posture of the dipluran and presence of cercal secretions can only be explained as an attack movement (cf. San Martín, 1969: 127, fig. 14; San Martín, 1963). Fluid resin probably flowed over the pair of hexapods just as the dipluran was attacking. Occurrence: Mid-Cretaceous Burmese amber (latest Cenomanian) near Myitkyina, in Kachin Province. Diagnosis: As for genus, by monotypy. Description Sex unknown, adult. Body: Length 2.49 mm, slender. Cuticle altered by preservation, with no visible details. Clothing setae smooth; macrosetae (M) as indicated below. Head: Length 0.29 mm, 0.12× length of body; dorsum with few smooth setae. A structure that resembles a labial palpus protrudes from the inferior part of the head. Both antennae (Fig. 2D) complete (non-regenerated), elongate, reaching second abdominal segment, length 1.23 mm, 0.49× length of body, slightly tapered, with 27 antennomeres; antennomeres I and II subrectangular, distinctly longer than others, together 0.15 mm, antennomere II somewhat swollen distally probably owing to preservation; antennomere III onwards moniliform; medial antennomeres length 0.05 mm, width 0.03 mm; antennomeres II and III without secondary sexual characters, antennomere VII without pyriform sensillum; trichobothria not visible; whorls of setae poorly visible, the few visible setae smooth. Thorax: 0.29× length of body; pronotum 0.17 mm, mesonotum 0.25 mm, metanotum 0.30 mm. Macrosetae visible in anterior and posterior position in pro-, meso- and metanotum, exact pattern not discernible owing to preservation, all macrosetae long and with thin barbs along distal half or two-thirds of each macroseta. Legs slender and elongate, with metathoracic leg distinctly longer than others, reaching abdominal segment III; some segments of legs difficult to distinguish owing to preservation; femur longest segment; tibia and tarsus equal in length; tibia with two typical large, stout, simple calcar spurs near ventral apex; tarsus with few visible smooth setae on ventral side; pretarsus with subequal claws, empodium lacking. Abdomen: Length 1.48 mm, 0.59× length of body. Macrosetae anterior (A) and posterior (P) visible at least from tergites I to VI; macrosetae P visible from tergites VIII to X; all macrosetae long and with thin barbs along distal half or two-thirds of each macroseta. Sternite I with subcylindrical lateral subcoxal appendages, slightly tapered toward tip; styli present from sternites I to VII; measurements of styli and subcoxal appendage not possible owing to preservation; genital papilla not visible. Cercus (Fig. 2B) short, length 0.33 mm, 0.13× length of body, slender (segments not compacted), with greatest width in visible segment II, with at least seven segments (basal portion not clearly visible) not counting the infundibuliform complex, without tegumentary expansions or modified setae or other secondary sexual characters on inner side; visible segments I–VII with a distal whorl of smooth setae; visible segment VII terminating in a longitudinally striated excretory tube, the infundibuliform complex., Published as part of Sánchez-García, Alba, Sendra, Alberto, Davis, Steven & Grimaldi, David A., 2023, Fossil diversity in ' dawn' hexapods (Diplura: Projapygoidea), with direct evidence for being chemically predaceous in the Cretaceous, pp. 847-870 in Zoological Journal of the Linnean Society 198 (3) on pages 850-852, DOI: 10.1093/zoolinnean/zlac101, http://zenodo.org/record/8146868, {"references":["San Martin PR. 1969. Descripcion de una nueva especie de Projapygidae del Uruguay y estudio de los cercos en la familia (Diplura). ReVista Brasileira de Biologia 29: 121 - 134."]}

SYMPHYLURINOPSIS PUNCTATUS SP.NOV. (FIGS 3–5) Zoobank registration: urn: lsid: zoobank. org:act: 12E8CFF1-2EDD-4571-B332-498D626E0596 Diplura: Campodeidae; Grimaldi & Engel (2005): p. 118, fig. 3.34. Etymology: The name is derived from the Latin punctatus, dotted, in reference to the cuticular punctures. Holotype: M-2232 (Figs 3–5), Ettore Morone collection, Turin, Italy (available through the AMNH), male, adult; specimen entirely preserved and observable dorsally and ventrally; the piece is a transparent, orange in colour, 23 mm × 11 mm in largest dimensions; without syninclusions., Published as part of Sánchez-García, Alba, Sendra, Alberto, Davis, Steven & Grimaldi, David A., 2023, Fossil diversity in ' dawn' hexapods (Diplura: Projapygoidea), with direct evidence for being chemically predaceous in the Cretaceous, pp. 847-870 in Zoological Journal of the Linnean Society 198 (3) on page 852, DOI: 10.1093/zoolinnean/zlac101, http://zenodo.org/record/8146868, {"references":["Grimaldi D, Engel MS. 2005. EVolution of the insects. Cambridge: Cambridge University Press."]}

SÁNCHEZ-GARCÍA, ALBA, RÍOS, MARÍA, ARNAL, FERNANDO ANTONIO MARTÍN, GAMONAL, ARTURO, CRUZADO-CABALLERO, PENÉLOPE, GONZÁLEZ-DIONIS, JAVIER, VLACHOS, EVANGELOS, GUERRERO-ARENAS, ROSALÍA, CRESPO, VICENTE D. (2023): The 4 Palaeontological Virtual Congress. Palaeoentomology 6 (1): 1-2, DOI: 10.11646/palaeoentomology.6.1.1, URL: http://dx.doi.org/10.11646/palaeoentomology.6.1.1

Anatoliacampa diclensis Sendra, Tusun & Satar sp. nov. Etymology. The specific epithet refers to both the name of the cave and the name of the region of the type’ s locality. Type locality. Dicle Cave, Bozaba village, Dicle district, Diyarbakır province, Southeastern Anatolian region, Turkey. Holotype. TURKEY. ♀ (labelled as holotype- ♀ DUZM-2121). Dicle Cave, Bozaba village, Dicle district, Diyarbakır province, Southeastern Anatolian region, 38 ◦ 19 ′ 23 ′′ N, 40 ◦ 06 ′ 25 ′′ , E. Sadreddin TUSUN and Ali SATAR leg. Paratype. TURKEY. 6 (labelled as paratype-61 MZB (MCNB) 2022- 5694), same locality, date, and collectors as holotype. Other material. ♀ (coll. AS), same locality, date and collectors as types; specimen used for scanning electron microscopy (SEM) and DNA analysis. Description. Body . Body length 5.6 mm (female, holotype) and 5.8 mm (male, paratype). Epicuticle smooth under compound microscope and SEM; with thin, medium-sized clothing covered by thin distal barbs. Head. Antennae complete (Figs. 1 and 2), with 41 antennomeres. Small, subcylindrical sensillum on third antennomere located in ventral position between c and d macrosetae; flagellum of the trichobothria very long (Fig. 2b). Central antennomeres 2.3 × as long as wide, with large barbed and short setae distributed in two to three whorls, in addition to one distal whorl of about 12 thin gouge sensilla 40–50 μm long (Fig. 2a, c, d). Apical antennomere with sensorial equipment of: cupuliform organ with about 12 complex olfactory chemoreceptors (each chemoreceptor is composed of a central column with the typical apical hole surrounded by irregular folds in a cauliflower shape and its surface is entirely reticulated and perforated) (Fig. 1a and b); numerous gouge sensilla 25–50 