taxonID	type	description	language	source
03B7D164FFE4FFC4F540F9CAFE1333E0.taxon	description	urn: lsid: zoobank. org: act: 9270 D 5 F 9 - 8013 - 4 CA 8 - 8 A 2 A- 9118 D 5 B 597 AD	en	Sendra, Alberto, Jiménez-Valverde, Alberto, Selfa, Jesús, Cupello, Mario (2026): New cave japygids (Diplura: Japygidae) from North America. European Journal of Taxonomy 1043: 166-198, DOI: 10.5852/ejt.2026.1043.3217, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3217/14261
03B7D164FFE4FFC4F540F9CAFE1333E0.taxon	type_taxon	Type species Quapawjapyx osage Sendra gen. et sp. nov.	en	Sendra, Alberto, Jiménez-Valverde, Alberto, Selfa, Jesús, Cupello, Mario (2026): New cave japygids (Diplura: Japygidae) from North America. European Journal of Taxonomy 1043: 166-198, DOI: 10.5852/ejt.2026.1043.3217, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3217/14261
03B7D164FFE4FFC4F540F9CAFE1333E0.taxon	diagnosis	Diagnosis Slightly elongate and large body, over 20 mm in large adults. Epicuticle smooth visible under optical microscope. Set of macrosetae (M) in a resemble as the typical pattern in head, thorax and abdomen sclerites and appendages in Japygidae – dorsal head A, S, M, I, L and P macrosetae; 1 + 1 macrosetae on submentum; 5 + 5, 6 + 6 and 5 + 5 macrosetae on pronotum, mesonotum and metanotum; 7 + 7, 8 + 8, 8 + 8 - 9 + 9 macrosetae on pro. meso. and metathoracic sternum; 1 + 1 ma setae and 5 + 5 macrosetae on most urotergites; 2 + 2 macrosetae D on urite X; 13 + 13 - 15 + 14 macrosetae on first urosternites with three horizontal rows up to 16 + 16 macrosetae. Dorsal side of head and antennomeres with micropores on the surface. Lacinia falciform, well sclerotized, with the five laminae pectinate. Antennae with 26 antennomeres. Trichobothria with 2 dorsal / 1 ventral on IV and VI antennomeres, dorsal trichobothria (a) of IV on medial position. Prosternum and interstermun well defined in thorax. Pretarsus with two unequal claws and a sharp medial unguiculus. Lateral urotergites angles V – VIII angles more and more conspicuous with small point on VIII urotergites. Urite X with distinctly marked carinae with subparallel margins slightly converging towards posterior border; acropygium sub-quadrangular with borders slightly rounded. Lateral organ occupying half interstylar area; females with one row of GS, males with 2 – 3 GS rows, both setae shorter than stylus length. Median glandular organ with a few pseudospori with a microsetae. Median glandular organ with a few pseudospori (microsetae). Male and female genital papillae have typical shape as in Japygidae; male papillae show large and short lateral appendages. Cerci symmetric with two main teeth, Cerci symmetric, with one proximal tooth in cercus r (predental / postdental length) 0,5 – 0,7; right and left cercus with predental margin with two rows of denticles.	en	Sendra, Alberto, Jiménez-Valverde, Alberto, Selfa, Jesús, Cupello, Mario (2026): New cave japygids (Diplura: Japygidae) from North America. European Journal of Taxonomy 1043: 166-198, DOI: 10.5852/ejt.2026.1043.3217, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3217/14261