μm long (Fig. 1c, f); a few short bacilliform sensilla; several glandular setae on the edge of the hole of the cupuliform organ (Fig. 1a, d, e); and numerous long barbed setae. Frontal process slightly protruding, plain, and with one distal and two posterior non-tubercular setae; macrosetae along the insertion line of antennomere I similar in length, x setae the longest (a/i/p/x with relative lengths of 23/25/21/ 40 in holotype), all with very thin distal barbs. Labial palpus suboval, with a bacilliform latero-external sensillum, two guard setae on internal side, up to 8 setae on anterior border, and up to 150 neuroglandular setae in holotype, and 180 neuroglandular setae in paratype. Thorax. Macrosetal distribution (Fig. 3): pronotum with 1 + 1 ma, 2 + 2 la 1,3 , 2 + 2 lp 2,3 ; mesonotum with 1 + 1 ma, 2 + 2 la 1,2 , 2 + 2 lp 2,3 ; metanotum with 1 + 1 ma, 2 + 2 la 1,2 , 2 + 2 lp 2,3 . All notal macrosetae long and with thin barbs along distal half to two thirds; marginal setae longer than clothing setae, with thin barbs along distal half. Legs elongated, pretarsus of metathoracic leg overpasses end of abdomen. Lengths of metathoracic leg segments on holotype / paratype: coxa 0.26/0.30, trochanter 0.20/0.25, femur 0.98/1.05, tibia 1.19/1.30, tarsus 0.88/ 0.90; total: 3.51/3.80. Femora I–III with two long barbed dorsal macrosetae in the distal third of femur and one shorter barbed ventral macroseta inserted near central position (Fig. 4a). Tibia I with one or two short barbed ventral macrosetae; tibiae II–III without macrosetae. Calcars on tibiae covered with thin, short barbs. Tarsi with two rows of thick ventral setae with very thin barbs except on the apical portion. Four long barbed setae on dorsal subapical end of the tarsi. Claws (Fig. 4b–f) subequal with well-developed crests and a slightly backward overhang on both claws; the dorsal side is almost smooth and the ventral side has thin longitudinal grooves. Pretarsal lateral process start in laminar shape and extend into a narrow axis overpassing the end of the claws, which is divided into multiple fringes: simple or subdivided and with a hook ending. Abdomen. Distribution of macrosetae on tergites: 1 + 1 post 1 on I–IV; 0 + 0, 0 + 1 la, 3 + 3 post 1–3 or 4 + 4 post 1–4 on V; 0 + 0, 0 + 1, 1 + 1 la, 4 + 4 post 1–4 on VI–VII; 5 + 5 or 6 + 6 post 1–6 on VIII, and 8 + 8 post 1–8 on abdominal segment IX. All post urotergal macrosetae long and covered by thin barbs along distal four-fifths; la urotergal macrosetae shorter than post macrosetae, covered by barbs along distal half. Urosternite I with 11 + 12 macrosetae on holotype and 15 + 17 macrosetae on paratype (Fig. 5a and b); urosternites II–VII with 7 + 7–8 + 8 macrosetae; urosternite VIII with 3 + 3 macrosetae; all urosternal macrosetae robust and large, covered by long barbs along distal third to four-fifths. Styli with apical, subapical, and ventromedial setae completely surrounded by thin and short barbs (Fig. 6b). Eversible vesicles large, with two distinct zones: the distal one with an almost smooth surface with a sinuous border and the proximal one with a rough surface densely covered by minute dots (Fig. 6a). Gonopore of the genital papilla surrounded by 19 short setae (in paratype male). Cerci lost in all three specimens. Secondary sex characters. Male urosternite I (Fig. 5a) with very large appendages, almost subcylindrical, with up to 200 glandular a 1 setae. Female urosternite I (Fig. 5b) with thinner subcylindrical appendages, almost coniform, with up to 11 glandular a 1 setae. 3.2. Molecular analysis The new COI sequences have been uploaded to Genbank with codes: OM680964-OM680970. The selected nucleotide substitution model after alignment was GTR + G + I (BIC = 8561.95), with the proportion of invariant sites (I = 0.45) and estimated alpha parameter for the gamma distribution (α = 0.99), indicating a significant heterogeneity in the DNA substitution among sites. The ML phylogenetic tree showed Plusiocampinae sequences form a well-supported clade, nested within Campodeidae and clearly distinct from Japygidae (Fig. 9). Anatoliacampa diclensis Sendra, Tusun & Satar sp. nov grouped with Plusiocampa (Plusiocampa) imereti Sendra & Barjadze, 2021 (Sendra et al., 2021d) from Georgia and Plusiocampa (Stygiocampa) bureschi Silvestri, 1931 from Bulgaria, whereas Plusiocampa taxa from Iberian Peninsula [e.g. Plusiocampa (Plusiocampa) gadorensis Sendra, 2001, Plusiocampa (Plusiocampa) baetica Sendra, 2004 (Sendra et al., 2004) or Cestocampa iberica Sendra & Cond´e, 2012 (Sendra et al., 2012)] clustered in separate clades. 3.3. Habitat Dicle cave is excavated in carbonate rocks of the Fırat formation, Eocene to Miocene in age. It has one artificial entrance, 2 m × 1.5 m size, protected by a metal grid. It gives access to a large room with plenty of speleothems, 80 m long and 60 m wide, which goes down to a depth of 13 m (Fig. 7). The cave has no water pools or watercourse but some deep corners have hidden places where specimens of Anatoliacampa diclensis Sendra, Tusun & Satar gen. et sp. nov. were sampled. In summer, the outside environment reaches 35 ◦ C and 27% humidity, while the general atmosphere inside the cave stays at 23 ◦ C and 60% humidity., Published as part of Sendra, Alberto, Palero, Ferran, Sánchez-García, Alba, Selfa, Jesús, Tusun, Sadreddin & Satar, Ali, 2022, New evidence for an Anatolian bridge: Colonization of Euromediterranean lands by cave-adapted Plusiocampinae (Diplura, Campodeidae), with establishment of a new genus, pp. 205-214 in Zoologischer Anzeiger 301 on pages 208-210, DOI: 10.1016/j.jcz.2022.10.006, http://zenodo.org/record/8164054, {"references":["Sendra, A., Palero, F., S´anchez-Garcia, A., Jim´enez-Valverde, A., Selfa, J., Maghradze, E., Barjadze, S., 2021 d. A new Diplura species from Georgia caves, Plusiocampa (Plusiocampa) imereti (Diplura, Campodeidae), with morphological and molecular data. Eur. J. Taxon. 778, 71 - 85. https: // doi. org / 10.5852 / ejt. 2021.778.1567.","Silvestri, F., 1931. Contributo alla conoscenza dei Campodeidae (Thysanura) delle grotte della Bulgaria. Bulletin des Institutions royales d' Histoire naturelle a´Sofia 6, 97 - 107.","Sendra, A., 2001. Dipluros campodeidos (Diplura: Campodeidae) de las grutas almerienses (Almeria, Espatna). Zool. Baetica 12, 71 - 82.","Sendra, A., Lara, M. D., Ruiz Aviles, F., Tinaut, A., 2004. Une nouvelle esp`ece du genre Plusiocampa Silvestri, 1912 (Diplura, Campodeidae) et donn´ees pour sa reconstruction pal´eobiog´eographique dans les B´etiques. Subterr. Biol. 2, 113 - 122.","Sendra, A., Arnedo, M. A., Ribera, C., Teruel, S., Bidegaray-Batista, L., Conde´, B., 2012. Revision of Cestocampa Cond´e (Diplura, Campodeidae), with description of a new species from caves in the eastern iberian Peninsula. Zootaxa 3252, 43 - 56. https: // doi. org / 10.11646 / zootaxa. 3252.1.2."]}

Genus Anatoliacampa Sendra, Tusun & Satar gen. nov. Type species. Anatoliacampa diclensis Sendra, Tusun & Satar sp. nov. Etymology. The genus is named in reference to the Anatolian Peninsula and the suffix -campa, traditionally used in Campodeidae taxonomy. The gender of the name is feminine. Diagnosis. Troglomorphic body and appendages. Cupuliform organ of antennae with complex olfactory chemoreceptors; frontal process of head slightly protruding, with non-tubercular setae. Pronotum 1 + 1 ma, 2 + 2 la 1,3 , 2 + 2 lp 2,3 ; mesonotum and metanotum 1 + 1 ma, 2 + 2 la 1,2 , 2 + 2 lp 2,3 . Femora I–III with two dorsal macrosetae. Tibiae II–III without macrosetae. Claws subequal, with well-developed crests. Pretarsal process start in a laminar shape and extend to a narrow axis, divided into multiple fringes: simple or subdivided with a hook ending. Urotergites 1 + 1 post 1 on I–IV; 0 + 0 or 1 + 1 la, 3 + 3–4 + 4 post 1–4 on V–VII, 5 + 5–6 + 6 post 1–6 on VIII, and 8 + 8 post 1–8 on IX. Urosternite I with up to 32 macrosetae; urosternites II–VII with up to 16 macrosetae; urosternite VIII with 3 + 3 macrosetae. Male urosternite I appendages subcylindrical, with up to 200 glandular a 1 setae. Female urosternite I appendages coniform, with up to 11 glandular a 1 setae., Published as part of Sendra, Alberto, Palero, Ferran, Sánchez-García, Alba, Selfa, Jesús, Tusun, Sadreddin & Satar, Ali, 2022, New evidence for an Anatolian bridge: Colonization of Euromediterranean lands by cave-adapted Plusiocampinae (Diplura, Campodeidae), with establishment of a new genus, pp. 205-214 in Zoologischer Anzeiger 301 on page 207, DOI: 10.1016/j.jcz.2022.10.006, http://zenodo.org/record/8164054

Anatoliacampa diclensis Sendra, Tusun & Satar sp. nov. Etymology. The specific epithet refers to both the name of the cave and the name of the region of the type’ s locality. Type locality. Dicle Cave, Bozaba village, Dicle district, Diyarbakır province, Southeastern Anatolian region, Turkey. Holotype. TURKEY. ♀ (labelled as holotype- ♀ DUZM-2121). Dicle Cave, Bozaba village, Dicle district, Diyarbakır province, Southeastern Anatolian region, 38 ◦ 19 ′ 23 ′′ N, 40 ◦ 06 ′ 25 ′′ , E. Sadreddin TUSUN and Ali SATAR leg. Paratype. TURKEY. 6 (labelled as paratype-61 MZB (MCNB) 2022- 5694), same locality, date, and collectors as holotype. Other material. ♀ (coll. AS), same locality, date and collectors as types; specimen used for scanning electron microscopy (SEM) and DNA analysis. Description. Body . Body length 5.6 mm (female, holotype) and 5.8 mm (male, paratype). Epicuticle smooth under compound microscope and SEM; with thin, medium-sized clothing covered by thin distal barbs. Head. Antennae complete (Figs. 1 and 2), with 41 antennomeres. Small, subcylindrical sensillum on third antennomere located in ventral position between c and d macrosetae; flagellum of the trichobothria very long (Fig. 2b). Central antennomeres 2.3 × as long as wide, with large barbed and short setae distributed in two to three whorls, in addition to one distal whorl of about 12 thin gouge sensilla 40–50 μm long (Fig. 2a, c, d). Apical antennomere with sensorial equipment of: cupuliform organ with about 12 complex olfactory chemoreceptors (each chemoreceptor is composed of a central column with the typical apical hole surrounded by irregular folds in a cauliflower shape and its surface is entirely reticulated and perforated) (Fig. 1a and b); numerous gouge sensilla 25–50 μm long (Fig. 1c, f); a few short bacilliform sensilla; several glandular setae on the edge of the hole of the cupuliform organ (Fig. 1a, d, e); and numerous long barbed setae. Frontal process slightly protruding, plain, and with one distal and two posterior non-tubercular setae; macrosetae along the insertion line of antennomere I similar in length, x setae the longest (a/i/p/x with relative lengths of 23/25/21/ 40 in holotype), all with very thin distal barbs. Labial palpus suboval, with a bacilliform latero-external sensillum, two guard setae on internal side, up to 8 setae on anterior border, and up to 150 neuroglandular setae in holotype, and 180 neuroglandular setae in paratype. Thorax. Macrosetal distribution (Fig. 3): pronotum with 1 + 1 ma, 2 + 2 la 1,3 , 2 + 2 lp 2,3 ; mesonotum with 1 + 1 ma, 2 + 2 la 1,2 , 2 + 2 lp 2,3 ; metanotum with 1 + 1 ma, 2 + 2 la 1,2 , 2 + 2 lp 2,3 . All notal macrosetae long and with thin barbs along distal half to two thirds; marginal setae longer than clothing setae, with thin barbs along distal half. Legs elongated, pretarsus of metathoracic leg overpasses end of abdomen. Lengths of metathoracic leg segments on holotype / paratype: coxa 0.26/0.30, trochanter 0.20/0.25, femur 0.98/1.05, tibia 1.19/1.30, tarsus 0.88/ 0.90; total: 3.51/3.80. Femora I–III with two long barbed dorsal macrosetae in the distal third of femur and one shorter barbed ventral macroseta inserted near central position (Fig. 4a). Tibia I with one or two short barbed ventral macrosetae; tibiae II–III without macrosetae. Calcars on tibiae covered with thin, short barbs. Tarsi with two rows of thick ventral setae with very thin barbs except on the apical portion. Four long barbed setae on dorsal subapical end of the tarsi. Claws (Fig. 4b–f) subequal with well-developed crests and a slightly backward overhang on both claws; the dorsal side is almost smooth and the ventral side has thin longitudinal grooves. Pretarsal lateral process start in laminar shape and extend into a narrow axis overpassing the end of the claws, which is divided into multiple fringes: simple or subdivided and with a hook ending. Abdomen. Distribution of macrosetae on tergites: 1 + 1 post 1 on I–IV; 0 + 0, 0 + 1 la, 3 + 3 post 1–3 or 4 + 4 post 1–4 on V; 0 + 0, 0 + 1, 1 + 1 la, 4 + 4 post 1–4 on VI–VII; 5 + 5 or 6 + 6 post 1–6 on VIII, and 8 + 8 post 1–8 on abdominal segment IX. All post urotergal macrosetae long and covered by thin barbs along distal four-fifths; la urotergal macrosetae shorter than post macrosetae, covered by barbs along distal half. Urosternite I with 11 + 12 macrosetae on holotype and 15 + 17 macrosetae on paratype (Fig. 5a and b); urosternites II–VII with 7 + 7–8 + 8 macrosetae; urosternite VIII with 3 + 3 macrosetae; all urosternal macrosetae robust and large, covered by long barbs along distal third to four-fifths. Styli with apical, subapical, and ventromedial setae completely surrounded by thin and short barbs (Fig. 6b). Eversible vesicles large, with two distinct zones: the distal one with an almost smooth surface with a sinuous border and the proximal one with a rough surface densely covered by minute dots (Fig. 6a). Gonopore of the genital papilla surrounded by 19 short setae (in paratype male). Cerci lost in all three specimens. Secondary sex characters. Male urosternite I (Fig. 5a) with very large appendages, almost subcylindrical, with up to 200 glandular a 1 setae. Female urosternite I (Fig. 5b) with thinner subcylindrical appendages, almost coniform, with up to 11 glandular a 1 setae. 3.2. Molecular analysis The new COI sequences have been uploaded to Genbank with codes: OM680964-OM680970. The selected nucleotide substitution model after alignment was GTR + G + I (BIC = 8561.95), with the proportion of invariant sites (I = 0.45) and estimated alpha parameter for the gamma distribution (α = 0.99), indicating a significant heterogeneity in the DNA substitution among sites. The ML phylogenetic tree showed Plusiocampinae sequences form a well-supported clade, nested within Campodeidae and clearly distinct from Japygidae (Fig. 9). Anatoliacampa diclensis Sendra, Tusun & Satar sp. nov grouped with Plusiocampa (Plusiocampa) imereti Sendra & Barjadze, 2021 (Sendra et al., 2021d) from Georgia and Plusiocampa (Stygiocampa) bureschi Silvestri, 1931 from Bulgaria, whereas Plusiocampa taxa from Iberian Peninsula [e.g. Plusiocampa (Plusiocampa) gadorensis Sendra, 2001, Plusiocampa (Plusiocampa) baetica Sendra, 2004 (Sendra et al., 2004) or Cestocampa iberica Sendra & Cond´e, 2012 (Sendra et al., 2012)] clustered in separate clades. 3.3. Habitat Dicle cave is excavated in carbonate rocks of the Fırat formation, Eocene to Miocene in age. It has one artificial entrance, 2 m × 1.5 m size, protected by a metal grid. It gives access to a large room with plenty of speleothems, 80 m long and 60 m wide, which goes down to a depth of 13 m (Fig. 7). The cave has no water pools or watercourse but some deep corners have hidden places where specimens of Anatoliacampa diclensis Sendra, Tusun & Satar gen. et sp. nov. were sampled. In summer, the outside environment reaches 35 ◦ C and 27% humidity, while the general atmosphere inside the cave stays at 23 ◦ C and 60% humidity.