03B7D164FFE4FFC4F540F9CAFE1333E0.taxon	etymology	Etymology The generic name is in honor of the Quapaw Nation of Native Americans, who thrived along the lowlands of the Arkansas river five hundred years ago. Today, they live alongside much later European settlers from other nations.	en	Sendra, Alberto, Jiménez-Valverde, Alberto, Selfa, Jesús, Cupello, Mario (2026): New cave japygids (Diplura: Japygidae) from North America. European Journal of Taxonomy 1043: 166-198, DOI: 10.5852/ejt.2026.1043.3217, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3217/14261
03B7D164FFE5FFC8F561FC0AFAFA3218.taxon	description	urn: lsid: zoobank. org: act: 9 EB 82 F 6 A- 341 C- 4950 - 802 E- 2 EEF 37 CE 0 D 2 B Figs 1 – 9, 19	en	Sendra, Alberto, Jiménez-Valverde, Alberto, Selfa, Jesús, Cupello, Mario (2026): New cave japygids (Diplura: Japygidae) from North America. European Journal of Taxonomy 1043: 166-198, DOI: 10.5852/ejt.2026.1043.3217, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3217/14261
03B7D164FFE5FFC8F561FC0AFAFA3218.taxon	diagnosis	Diagnosis Slightly elongate and large body; over 20 mm in larger adults. Dorsal side of the head with abundant micropores mostly between two Y branches; a set of 14 + 14 M (A 1, 3; S 2, 4, 6; M 2, 3, 5; I 1 – 3, 5; L 3 and P 1 – 2). Labial palp 3 × as long as wide; lacinia falciform with five laminae pectinate. Antenna 0.32 – 0.35 × as long as body, with 26, 28 antennomeres; first and second antennomeres shorter than wide, third and fourth antennomeres slightly longer than wide with tiny reinforced borders; medial and distal antennomeres slight longer than wide. Proximal antennomeres with abundant micropores that disappear in medial and distal antennomeres; apical antennomere with about 8 placoid sensilla uneven distributed. Trichobothria with 2 dorsal / 1 ventral on IV and VI antennomeres, dorsal trichobothria of IV antennomere overpassing whorl of M. Thoracic segments slightly elongated, pronotum with 1 + 1 ma, 1 + 1 la, 3 + 3 lp 1 - 3; prescutum of mesonotum with 1 + 1 M; mesonotum with 1 + 1 ma, 1 + 1 la, 1 + 1 mp, 3 + 3 lp 1 - 3; prescutum of metanotum with 1 + 1 M, scutum of the metanotum with 1 + 1 ma, 1 + 1 la, 1 + 1 mp, 3 + 3 lp 1 – 3. Propresternum with 1 + 1 M; prosternum with 3 + 3 la, 2 + 2 ma and mp and 2 + 2 lp; meso-poststernum with 4 + 4 M; meso-intersternum with 2 + 2 M; mesosternum with 3 + 3 la, 3 + 3 ma, mi and mp plus one sagM intermediate and 2 + 2 lp; metapoststernum with 3 + 3 M; meta-intersternum with 2 + 2 M; metasternum with 4 + 4 – 3 + 3 la, 3 + 3 ma, mi and mp and one single sagM, and 2 + 2 lp. Legs slightly long, hind leg reaching posterior border of third abdominal segment. Pretarsus with two unequal claws (1.2 – 1.3 anterior to posterior claw) and sharp medial unguiculus. Urotergite I with 1 + 1 M, scutum with 1 + 1 M 5; urotergite II – VII with 1 + 1 ma setae, 5 + 5 M 1 – 5 and two additional M; urotergite VIII with 3 + 3 M 2,4 – 5 and one additional M; urite IX without M on dorsal position 8 + 8; urite X 1.1 – 1.3 × as long as wide, with distinctly marked carinae slightly converging towards posterior border; dorsal side with 2 + 2 D 1 and D 4 intracarinal; acropygium sub-quadrangular with borders slightly rounded; lateral side with 3 – 4 + 3 – 4 L; ventral side with 10 + 10 M setae arranged in 3 + 3 rows from right lateral side to left lateral side. Lateral urotergites I ‒ IV with blunt, slightly rounded posterolateral angles; from urotergites V – VIII angles more and more conspicuous with small point on VIII urotergites. Prescutum of urosternite I with 3 + 3 M; scutum