Campodea (Dicampa) catalana Denis, 1930 Campodea (Dicampa) catalana Denis, 1930: 28, figs 7���13. Material examined KYRGYZSTAN ��� 6 ♂♂, 11 ♀♀; Osh Province, Nookat District, Abshir Say River; 40��08���50��� N, 72��21���52��� E; alt. 1851 m; 21 Jul. 2019; Alberto Sendra leg.; endogean habitat near Cupressus tree; Coll. AS. Remarks Campodea (Dicampa) catalana is an abundant species in Western Mediterranean soil habitats. This finding seems to show a disjunct distribution on both sides of the Mediterranean region., Published as part of Sendra, Alberto, S��nchez-Garc��a, Alba, Selfa, Jes��s, Milko, Dmitry A. & Ferreira, Rodrigo Lopes, 2021, Campodeidae (Hexapoda: Diplura) from Kyrgyzstan, Central Asia, with the description of a remarkable new genus and species, pp. 1-20 in European Journal of Taxonomy 782 on page 4, DOI: 10.5852/ejt.2021.782.1585, http://zenodo.org/record/5761387, {"references":["Denis J. R. 1930. Sur la faune francaise des Apterygotes, XIe note: Diploures avec tableau de determination des especes francaises. Bulletin de la Societe zoologique de France 55: 19 - 41."]}

Genus Kyrgyzstancampa Sendra & Ferreira gen. nov. urn:lsid:zoobank.org:act: 8647AF6F-06B9-4D69-A01B-89BF113788C5 Type species Kyrgyzstancampa sanare Sendra & Ferreira gen. et sp. nov. Diagnosis Sensilla of cupuliform organ paddle-shaped. Notal macrosetae pattern with 3+3 ma, la, and lp on pronotum and mesonotum. Femora with one dorsal macroseta and tibiae with one ventral macroseta. Claws subequal and regularly curved with ventral and lateral microspines. Laminar lateral processes striate on dorsal side with ridges surpassing apex and with short barbs on ventral side. Urotergites I– VII with up to 1+1 la and 2+2 lp macrosetae. First urosternite with 5+5 macrosetae, second to seventh urosternites with 3+3, and eighth urosternite with 1+1 macrosetae. First urosternite in males with glandular g 1, a 2 and a 1 setae; first urosternite in females with glandular a 1 setae. Etymology The genus name is a combination of ‘Kyrgyzstan’, the country where the material was found, and ‘campa’, a commonly applied suffix to dipluran generic names.

Turkmenocampa edaphica Sendra & S��nchez-Garc��a sp. nov. urn:lsid:zoobank.org:act: 6B28F41D-4B21-4E3C-A9B6-23FA7F8B0E17 Figs 27���31 Etymology The specific epithet refers to the habitat of the new species. Type material Holotype KYRGYZSTAN ��� ♀; ���holotype-♀07 MZB (MCNB) 2021-2340 ���; Jalal-Abad Region, Suzak Province, Kara Alma; 41��15���59��� N, 73��22���43��� E; alt. 1661 m; 16 Jul. 2019; A. Sendra leg.; endogean habitat near tree roots; MZB (MCNB) 2021-2340. Paratypes KYRGYZSTAN ��� 1 ♂, mounted in Marc Andr�� II solution; ���paratype-♂01 MZB (MCNB) 2021-2341���; same collection data as for holotype; MZB (MCNB) 2021-2341 ��� 1 ♂, mounted in Marc Andr�� II solution; ���paratype-♂02 Coll. AS���; same collection data as for preceding; Coll. AS ��� 1 ♀, mounted in Marc Andr�� II solution; ���paratype-♀05 Coll. AS���; same collection data as for preceding; Coll. AS ��� 1 ♀, mounted in Marc Andr�� II solution; ���paratype-♀08���; same collection data as for preceding; Coll. AS ��� 1 ♀, mounted in Marc Andr�� II solution; ���paratype-♀09 Coll. AS ���; same collection data as for preceding; Coll.AS ��� 1 ♀, mounted in Marc Andr�� II solution; ���paratype-♀10 MZB (MCNB) 2021-2342 paratype ���; same collection data as for preceding; MZB (MCNB) 2021-2342 ��� 1 ♀; ���♀01- paratype MCB (MCNB) 2021-2343 ���; Osh Province, Nookat District, Abshir Say River; 40��08���50��� N, 72��21���52��� E; alt. 1851 m; 21 Jul. 2019; A. Sendra leg.; endogean habitat near Cupressus tree; MCB (MCNB) 2021- 2343 ��� 1 ♀; ���paratype-♀02 MZB (MCNB) 2021-2344 ���; same collection data as for preceding; MZB (MCNB) 2021-2344 ��� 1 ♀; ���paratype-♀03 Coll AS ���; same collection data as for preceding; Coll AS ��� 1 ♀; ���paratype-♀04���; same collection data as for preceding; Coll. AS ��� 1 juv.; ���paratype-J01 Coll AS���; same collection data as for preceding; Coll AS ��� 1 juv.; ���paratype-J02 Coll AS���; same collection data as for preceding; Coll AS. Other material KYRGYZSTAN ��� 2 specs, mounted on an aluminium stage and coated with palladium-gold; same collection data as for holotype; Coll. AS. Description BODY. Length 3.0 and 3.1 mm in two males, 3.5���4.8 mm in nine females, 2.2 and 2.3 mm in two juveniles (Table 1). Epicuticle smooth under optical microscope but slightly reticulated at high magnifications with irregular polygonal structures of variable size. Body with short to middle-sized smooth clothing setae. HEAD. Antennae with 25���30 antennomeres in 10 complete intact antennae; antennae 0.6���0.8 �� length of body in adults and 0.9�� in juveniles (Table 1). Medial antennomeres 1.4�� as long as wide, apical antennomere 2.3�� as long as wide. Cupuliform organ with about ten oviform sensilla of types I and II and an unknown number of tree-shaped sensilla (type III) in this olfactory complex (Fig. 27). Distal and central antennomeres with five whorls of barbed macrosetae and scattered smooth setae, plus single distal whorl of about ten short, thin gouge sensilla 15���18 ��m long (Fig. 28). Proximal antennomeres with typical trichobothria, plus small and slightly shallow, 7 ��m long sensillum on 3 rd antennomere in ventral position. Plain frontal process with one frontal and two posterior macrosetae with length ratios a/p 45/30; three macrosetae along each side of insertion line of antennomere and setae x with thin distal barbs; length ratios a/i/p/x 28/38/27/ 30 in paratype ♀ 05 IBB-92102. Occiput of the head dorsally with 6+6 macrosetae, including 3+3 la, lp and mp macrosetae, longer than clothing setae and with few distal barbs. Large suboval labial palps each with microsensillae on the surface, small shallow latero-external sensillum, two guard setae, and up to ten clothing setae in anterior position, with up to 120 neuroglandular setae in medial and posterior positions, in holotype. THORAX. Thoracic macroseta distribution: pronotum and mesonotum with 1+1 ma, 1+1 la, 2+2 lp macrosetae; metanotum with 1+1 ma, 2+2 lp macrosetae. All macrosetae long and slightly thickened, with barbs along distal five-fourths of each seta; marginal setae up to twice as long as and thicker than clothing setae, well barbed near base (Fig. 29). Metathoracic legs reaching abdominal segment VI, about 0.3���0.4 �� as long as body length (Table 1). Femora II���III each with one long, thick dorsal macroseta with barbs along distal half and with two long ventral macrosetae. Calcars with long barbs along one side. Tibiae I���III with two short, thick ventral macrosetae with barbs along distal two-thirds. Two rows of ventral barbed setae. Three smooth, dorsal distal tarsal setae longer than rest. Subequal claws with large basal half with tiny dorsal spines and distal half curved and thinner. Laminar processes of pretarsus smooth on dorsal side and with long, thin, ending curved and with enlarged on ventral side. ABDOMEN. Distribution of abdominal macrosetae on tergites: 1+1 post 1 on I���II; 2+2 post 1���2 on III; 4+4 post 1���4 on IV���VII; 5+5 post 1���5 on VIII; and 7+7 post 1���7 on abdominal segment IX. All tergal abdominal macrosetae long, thick and short, with thin barbs along the distal fourth-fifths. Urosternite I with 8+8 macrosetae (Figs 30���31); urosternites II���VII with 4+4 macrosetae; urosternite VIII with 1+1 macrosetae; urosternal macrosetae of medium length or longer, with long barbs in single row along distal one-fourth to three-fourths. Stylus with apical seta, subapical seta and ventromedial seta with few long barbs arranged in one row along distal half to four-fifths. Cerci 0.6���0.85 �� length of body, with 5 and 7 primary articles, not counting multi-divided basal article (Table 1). Each primary article covered with unarranged whorls of barbed macrosetae and typical whorl of short setae with tiny distal barbs. SECONDARY SEX CHARACTERS. Female first urosternite with slightly thickened cylindrical appendages, each bearing microsensillae and 32���64 glandular a1 setae in a distal field (Table 1; Fig. 31). Male first urosternite with short subcylindrical appendages, each bearing microsensillae and 19���22 glandular a 1 setae in distal field (Table 1; Fig. 30). Type locality Kyrgyzstan, Kara Alma Village, Suzak Province, Jalal-Abad Region, 41��15���59��� N, 73��22���43��� E, 1661 m a.s.l. Habitat The morphological features and locations where Turkmenocampa edaphica Sendra & S��nchez-Garc��a sp. nov. has been found are congruent with those of a soil-dwelling species. In all cases, the species has been found in endogean habitats: under stones or among tree roots, always in humid places at 1661 m a.s.l. and 1851 m a.s.l., which is an average elevation for this mountainous country. Phyletic affinities Turkmenocampa edaphica Sendra & S��nchez-Garc��a sp. nov. is the second known species of a genus known previously from a troglobitic species inhabiting Kaptarhana Cave in Eastern Turkmenistan, Turkmenocampa mirabilis Sendra & Pavel, 2017, which is characterized by a Plusiocampinae pattern of macrosetae on the thorax and abdomen. Turkmenocampa also has a unique pretarsus consisting of subequal claws comprised of a large basal half with tiny dorsal spines, a thin, curved distal half and lateral laminar processes with long ventral barbs. Turkmenocampa mirabilis shows slight troglobiomorphic features: 30���32 antennomeres; up to twenty oviform sensilla on the cupuliform organ; gouge sensilla 18���26 ��m long; middle antennomeres 2���2.5�� as long as wide; legs slightly elongated; metathoracic legs reaching abdominal segment VIII; and much longer than wide appendages of the first urosternite, both in males and females (Sendra et al. 2017). However, T. edaphica Sendra & S��nchez-Garc��a sp. nov. shows body characters of a soil dweller: 25���30 antennomeres; up to ten oviform sensilla on the cupuliform organ; middle antennomeres 1.4�� as long as wide; gouge sensilla 15���18 ��m long; metathoracic legs reaching abdominal segment VI; and much longer than wide appendages of the first urosternite in both females and males. Turkmenocampa edaphica Sendra & S��nchez-Garc��a sp. nov. differs from T. mirabilis by the greater thickness of barbed marginal setae and by the greater number of glandular a 1 setae on the first urosternite appendages in females: with 32���64 in T. edaphica Sendra & S��nchez-Garc��a sp. nov., 12���21 in T. mirabilis., Published as part of Sendra, Alberto, S��nchez-Garc��a, Alba, Selfa, Jes��s, Milko, Dmitry A. & Ferreira, Rodrigo Lopes, 2021, Campodeidae (Hexapoda: Diplura) from Kyrgyzstan, Central Asia, with the description of a remarkable new genus and species, pp. 1-20 in European Journal of Taxonomy 782 on pages 13-17, DOI: 10.5852/ejt.2021.782.1585, http://zenodo.org/record/5761387, {"references":["Sendra A., Sket B. & Stoev P. 2017. A striking new genus and species of troglobitic Campodeidae (Diplura) from Central Asia. Subterranean Biology 23: 47 - 68. https: // doi. org / 10.3897 / subtbiol. 23.14631"]}