with 13 + 13 – 14 + 15 M and posterior field of short slightly thicker setae preceding each lateral subcoxal organ. Urosternites II ‒ III with 15 – 16 + 15 – 16 M arrange in three horizontal rows A – C; urosternite VIII with about 7 + 7 M between two well-defined carinae plus 3 + 3 M on lateral side of carinae; ventral side of abdominal segment IX with 3 + 3 – 4 M. Median glandular organ with 4 – 9 pseudospori with microsetae. Lateral subcoxal organ, with one row of 52 – 54 GS and one row of 35 – 38 SS, organ occupying 0.58 × of interstylar area; GS / st 1 (stylus of first urosternite) = 0.33; SS / st 1 = 0.22, each organ with 2 – 3 rows of 124 GS and one row of 26 SS, organ occupying 0.58 × of interstylar area; GS / st 1 (stylus of first urosternite) = 0.25; SS / st 1 = 0.15. In addition to these glandular organs, only males showing impair of central glandular organ located in anterior position of third urosternite constituted by open narrow channel surrounding by long barbed setae. Male and female genital papillae with typical shape as in Japygidae. Cerci symmetric, 0.08 – 0.1 × as long as body, with one proximal tooth in cercus r (predental length / postdental length) = 0,5 – 0,7. Right cercus with proximal pointed tooth; predental margin with two rows of 6 + 5 round denticles, postdental crenelated margin by one row of 25 – 30 round denticles. Left cercus with proximal tooth; predental margin with two rows of 5 + 5 round denticles; postdental margin with two crenelated rows of tiny scale shape denticles.	en	Sendra, Alberto, Jiménez-Valverde, Alberto, Selfa, Jesús, Cupello, Mario (2026): New cave japygids (Diplura: Japygidae) from North America. European Journal of Taxonomy 1043: 166-198, DOI: 10.5852/ejt.2026.1043.3217, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3217/14261
03B7D164FFE5FFC8F561FC0AFAFA3218.taxon	etymology	Etymology The specific epithet is in honor of the Osage Nation of Native Americans, who now reside in Osage County, Oklahoma. They arrived and thrived a few hundred years ago in the northern mountains of Arkansas.	en	Sendra, Alberto, Jiménez-Valverde, Alberto, Selfa, Jesús, Cupello, Mario (2026): New cave japygids (Diplura: Japygidae) from North America. European Journal of Taxonomy 1043: 166-198, DOI: 10.5852/ejt.2026.1043.3217, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3217/14261
03B7D164FFE5FFC8F561FC0AFAFA3218.taxon	materials_examined	Type material Holotype USA – Arkansas • ♀; Marion County, Dogman Cave; 26 Jul. 2002; M. E. Slay leg.; TAMUIC, TAMU-ENTO X 1831147. Paratypes USA – Arkansas • 1 ♂; Newton County, Filton cave; 13 Dec. 2004; M. E. Slay, C. Bitting, G. O. Groening and D. Fenolio leg.; TAMUIC, TAMU-ENTO X 1832015 • 1 ♂; Newton County, Pretty Clean cave; 7 Jul. 2001; M. E. Slay and C. Bitting leg.; MUVHN, MUVHN-ZE 8000 • 1 ♂; Newton County, Toney Bend Mine # 2; 27 May 2004; M. E. Slay and C. Bitting leg.; ASPC.	en	Sendra, Alberto, Jiménez-Valverde, Alberto, Selfa, Jesús, Cupello, Mario (2026): New cave japygids (Diplura: Japygidae) from North America. European Journal of Taxonomy 1043: 166-198, DOI: 10.5852/ejt.2026.1043.3217, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3217/14261