Kyrgyzstancampa sanare Sendra & Ferreira gen. et sp. nov. urn:lsid:zoobank.org:act: 012CDB29-579C-432F-B6C0-2DEE0F408BE5 Figs 8–26 Etymology The specific epithet is taken from the Latin ‘sanare’, meaning ‘cure’ and is related to the cave where the species was found, which is used for therapeutic purposes. This should be treated as a noun in apposition. Type material Holotype KYRGYZSTAN • ♀; “holotype- ♀ IBB 92101 ”; Ak-Turpak Cave; 40º10′35.18″ N, 71º03′45.36″ E; alt. 900 m; 12 Aug. 2019; R.L. Ferreira leg.; IBB 92101. Paratypes KYRGYZSTAN • 1 ♀, mounted in Marc André II solution; “paratype- ♀ 01 MZB (MCNB) 2021-2338”; same collection data as for holotype; MZB (MCNB) 2021-2338 • 1 ♀, mounted in Marc André II solution; “paratype- ♀ 02 Coll. AS”; same collection data as for preceding; Coll. AS • 1 ♂, mounted in Marc André II solution; “paratype-♂1 MZB (MCNB) 2021-2339”; same collection data as for preceding; MZB (MCNB) 2021-2339 • 1 ♂, mounted in Marc André II solution; “paratype-♂02 Coll. AS”; same collection data as for preceding; Coll. AS. Other material Two specimens with the same data as the holotype were mounted on an aluminium stage and coated with palladium-gold. Description BODY. Length 3–3.9 mm (3–3.9 mm in females; 3.1 and 3.4 mm in males; 3.9 mm in holotype) (Figs 8–9). Epicuticle smooth under optical microscope on dorsal side of nota and legs, but, at high magnification, slightly reticulate with irregular polygonal structures of variable size (Figs 16–17). Body sparsely covered with short clothing setae bearing 0–3 tiny distal barbs. HEAD. Two apparently intact antennae with 27–28 antennomeres; antennae 0.28–0.29 × length of body, with medial antennomeres 1.1× as long as wide; apical antennomere 1.9 × as long as wide. Cupuliform organ with about eight plain paddle-shaped olfactory chemoreceptor sensilla, 7 µm long (Fig. 10). Distal and medial antennomeres with two whorls of barbed macrosetae and scattered smooth setae, plus 2–4 short thin gouge sensilla 8–9 µm long (Figs 11–12). Proximal antennomeres with typical trichobothria, plus small bacilliform sensillum 6–7 µm long on 3 rd antennomere in ventral position (Figs 13–14). Plain frontal process with one anterior and three posterior smooth setae; length ratios of a / p 53/ 23 in holotype. Four short, smooth macrosetae along each side of antennomere insertion line with length ratios of a / i1 and i2 / p 11/15/14/ 11 in holotype; no x setae observed (Fig. 15). Small subtrapezoidal labial palp with small subcylindrical latero-external sensillum; two guard setae, up to three simple setae on anterior border, and up to 35 neuroglandular setae, as well as short and coniform palpiform sensillum, in holotype. THORAX. Thoracic macrosetae distribution (Figs 16–18): pronotum and mesonotum with 1+1 ma, 1+1 la, 1+1 lp macrosetae; metanotum with 1+1 ma macrosetae. All macrosetae rather slender with short barbs along middle third; marginal setae similar to clothing setae (Fig. 17). Legs short, metathoracic legs reaching abdominal segment V, about 0.3 × length of body (Fig. 19). Large, deep joint between femur and tibia with longitudinal protrusion on inner side (Fig. 20). Femora I–III each with one middle-sized dorsal macroseta with few distal barbs, slightly longer than ventral macroseta. Calcars slightly thickened with long barbs on one side. Tibiae I‒III with one ventral macroseta with three or four distal barbs. Two rows of ventral barbed setae longer and thicker than clothing setae, with long thin barbs. Three smooth, distal dorsal tarsal setae longer than rest. Claws subequal, regularly curved, with tiny ventral and lateral microspines. Laminar lateral processes of pretarsus striated on dorsal side with ridges surpassing end of the apex, giving appearance of distal fringe, and with short barbs on ventral side (Figs 21–24). ABDOMEN. Distribution of abdominal macrosetae on tergites: 1+1 lp on urotergite III; 1+1 la, 2+2 lp on urotergites IV–VIII; 3+3 lp on abdominal segment IX; 5+5 macrosetae on abdominal segment X; all macrosetae long, with thin barbs along distal half. Urosternite I apparently with 5+5 macrosetae (Figs 25–26); urosternites II–VII with 3+3 macrosetae; urosternite VIII with 1+1 macrosetae; longsized urosternal macrosetae with few distal barbs. Stylus with apical seta with two long basal teeth, subapical seta and ventromedial seta, each bearing a row of barbs along distal half, more abundant on ventromedial setae. CERCI. 0.71 × length of body (on a cercus apparently intact in the holotype), with basal article divided into four secondary articles plus 11 primary articles; each primary article with central constriction bearing whorl of long macrosetae with thin barbs on distal part and one or two whorls of thin smooth setae; each primary article ending in whorl of thin setae, including apical article. SECONDARY SEX CHARACTERS. Female urosternite I with short subcylindrical appendages, each bearing up to 11–13 glandular a 1 setae in distal field (Fig. 27). Male urosternite I with elongated subtrapezoidal appendages, each bearing up to 8 glandular a 1 setae in distal field and larger posterior field with up to 70 glandular a2 setae; posterior edge of first urosternite with field of up to 44 glandular g1 setae arranged in two rows (Fig. 25). Type locality Kyrgyzstan, Kadamjay District, Batken Region, Ak-Turpak Cave, gypsum cave located south of Ak-Turpak village; 40º10′35.18″ N, 71º03′45.36″ E. Habitat The specimens were observed only in the deep zone of Ak-Turpak Cave, located near the western margin of the Kadamjay District, Batken Province, Kyrgyzstan, which is located about 2.5 km south of the village of Ak-Turpak (northwestern part of Alai Mts.). The name of the locality means ʻwhite landʼ in the local Turkish dialect and reflects the prevalence of the whitish, pinkish, or reddish clayey ground surface. Its entrance is located about 400 m from the right bank of the river Sokh (Kozheshken) (Fig. 2), approximately 40–50 m a.s.l. The Sokh River divides the northern macroslopes of the Turkestan Mt Range and Alai (or Alay) Mt System. This area can also be considered as the southern edge of the Fergana Depression. The cave entrance is surrounded by a hilly relief, without any tops above 1000 m a.s.l. in a one-kilometre-neighbourhood. The landscape surrounding the Kyzyl-Unkuyr Cave is quite dry (Figs 2−3), with only sparse shrubby vegetation typical of rocky outcrops, where the soil is extremely shallow when present. On the other hand, the Sokh River floodplain, located quite close to the cave, is moist although it is currently very altered due to the presence of crops and small villages. However, suitable habitats for soil invertebrates certainly occur along this floodplain. It is worth mentioning that although Kyrgyzstancampa sanare Sendra & Ferreira gen. et sp. nov. was found in a cave, it does not show any troglomorphic morphological characters. Thus, it is likely that the species is not troglobitic, although further sampling in the external area surrounding the cave (especially along the floodplain of the Sokh River) is needed to confirm this hypothesis. The Ak-Turpak cave has a single entrance, where a metallic structure was installed to safeguard and protect the cave’s entrance (Figs 3−4). From the entrance inwards, stairs were built to facilitate access for visitors. The cave gallery is comparatively simple and oriented east-northeast, with 137 m of linear extension and about 40 m deep (Gvozdetskij 1981; Dudashvili & Mikhailyov 1990). The area of the cave was estimated to be 2400 m 2 and the volume is 8393 m 3 (Mamatkulov 1978). The cave is situated in a gypsum stratum (Gvozdetskij 1981) in the trough zone, where karstified rocks are represented by gypsum, marls, marlstones, limestones, and dolomites of Cretaceous and Paleogene ages (Beloglazova & Smirnova 1987; Sultanov 1972). The origin of all karst forms in Southern Fergana is related to tectonic faults and sedimentary breccias, and they often developed as a result of repeated and sometimes overlapping karst processes (Sultanov 1972). In the upper part of the cave conduit, there is a noticeable proportion of soft marl that is somewhat dilapidated (during the last 5–10 years, this part of the gallery was equipped with a cement staircase and the walls were partly reinforced with rubble masonry panels to reduce dust and for balneological and recreational use). In the deeper parts of the gallery, the cave vaults are formed by fine-crystalline selenite (calcium sulphate dihydrate CaSO4•2H2O) of several, sometimes contrasting, colour shades. The north side of the cave is preferentially formed by argillite. The atmosphere of the Ak-Turpak cave is rather dry and there are no traces of thermokarst processes (Dudashvili & Mikhailyov 1990); however, the cave vaults are somewhat crumbled after recent earthquakes. Over the last decades, local residents (≈ 100–330 per year) have used the cave for therapeutic purposes (respiratory treatments: asthma, bronchitis, etc.) as word of mouth on the cave’s ʻhealing propertiesʼ has spread among them. Hence, one can see, especially in the entrance, small platforms and mattresses (Fig. 5). Visitors mostly use the entrance area, but stairs have also been installed deep inside the cave (Fig. 6), in which some mattresses were observed, indicating that the entire cave has been used for therapeutic purposes. Specimens of Kyrgyzstancampa sanare Sendra & Ferreira gen. et sp. nov. were only found in the last chamber of the cave, and always associated with old bat guano (Fig. 7). Several individuals were observed amidst the guano (Figs 8−9), rapidly escaping when disturbed. In these cases, they tended to enter the small spaces between the chitin fragments observed in the pile, so it was difficult to capture specimens without injuring them. The only organic resource observed inside the cave was bat guano from species of Rhinolophus Lacépède, 1799 (horseshoe bats) and a few organic materials left by visitors (such as cardboard pieces and wood). The cave is not well preserved as many accesses were built, thus deeply altering the pristine substrates. However, considering the lack of troglomorphic traits in the species (indicating that the cave is not its unique habitat) and given that apparently few visitors access the deeper parts of the cave, the species does not appear to be threatened. Phyletic affinities Kyrgyzstancampa Sendra & Ferreira gen. nov. has similarities with several species of the paraphyletic genus Eutrichocampa Silvestri, 1902. In his diagnosis of Eutrichocampa, the tarsus ends abruptly instead of being acuminate towards the apex, which he considered to be a feature differentiating Eutrichocampa and Campodea (Silvestri 1902). For more than a century, several authors have been adding species to this genus, such as Wygodzinsky (1941, 1943), Condé (1947, 1994), Ionsecu (1955), Loksa (1960), García-Gómez (2016) and also Silvestri (1931a, 1932a, 1932b, 1933a), resulting in the current fifteen species of Eutrichocampa (Sendra et al. 2021). These species were described from localities scattered in the Americas, Africa, Asia, and Europe. In all of these contributions, the entire pretarsus shape is referred to as the differential character for Eutrichocampa: regularly curved claws with laminar or subcylindrical lateral processes with abundant barbs. Since Wygodzinsky (1941), Eutrichocampa has been considered a heterogeneous genus showing a wide variation in macrosetal patterns on the thorax and abdomen, including the presence or absence of dorsal macrosetae on the femora. In spite of the effort made by Condé (1956) to keep Eutrichocampa as a homogeneous taxon, several authors (including Condé himself) have tried to arrange it into several genera and subgenera (Paclt 1957), proposing other genera with the same pretarsus trait and thoracic macrosetae of the Campodea pattern; for instance, Parallocampa Silvestri, 1933b with eleven species from North America, and Remycampa Condé, 1952, with two species from northwest Africa and the Canary Islands, and four monotypic genera: Allocampa Silvestri, 1931b from Cuba; Edriocampa Silvestri, 1933a from the South Aegean islands and Anatolian Peninsula; Libanocampa Condé, 1955 from Lebanon and Anatolia; and Pseudolibanocampa Xie & Yang, 1991 from Guangdong and Yunnan in China. In 1957, Paclt proposed an artificial arrangement of Eutrichocampa by dividing it into four subgenera and the genus Leniwytsmania Paclt, 1957 for two species, both from China: L. orientalis (Silvestri, 1931a) and L. helvetica (Wygodzinsky, 1941). Our proposal of Kyrgyzstancampa Sendra & Ferreira gen. nov. is another effort to unravel the diversity within the subfamily Campodeinae, in which this new genus can be included. Several characters define Kyrgyzstancampa Sendra & Ferreira gen. nov., such as the pretarsus with a regularly curved claw with tiny ventral and lateral microspines; the laminar lateral processes, striated on the dorsal side with short barbs on the ventral side (Figs 21–24); a unique femur-tibia joint; the macroseta pattern on the nota: 3+3 ma, la, lp macrosetae on the pronotum and metanotum, 2+2 ma, lp on the metanotum plus one dorsal femoral macroseta; and 1+1 la and 2+2 lp on urotergites IV–VIII. Other notable features are the sparse clothing setae on the body, the plain paddle-shaped sensilla on the cupuliform organ, and the secondary sexual characters on the first urosternite. This combination of characters delineates Kyrgyzstancampa Sendra & Ferreira gen. nov. from other genera of Campodeinae and all species of Eutrichocampa. The closest species to K. sanare Sendra & Ferreira gen. et sp. nov. seems to be Eutrichocampa birabei Wygodzinsky, 1943, described from San Antonio de Arredondo, Córdoba in Argentina. Both share the shape of the pretarsus, the distribution of macrosetae on the nota and urotergites, and a dorsal macroseta on the femora. However, K. sanare Sendra & Ferreira gen. et sp. nov. and E. birabei differ in the number of urotergal macrosetae and in the secondary sexual characters of the first urosternite. Furthermore, reuniting both species in Kyrgyzstancampa Sendra & Ferreira gen. nov. would be a far-fetched approach, and new material on the South American species will be necessary to provide a more accurate description.

Plusiocampa (Plusiocampa) imereti Sendra & Barjadze sp. nov. urn:lsid:zoobank.org:act: 40C3DB96-19CD-4354-ACA9-434F0ADA34BB Figs 2���12 Diagnosis Troglomorphic species. Antennae with 39���45 antennomeres; 12 complex olfactory chemoreceptors within cupuliform organ; non-protruding frontal process slightly protruding, plain, with non-tubercular setae or just slightly tubercular. Pronotum 1+1 ma, 2+2 la 1,2 , 2+2 lp 2,3 ; mesonotum with 1+1 ma, 3+3 la 1���3 , 2+2 lp 2,3 , 1+1 mp; metanotum with 1+1 ma, 2+2 la 1,2 or sla 1,2 , 1+1 lp 2 , 1+1 mp; thin; all notal macrosetae long and covered by thin barbs on half to distal portions; thin, middle-sized clothing setae covered by 0���4 thin distal barbs. Legs elongated, pretarsus of metathoracic legs slightly overpasses end of abdomen. Femora I���III with one long, barbed dorsal macroseta and one shorter, barbed ventral macroseta. Tibiae I���III with two short barbed ventral macrosetae. Claws unequal (posterior claw 1.3 �� as long as anterior one); large, backward overhang on posterior claw; lateral crests well-developed. Pretarsal process long and setiform. Urotergites 1+1 post 1 on I���II; 0+0, 0+1 or 1+1 la, 1+1 or 2+2 post 1,2 on III; 1+1 la 3 , 2+2 to 4+4 post 1���4 on IV; 2+2 la 2,3 , 4+4 post 1���4 on V���VII; 6+6 post 1���6 on VIII and 8+8 or 8+7 post 1���8 on abdominal IX. Urosternite I with 8+8���7+7 macrosetae (Figs 8���9); urosternites II���VII with 6+6 macrosetae; urosternite VIII with 2+2 macrosetae. Male urosternite I (Fig. 8) with slightly enlarged subcylindrical appendages, each bearing up to 21 glandular a 1 setae. Female appendages slightly thinner, with up to 11 glandular a 1 setae. Etymology The specific epithet refers to the Imereti region, the location of the Shvilobisa Cave, treated as a noun in apposition. Type material Holotype GEORGIA ��� ♀, ���holotype-♀ IZISU-TD-T-00001���; Shvilobisa Cave, Bunikauri village, Chiatura Municipality, Imereti region, Zemo Imereti Plateau; 42��19���31.44��� N, 43��16���4.33��� E; 24 Feb. 2018; Shalva Barjadze and Eter Maghradze leg.; IZISU-TD-T-00001. Paratypes GEORGIA ��� 1 ♂, ���paratype-♂1 IZISU-TD-T-00002���; same collection data as for holotype; IZISU- TD-T-00002 ��� 1 ♀, ���paratype-♀1 MZB (MCNB) 2021-2336���; same locality as for holotype; 20 Jul. 2020; Eter Maghradze leg.; MZB (MCNB) 2021-2336 ��� 1 ♀, ���paratype-♀2 MZB (MCNB) 2021-2337���; same collection data as for preceding; MZB (MCNB) 2021-2337 ��� 1 ♂, ���paratype-♂2 Coll AS���; same collection data as for preceding; Coll AS. Other material GEORGIA ��� 2 specs, unknown sex [for SEM photography and one specimen for DNA analysis]; Shvilobisa Cave; 24 Feb. 2018; Shalva Barjadze and Eter Maghradze leg. ��� 2 specs [for SEM photography and one for DNA analysis]; same collection data as for preceding; 20 Jul. 2020; Eter Maghradze leg. Other material from two other caves (all Coll AS) GEORGIA ��� 1 ♀; Kumistavi village Tskaltubo Municipality, Imereti Region, Sataplia-Tskaltubo karst massif, Datvis (Bear) Cave; 42��22���28������ N, 42��35���45��� E; 5 Jul. 2018; Eter Maghradze leg. ��� 1 ♀; same collection data as for preceding; 1 Sept. 2019 ��� 1 ♂; near Melouri village, Tskaltubo Municipality, Imereti Region, Sataplia-Tskaltubo karst massif, Melouri Cave; 42��23���15.1��� N, 42��37���41.5��� E; 1 Nov. 2018; Eter Maghradze leg. Description BODY. Body length 4.3���7.2 mm (females) and 4.9���5.2 mm (males) (Table 1). Epicuticle smooth under optical microscope and SEM; body with thin, middle-sized clothing setae covered by 0���4 thin distal barbs. HEAD. Three intact antennae, all slightly longer than body length, with 39���45 antennomeres (Table 1). Small, thin, subcylindrical sensillum on third antennomere located in ventral position between c and d macrosetae. Central antennomeres 2.1 �� as long as wide, apical antennomere 3.0 �� as long as wide. Cupuliform organ occupying ⅓ of total length of apical antennomere, with about 12 complex olfactory chemoreceptors. Each olfactory chemoreceptor is composed of a complete fold surrounding a central cylinder with two lateral expansions, entirely reticulated and perforated (Fig. 5). Gouge sensilla 30��� 40 ��m long, in a single distal whorl of 13���16 sensilla on each medial and distal antennomere. Frontal process slightly protruding, plain, with non-tubercular setae or just slightly tubercular on distal portion (Fig. 3); macrosetae along the insertion line of antennomere and i macrosetae and x setae longer than other macrosetae (a / i / p/ x with relative lengths of 25 / 36 / 19 / 37 in holotype). Suboval labial palps with a bacilliform latero-external sensillum, two guard setae, up to 7 setae on anterior border, and up to 130 neuroglandular setae in holotype. THORAX. Thoracic macrosetal distribution (Figs 2, 4, 6): pronotum with 1+1 ma, 2+2 la 1���2 , 2+2 lp 2,3 ; mesonotum with 1+1 ma, 3+3 la 1���3 , 2+2 lp 2,3 , 1+1 mp; metanotum with 1+1 ma, 2+2 la 1,2 or sla 1,2 , 1+1 lp 2 , 1+1 mp. All notal macrosetae are long and covered by thin barbs on half to distal portions (Figs 4, 6); submacrosetae sla are thinner and shorter than notal macrosetae; marginal setae are similar to clothing setae, and covered by 1���8 thin distal barbs. Legs elongated, pretarsus of metathoracic legs slightly overpasses end of abdomen (Table 1). Femora I���III with one long, barbed dorsal macroseta and one shorter, barbed ventral macroseta. Tibiae I���III with two short barbed ventral macrosetae. Calcars with 4���5 long barbs. Tarsi with two rows of thicker ventral setae with 2���3 very thin barbs on middle portion. Two dorsal and one ventral, smooth, subapical tarsal setae. Claws are unequal (posterior claw 1.3 �� as long as anterior one); large, backward overhang on posterior claw; lateral crests well-developed. Pretarsal process long and setiform, overpassing end of claws. ABDOMEN. Distribution of abdominal macrosetae on tergites (Fig. 7): 1+1 post 1 on I���II; 0+0, 0+1 or 1+1 la, 1+1 or 2+2 post 1,2 on III; 1+1 la 3 , 2+2 to 4+4 post 1���4 on IV; 2+2 la 2,3 , 4+4 post 1���4 on V���VII; 6+6 post 1���6 on VIII and 8+8 or 8+7 post 1���8 on abdominal segment IX. All post urotergal macrosetae long and covered by thin barbs along distal four-fifths (Fig. 10); la urotergal macrosetae shorter than post macrosetae, covered by barbs along distal half. Urosternite I with 8+8���7+7 macrosetae (Figs 8���9); urosternites II���VII with 6+6 macrosetae; urosternite VIII with 2+2 macrosetae (Fig. 12); all urosternal macrosetae robust and large, covered by long barbs along distal third to four-fifths. Apical, subapical and ventromedial setae with a few (two to four) thin, short and long barbs (Fig. 11). SECONDARY SEX CHARACTERS. Male urosternite I (Fig. 8) with slightly enlarged subcylindrical appendages, each bearing up to 21 glandular a1 setae. Female appendages slightly thinner, with up to 11 glandular a 1 setae. Spermatozoid fascicles 40 ��m in diameter without apparently spiral filament. Molecular analysis The nucleotide substitution model selected was GTR+G+I (BIC = 6998.6), with the proportion of invariant sites (I = 0.46) and estimated alpha parameter for the gamma distribution (�� = 1.39), indicating a significant heterogeneity in the DNA substitution among sites. The Campodeidae sequences formed a well-supported clade, clearly distinct from that of Japygidae (Fig. 13). Although bootstrap values were low, the ML phylogenetic tree grouped Plusiocampa (P.) imereti Sendra & Barjadze sp. nov. with Eastern Europe taxa such as Plusiocampa (Plusiocampa) aff. elongata Ionescu, 1955 and Plusiocampa (Plusiocampa) humicola Ionescu, 1955, whereas Iberian Peninsula taxa (Plusiocampa (Plusiocampa) gadorensis Sendra, 2001, Plusiocampa (Plusiocampa) baetica Sendra, 2004 and Cestocampa iberica Sendra & Cond��, 2012) clustered in a distinct clade. K2P genetic distances also showed P. (P.) aff. elongata (0.206�� 0.027) and P. (P.) humicola (0.205 �� 0.028) to be the closest species to the new Plusiocampa (P.) imereti Sendra & Barjadze sp. nov. from Georgia. Habitat Plusiocampa (Plusiocampa) imereti Sendra & Barjadze sp. nov. inhabits the deep zone (over 50 m from the entrance) of three caves. The Shvilobisa Cave, the type locality, is a 1000 m long, tunnellike, easily accessible sub-horizontal cave with a small subterranean water stream (Tatashidze et al. 2009b). The others two nearby caves are about 55 km away from the Shvilobisa Cave; the Melouri Cave is 5300 meters long and has the status of natural monument (Tatashidze et al. 2009b), whereas the Datvis Cave is a poorly known cavern (K. Tsikarishvili, pers. comm.). The distance between the Datvis and Melouri caves is ca 3.5 km. The Melouri Cave ��� easily accessible ��� has dried halls and a permanent subterranean water stream near its end. This cave has gigantic stalagmites and fallen stones. The Datvis Cave is a horizontal, dry, and easily accessible cave with several halls, which are rich in different speleothems like the Shvilobisa Cave (Tatashidze et al. 2009b). Invertebrate cavernicolous species of the studied caves The three caves (Shvilobisa, Datvis, and Melouri) which Plusiocampa (Plusiocampa) imereti Sendra & Barjadze sp. nov. inhabits are also the dwellings of other troglobitic arthropod species. Datvis and Melouri caves share three troglobitic species: the Diplopoda Leucogeorgia prometheus Antić & Reip, 2020, the Isopoda Colchidoniscus kutaissianus Borutzky, 1974, and the Pseudoscorpionida Chthonius satapliaensis Schawaller & Dashdamirov, 1988. In addition, the Insecta (Carabidae Coleoptera) Troglocimmerites imereti Dolzhanski & Ljovuschkin, 1985 dwells in the Datvis cave; and the Melouri Cave has five more species: the Opiliones Nemaspela melouri Martens, Maghradze & Barjadze, 2021, the Araneae Centromerus bulgarianus Drensky, 1931, the Hexapoda Collembola Pseudacherontides zenkevitchi Djanashvili, 1971; the Insecta (Carabidae Coleoptera) Inotrechus kurnakovi Dolzhanski & Ljovuschkin, 1989; and Troglocimmerites sp. 1. In the Shvilobisa Cave, eight troglobitic species dwell: the Diplopoda Leucogeorgia gioi Antić & Reip, 2020, the Isopoda Caucasonethes cf. borutzkyi Verhoeff, 1932 and Colchidoniscus sp., the Pseudoscorpionida Chthonius satapliaensis Schawaller & Dashdamirov, 1988, the Opiliones Nemaspela sp., the Hexapoda Collembola Oncopodura sp. and Pseudosinella sp., and the Insecta (Carabidae Coleoptera) Troglocimmerites sp. 2. (Barjadze et al. 2019; Maghradze et al. 2019; Antić & Reip 2020; Martens et al. 2021; Maghradze & Barjadze, unpublished data)., Published as part of Sendra, Alberto, Palero, Ferran, S��nchez-Garc��a, Alba, Jim��nez-Valverde, Alberto, Selfa, Jes��s, Maghradze, Eter & Barjadze, Shalva, 2021, A new Diplura species from Georgia caves, Plusiocampa (Plusiocampa) imereti (Diplura, Campodeidae), with morphological and molecular data, pp. 71-85 in European Journal of Taxonomy 778 on pages 74-79, DOI: 10.5852/ejt.2021.778.1567, http://zenodo.org/record/5675209, {"references":["Ionescu M. A. 1955. Diplura. In: Academia Republicii Socialiste Romania (ed.) Fauna Republicii Populare Romane, Insecta 7 (2): 1 - 48.","Sendra A., Lara M. D., Ruiz Aviles F. & Tinaut A. 2004. Une nouvelle espece du genre Plusiocampa Silvestri, 1912 (Diplura, Campodeidae) et donnees pour sa reconstruction paleobiogeographique dans les Betiques. Subterranean Biology 2: 113 - 122.","Tatashidze Z. K., Tsikarishvili K. D. & Jishkariani J. M. 2009 b. The Cadastre of the Karst Caves of Georgia. Petiti Publishing House, Tbilisi [in Georgian].","Antic D. Z & Reip H. R. 2020. The millipede genus Leucogeorgia Verhoeff, 1930 in the Caucasus, with descriptions of eleven new species, erection of a new monotypic genus and notes on the tribe Leucogeorgiini (Diplopoda: Julida: Julidae). European Journal of Taxonomy 713: 1 - 106. https: // doi. org / 10.5852 / ejt. 2020.713","Martens J., Maghradze E. & Barjadze Sh. 2021. Two new species of the genus Nemaspela Silhavy from caves in Georgia (Opiliones: Nemastomatidae). Zootaxa 4951 (3): 541 - 558. https: // doi. org / 10.11646 / zootaxa. 4951.3.7","Barjadze Sh., Arabuli T., Mumladze L., Maghradze E., Asanidze Z. & Shavadze R. 2019. Cave Biodiversity of Georgia, Open Access Database. Institute of Zoology at Ilia State University. Available from https: // cbg. iliauni. edu. ge / en / [accessed: 20 Apr. 2021].","Maghradze E., Faille A., Barjadze Sh. & Hlavac P. 2019. A new cavernicolous species of the genus Bergrothia Reitter, 1884 (Coleoptera, Staphylinidae, Pselaphinae) from Georgia. Zootaxa 4608 (2): 371 - 379. https: // doi. org / 10.11646 / zootaxa. 4608.2.11"]}

Autrigoniscus, new genus TYPE SPECIES: Autrigoniscus resinicola, new species. DIAGNOSIS: Male. An oniscidean with the following combination of characters: Body size small (Female. Unknown. ETYMOLOGY: The new genus group name is a combination of the pre-Roman tribe Autrigones (who lived in northern Spain near the region of the amber localities along with the Cantabri) and the Greek suffix – iskos, denoting a diminutive. The gender of the name is masculine., Published as part of Sánchez-García, Alba, Peñalver, Enrique, Delclòs, Xavier & Engel, Michael S., 2021, Terrestrial Isopods from Spanish Amber (Crustacea: Oniscidea): Insights into the Cretaceous Soil Biota, pp. 1-32 in American Museum Novitates 2021 (3974) on page 13, DOI: 10.1206/3974.1, http://zenodo.org/record/5356138

Heraclitus helenae, new species Figures 10, 11 DIAGNOSIS: As for the genus (above). DESCRIPTION: Body (fig. 10) more or less oblong, of “clinger” habitus; small sized, total length 1.76 mm, width not measurable due to orientation in amber; dorsal surface convex, rough, covered with large semicircular scales and densely crowded prominent triangularshaped scale setae arranged in transverse rows; gland pores not visible as preserved. Cephalon rounded, rather arcuate, rough, with same ornamentation as described for body (fig. 11B); lateral lobes not visible as preserved (probably small); antennal lobes absent. Eyes not visible as preserved. Pereon about 0.73× of body length; pereonites overlapping, lamellar; lateral parts of pereonites (i.e., epimeral plates) only moderately prominent, those of two posterior pereonites with posterior corners acutely triangular, all edged with small appressed spikes and distinctly prominent triangular-shaped scale setae (as a continuity to those of dorsal surface). Pleon about 0.24× of body length, distinctly narrower than pereon (abruptly contracted); pleotelson short, subtriangular, with rounded apex bearing two short distal setae. Mouthparts considerably well developed but without visible details. Antennule poorly visible; distal article with three short thick spines distally. Antenna (fig. 11A) very spinose in appearance, packed with small scale setae and spines, rather stout, with five peduncular articles and a multiarticulate flagellum; articles 1–3 partly obscured by head, apparently short and stout; fourth article elongate, stout, 0.18 mm long, 2.25× longer than broad; fifth article elongate and much slender than fourth article, 0.22 mm long, 1.22× length of preceding article, 5.5× longer than broad; flagellum partially preserved, with at least three articles not clearly delimited. Pereopods all alike, of moderate size; all pereopods without sexual modifications; basis and ischium mostly obscured by body; merus, carpus, and propodus with elongate, thick, nearly straight spines generally arranged in a longitudinal row on sternal margins giving them a rippled appearance; propodus and carpus length apparently subequal; dactylus with enlarged outer claw and minute inner claw; dactylar setae unbranched, glabrous, long, and slender, distinctly expanded distally; ungual setae not visible as preserved. Pleopods not visible as preserved. Uropod (fig. 11C) moderately elongate, about 0.28× of body length as preserved, freely projecting caudally; protopod produced inside pleotelson, subrectangular, long and stout, 0.15 mm long, with endopod and exopod inserted at same level; exopod styliform, long and slender, width not measurable due to poor preservation, 0.35 mm long as preserved (distal part not preserved), distinctly longer than endopod, with some very short thin setae sparsely distributed; endopod conical, maximum width at base 0.03 mm, tapering distally, 0.21 mm long (excluding terminal tuft of setae), with some very short thin setae sparsely distributed, and a tuft of long setae apically (0.10 mm long). HOLOTYPE: MCNA 12546, sex unknown, exposed dorsolaterally, ventrolaterally, and laterally. Preserved in darkened orange-colored amber, trimmed to 3.5×1.7× 1.2 mm (in an epoxy trapezoid 23.0×15.0× 1.5 mm), surrounded by numerous pseudoinclusions (Lozano et al., 2020) and particles of detritus; no other major inclusions. The specimen is nearly complete, but some areas are poorly visible owing to the position of the animal during fossilization. The distal parts of the uropodal endopod and the antennulae are not preserved. OCCURRENCE: Peñacerrada I amber site [Peñacerrada I = Moraza], eastern margin of the Basque-Cantabrian Basin, Burgos, northern Spain; Early Cretaceous (Late Albian). ETYMOLOGY: The specific epithet refers to Helen of Troy, a figure from Greek mythology known for her considerable beauty and having “launched a thousand ships” (à la Marlowe) as Menelaus and Agamemnon initiated the Trojan War to return her to Achaea. REMARKS: Heraclitus helenae may be clearly differentiated from A. resinicola by the roughened cuticle, the spinose antennulae, the dactylus with two claws, and the longer uropodal endopod. It is perhaps a representative of Crinocheta: Detonidae? owing to the antennule with a 5-jointed peduncle and a 4-jointed flagellum, the antennulae with conspicuous scaly tubercles, the inner claw small, the uropodal protopod surpassing the pleotelson, the exopodites leaf shaped, and the endopodites styliform (Schmidt, 2002). Oniscidea Indet. SPECIMEN MCNA 14907: (fig. 12A) is preserved in darkened orange-colored amber, trimmed to 10.5×6.0× 1.5 mm (in an epoxy trapezoid 21.1×13.5× 2.2 mm), and surrounded by numerous pseudoinclusions (Lozano et al., 2020), particles of detritus, fungal hyphae, and three coprolites. The specimen is exposed dorsally and ventrally, poorly preserved, with the cuticle virtually degraded except for the lateral parts of the segments, which are edged with small appressed spikes and some small triangular-shaped scale setae. Worthy of note are the oval-shaped medium-sized body (total length 4.27 mm as preserved, maximum width 2.27 mm), and the moderately large eyes composed of multiple ommatidia (only the left eye is visible). The specimen also possesses a partly preserved antenna (cut just below the articulation between the fourth and fifth articles) that is elbowed between the fourth and fifth articles. The first peduncular article is partly concealed by the head, whereas the measurements of the rest of articles are as follows: second article 0.22 mm long, about as long as broad; third article 0.25 mm long, 1.25× longer than broad, with one long, thick spine; fourth article elongate, 0.58 mm long, 4.83× longer than broad. Only some anterior pereopods are visible, but these are so thoroughly obscured owing to preservation as to provide no useful delineation of features. However, long, thick, nearly straight spines, a dactylus with two claws (the inner claw minute, distinctly smaller than in H. helenae), and an unbranched, glabrous, long, and slender dactylar seta are visible on some pereopods. The specimen is difficult to place accurately among Oniscidea, but placement within the family Ligiidae seems most appropriate owing to the presence of large eyes composed of multiple ommatidia, the comparatively elongate antennae, and a dactylus with two claws. Interestingly, the specimen corresponds with the anterior half of an exuvia. Isopods perform a biphasic molt, which consists of the shedding of the posterior and then the anterior half of the body. The boundary between the two halves is between the fourth and fifth pereonites, which agrees with that observed in the fossil. SPECIMEN MCNA 9458: (fig. 12B) is preserved in darkened yellow-colored amber, trimmed to 5.2×3.5×1.0 mm (in an epoxy trapezoid 19.1×14.1× 1.1 mm), and surrounded by numerous pseudoinclusions and particles of detritus; the amber is also darkened near the inclusion. The specimen is exposed dorsally and ventrally, and although a vague outline of the pereopods armed with long, stout, nearly straight spines can be seen, these are so thoroughly obscured owing to preservation and by the adjacent body mass as to provide no useful delineation of features. It possesses a more or less oblong, small body (total length 2.09 mm long as preserved, maximum width 0.87 mm), and some segments of the pereon with some small triangular-shaped scale setae. SPECIMEN MCNA 9924.2: (fig. 13; previously figured in Delclòs et al., 2007, Broly et al., 2013, and Sánchez-García et al., 2015) is preserved in darkened orange-colored amber, trimmed to 18.0×14.5× 3.5 mm (in an epoxy trapezoid 22.5×15.0× 4.8 mm). Syninclusions include one acariform Bdellidae, one Blattodea, six Diptera (one Ceratopogonidae, two Dolichopodidae, one Phoridae, two Psychodidae), one Tanaidacea (Alavatanais margulisae Sánchez-García, Peñalver, and Delclòs), two Archaeognatha, six coprolites (attributed to termites based on their general shape), and abundant fungal and plant remains (see details of syninclusions in Sánchez-García et al., 2015). The specimen is exposed laterally and partially complete (missing head), with the right side of the body and the right pereopods mostly polished off. The specimen possesses a medium-sized body (total length 3.78 mm as preserved), more or less oblong, only moderately convex dorsally. The pereon is about 0.73× the body length, with the lateral parts of the pereonites only moderately prominent, those of the four posterior segments with posterior corners acutely triangular, all edged with some small appressed spikes. The pleon is about 0.22× of the body length and distinctly narrower than the pereon (abruptly contracted). It also possesses spinous pereopods, a dactylus with two claws (the inner claw minute), and an unbranched, glabrous, long, and slender dactylar seta, distinctly expanded distally. Lastly, an elongate structure that could correspond to a short uropod is sticking out from under the pleon., Published as part of Sánchez-García, Alba, Peñalver, Enrique, Delclòs, Xavier & Engel, Michael S., 2021, Terrestrial Isopods from Spanish Amber (Crustacea: Oniscidea): Insights into the Cretaceous Soil Biota, pp. 1-32 in American Museum Novitates 2021 (3974) on pages 21-25, DOI: 10.1206/3974.1, http://zenodo.org/record/5356138, {"references":["Lozano, R. P., et al. 2020. Phloem sap in Cretaceous ambers as abundant double emulsions preserving organic and inorganic residues. Scientific Reports 10: 9751 [1 - 15].","Schmidt, C. 2002. Contribution to the phylogenetic system of the Crinocheta (Crustacea, Isopoda). Part 1. (Olibrinidae to Scyphaidae s. str.). Mitteilungen aus dem Museum fur Naturkunde in Berlin, Zoologische Reihe 78 (2): 275 - 352.","Delclos, X., et al. 2007. Fossiliferous amber deposits from the Cretaceous (Albian) of Spain. Comptes Rendus Palevol 6 (1 - 2): 135 - 149.","Broly, P., P. Deville, and S. Maillet. 2013. The origin of terrestrial isopods (Crustacea: Isopoda: Oniscidea). Evolutionary Ecology 27: 461 - 476.","Sanchez-Garcia, A., E. Penalver, R. Perez-de la Fuente, and X. Delclos. 2015. A rich and diverse tanaidomorphan (Crustacea: Tanaidacea) assemblage associated with Early Cretaceous resin-producing forests in North Iberia: palaeobiological implications. Journal of Systematic Palaeontology 13 (8): 645 - 676."]}

Autrigoniscus, new genus TYPE SPECIES: Autrigoniscus resinicola, new species. DIAGNOSIS: Male. An oniscidean with the following combination of characters: Body size small (Female. Unknown. ETYMOLOGY: The new genus group name is a combination of the pre-Roman tribe Autrigones (who lived in northern Spain near the region of the amber localities along with the Cantabri) and the Greek suffix – iskos, denoting a diminutive. The gender of the name is masculine.

Heraclitus helenae, new species Figures 10, 11 DIAGNOSIS: As for the genus (above). DESCRIPTION: Body (fig. 10) more or less oblong, of “clinger” habitus; small sized, total length 1.76 mm, width not measurable due to orientation in amber; dorsal surface convex, rough, covered with large semicircular scales and densely crowded prominent triangularshaped scale setae arranged in transverse rows; gland pores not visible as preserved. Cephalon rounded, rather arcuate, rough, with same ornamentation as described for body (fig. 11B); lateral lobes not visible as preserved (probably small); antennal lobes absent. Eyes not visible as preserved. Pereon about 0.73× of body length; pereonites overlapping, lamellar; lateral parts of pereonites (i.e., epimeral plates) only moderately prominent, those of two posterior pereonites with posterior corners acutely triangular, all edged with small appressed spikes and distinctly prominent triangular-shaped scale setae (as a continuity to those of dorsal surface). Pleon about 0.24× of body length, distinctly narrower than pereon (abruptly contracted); pleotelson short, subtriangular, with rounded apex bearing two short distal setae. Mouthparts considerably well developed but without visible details. Antennule poorly visible; distal article with three short thick spines distally. Antenna (fig. 11A) very spinose in appearance, packed with small scale setae and spines, rather stout, with five peduncular articles and a multiarticulate flagellum; articles 1–3 partly obscured by head, apparently short and stout; fourth article elongate, stout, 0.18 mm long, 2.25× longer than broad; fifth article elongate and much slender than fourth article, 0.22 mm long, 1.22× length of preceding article, 5.5× longer than broad; flagellum partially preserved, with at least three articles not clearly delimited. Pereopods all alike, of moderate size; all pereopods without sexual modifications; basis and ischium mostly obscured by body; merus, carpus, and propodus with elongate, thick, nearly straight spines generally arranged in a longitudinal row on sternal margins giving them a rippled appearance; propodus and carpus length apparently subequal; dactylus with enlarged outer claw and minute inner claw; dactylar setae unbranched, glabrous, long, and slender, distinctly expanded distally; ungual setae not visible as preserved. Pleopods not visible as preserved. Uropod (fig. 11C) moderately elongate, about 0.28× of body length as preserved, freely projecting caudally; protopod produced inside pleotelson, subrectangular, long and stout, 0.15 mm long, with endopod and exopod inserted at same level; exopod styliform, long and slender, width not measurable due to poor preservation, 0.35 mm long as preserved (distal part not preserved), distinctly longer than endopod, with some very short thin setae sparsely distributed; endopod conical, maximum width at base 0.03 mm, tapering distally, 0.21 mm long (excluding terminal tuft of setae), with some very short thin setae sparsely distributed, and a tuft of long setae apically (0.10 mm long). HOLOTYPE: MCNA 12546, sex unknown, exposed dorsolaterally, ventrolaterally, and laterally. Preserved in darkened orange-colored amber, trimmed to 3.5×1.7× 1.2 mm (in an epoxy trapezoid 23.0×15.0× 1.5 mm), surrounded by numerous pseudoinclusions (Lozano et al., 2020) and particles of detritus; no other major inclusions. The specimen is nearly complete, but some areas are poorly visible owing to the position of the animal during fossilization. The distal parts of the uropodal endopod and the antennulae are not preserved. OCCURRENCE: Peñacerrada I amber site [Peñacerrada I = Moraza], eastern margin of the Basque-Cantabrian Basin, Burgos, northern Spain; Early Cretaceous (Late Albian). ETYMOLOGY: The specific epithet refers to Helen of Troy, a figure from Greek mythology known for her considerable beauty and having “launched a thousand ships” (à la Marlowe) as Menelaus and Agamemnon initiated the Trojan War to return her to Achaea. REMARKS: Heraclitus helenae may be clearly differentiated from A. resinicola by the roughened cuticle, the spinose antennulae, the dactylus with two claws, and the longer uropodal endopod. It is perhaps a representative of Crinocheta: Detonidae? owing to the antennule with a 5-jointed peduncle and a 4-jointed flagellum, the antennulae with conspicuous scaly tubercles, the inner claw small, the uropodal protopod surpassing the pleotelson, the exopodites leaf shaped, and the endopodites styliform (Schmidt, 2002). Oniscidea Indet. SPECIMEN MCNA 14907: (fig. 12A) is preserved in darkened orange-colored amber, trimmed to 10.5×6.0× 1.5 mm (in an epoxy trapezoid 21.1×13.5× 2.2 mm), and surrounded by numerous pseudoinclusions (Lozano et al., 2020), particles of detritus, fungal hyphae, and three coprolites. The specimen is exposed dorsally and ventrally, poorly preserved, with the cuticle virtually degraded except for the lateral parts of the segments, which are edged with small appressed spikes and some small triangular-shaped scale setae. Worthy of note are the oval-shaped medium-sized body (total length 4.27 mm as preserved, maximum width 2.27 mm), and the moderately large eyes composed of multiple ommatidia (only the left eye is visible). The specimen also possesses a partly preserved antenna (cut just below the articulation between the fourth and fifth articles) that is elbowed between the fourth and fifth articles. The first peduncular article is partly concealed by the head, whereas the measurements of the rest of articles are as follows: second article 0.22 mm long, about as long as broad; third article 0.25 mm long, 1.25× longer than broad, with one long, thick spine; fourth article elongate, 0.58 mm long, 4.83× longer than broad. Only some anterior pereopods are visible, but these are so thoroughly obscured owing to preservation as to provide no useful delineation of features. However, long, thick, nearly straight spines, a dactylus with two claws (the inner claw minute, distinctly smaller than in H. helenae), and an unbranched, glabrous, long, and slender dactylar seta are visible on some pereopods. The specimen is difficult to place accurately among Oniscidea, but placement within the family Ligiidae seems most appropriate owing to the presence of large eyes composed of multiple ommatidia, the comparatively elongate antennae, and a dactylus with two claws. Interestingly, the specimen corresponds with the anterior half of an exuvia. Isopods perform a biphasic molt, which consists of the shedding of the posterior and then the anterior half of the body. The boundary between the two halves is between the fourth and fifth pereonites, which agrees with that observed in the fossil. SPECIMEN MCNA 9458: (fig. 12B) is preserved in darkened yellow-colored amber, trimmed to 5.2×3.5×1.0 mm (in an epoxy trapezoid 19.1×14.1× 1.1 mm), and surrounded by numerous pseudoinclusions and particles of detritus; the amber is also darkened near the inclusion. The specimen is exposed dorsally and ventrally, and although a vague outline of the pereopods armed with long, stout, nearly straight spines can be seen, these are so thoroughly obscured owing to preservation and by the adjacent body mass as to provide no useful delineation of features. It possesses a more or less oblong, small body (total length 2.09 mm long as preserved, maximum width 0.87 mm), and some segments of the pereon with some small triangular-shaped scale setae. SPECIMEN MCNA 9924.2: (fig. 13; previously figured in Delclòs et al., 2007, Broly et al., 2013, and Sánchez-García et al., 2015) is preserved in darkened orange-colored amber, trimmed to 18.0×14.5× 3.5 mm (in an epoxy trapezoid 22.5×15.0× 4.8 mm). Syninclusions include one acariform Bdellidae, one Blattodea, six Diptera (one Ceratopogonidae, two Dolichopodidae, one Phoridae, two Psychodidae), one Tanaidacea (Alavatanais margulisae Sánchez-García, Peñalver, and Delclòs), two Archaeognatha, six coprolites (attributed to termites based on their general shape), and abundant fungal and plant remains (see details of syninclusions in Sánchez-García et al., 2015). The specimen is exposed laterally and partially complete (missing head), with the right side of the body and the right pereopods mostly polished off. The specimen possesses a medium-sized body (total length 3.78 mm as preserved), more or less oblong, only moderately convex dorsally. The pereon is about 0.73× the body length, with the lateral parts of the pereonites only moderately prominent, those of the four posterior segments with posterior corners acutely triangular, all edged with some small appressed spikes. The pleon is about 0.22× of the body length and distinctly narrower than the pereon (abruptly contracted). It also possesses spinous pereopods, a dactylus with two claws (the inner claw minute), and an unbranched, glabrous, long, and slender dactylar seta, distinctly expanded distally. Lastly, an elongate structure that could correspond to a short uropod is sticking out from under the pleon.

Heraclitus, new genus TYPE SPECIES: Heraclitus helenae, new species. DIAGNOSIS: Sex unknown. An oniscidean with the following combination of characters: Body size small ( ETYMOLOGY: The genus group name honors the pre-Socratic philosopher Heraclitus of Ephesus (ca. 535–475 B.C.E.), early rationalist, empiricist, and founder of philosophical ontology. We have used the common Latinized form of his Greek name. The gender of the name is masculine.

Eoligiiscus, new genus TYPE SPECIES: Eoligiiscus tarraconensis, new species. DIAGNOSIS: Sex unknown. An oniscidean with the following unique combination of characters: Body size small ( ETYMOLOGY: The new genus group name is a combination of Greek Ἠώς (Eos), goddess of the dawn; Ligia Fabricius, type genus of the family; and the Greek suffix - iskos, a diminutive commonly applied as a suffix to woodlice generic names (as in Oniscus: itself a combination of Greek ὄνος [onos], meaning, “woodlouse”), and - iskos). The gender of the name is masculine.

Heraclitus, new genus TYPE SPECIES: Heraclitus helenae, new species. DIAGNOSIS: Sex unknown. An oniscidean with the following combination of characters: Body size small (, Published as part of Sánchez-García, Alba, Peñalver, Enrique, Delclòs, Xavier & Engel, Michael S., 2021, Terrestrial Isopods from Spanish Amber (Crustacea: Oniscidea): Insights into the Cretaceous Soil Biota, pp. 1-32 in American Museum Novitates 2021 (3974) on page 19, DOI: 10.1206/3974.1, http://zenodo.org/record/5356138

Eoligiiscus, new genus TYPE SPECIES: Eoligiiscus tarraconensis, new species. DIAGNOSIS: Sex unknown. An oniscidean with the following unique combination of characters: Body size small (Eos), goddess of the dawn; Ligia Fabricius, type genus of the family; and the Greek suffix - iskos, a diminutive commonly applied as a suffix to woodlice generic names (as in Oniscus: itself a combination of Greek ὄνος [onos], meaning, “woodlouse”), and - iskos). The gender of the name is masculine., Published as part of Sánchez-García, Alba, Peñalver, Enrique, Delclòs, Xavier & Engel, Michael S., 2021, Terrestrial Isopods from Spanish Amber (Crustacea: Oniscidea): Insights into the Cretaceous Soil Biota, pp. 1-32 in American Museum Novitates 2021 (3974) on page 5, DOI: 10.1206/3974.1, http://zenodo.org/record/5356138