03B7D164FFE5FFC8F561FC0AFAFA3218.taxon	description	Description BODY. Slightly elongate and large body; length 13 – 22 mm in adults; maximum width at urotergite II 2.2 mm (Fig. 1 A). Epicuticle smooth visible under optical microscope. Cuticle unpigmented, with sclerotized areas on mandible tips, claws, abdominal stylus ending, lateral apodemes of abdominal segments VII – X and cerci; and femoral, tibial and mostly cercal condyles. Body and appendages covered with normal setae (s), short setae (ss), long setae (ls) and microsetae (ms) mostly acting as proprioceptors in some body parts articulations, in addition to set of macrosetae (M) with typical pattern in head, thoracic and abdominal sclerites and appendages in Japygidae. Dorsal side of head and antennomeres with micropores on surface (Fig. 1 D). HEAD. Dorsal side with slightly sclerotized area between two branches of Y ecdysial line: few ms; with abundant micropores mostly between two Y branches; and 14 + 14 M: A 1, 3; S 2, 4, 6; M 2, 3, 5; I 1 – 3, 5; L 3 and P 1 – 2 (♂, paratype TAMU-ENTO X 1832015) (Fig. 2 A). On ventral side submentum with 1 + 1 M in posterior position; admentum with 3 + 3 M; mentum at base of labial palps with 4 + 4 M; external lobes of mentum with abundant thick setae. Labial palp 3 × as long as wide, with two apical M, one subapical M, one medial M and one proximal M and several sl. Labrum short with abundant short thick setae. Lacinia falciform, well sclerotized, with five laminae pectinate (Fig. 1 C). Mandible with four distensible teeth and one tiny tooth in interior position (Fig. 1 C). ANTENNAE. Antenna 0.32 – 0.35 × as long as body, with 26 antennomeres and 28 antennomeres of left antennae in ♀ holotype; first and second antennomeres shorter than wide, third and fourth antennomeres slightly longer than wide with tiny reinforced borders; medial and distal antennomeres slight longer than wide. Proximal antennomeres with abundant micropores mostly on dorsal side (Fig. 1 D), in addition to ms, ss, ls and one whorl of M; micropores disappear in medial and distal antennomeres; apical antennomere including additional whorl of M and about 8 placoid sensilla uneven distributed (Fig. 1 B). Trichobothria with 2 dorsal / 1 ventral on IV and VI antennomeres, dorsal trichobothria of IV antennomere overpassing whorl of M. THORAX. Thoracic segments slightly elongated, with few ls, ss and typical pattern of M: pronotum with 5 + 5 M (1 + 1 ma, 1 + 1 la, 3 + 3 lp 1 - 3) (Fig. 2 B); prescutum of mesonotum with 1 + 1 M; mesonotum with 6 + 6 M (1 + 1 ma, 1 + 1 la, 1 + 1 mp, 3 + 3 lp 1 - 3) (Fig. 2 C); prescutum of metanotum with 1 + 1 M, scutum of the metanotum with 5 + 5 M (1 + 1 ma, 1 + 1 la, 1 + 1 mp, 3 + 3 lp 1 – 3) (Fig. 3 A). Thoracic sternites, intersternites, and presternites well defined with few ss, sl, and several M; border of leg articulations with ms (Fig. 2 D). Pro-presternites and pro-, meso- and metasternites with internal Y-shaped cuticular structures (furcisternites) (Barlet & Carpentier 1962); only in propresternites prolongation of posterior branch (spine) is visible on surface (Denis 1949); propresternum with 1 + 1 M; prosternum with 3 + 3 la, 2 + 2 ma and mp and 2 + 2 lp; meso-poststernum with 4 + 4 M; meso-intersternum with 2 + 2 M; mesosternum with 3 + 3 la, 3 + 3 ma, mi and mp and one sagM intermediate and 2 + 2 la; meta-poststernum with 3 + 3 M; meta-intersternum with 2 + 2 M; metasternum with 4 + 4 – 3 + 3 la, 3 + 3 ma, mi and mi and one single sagM, and 2 + 2 lp (Fig. 2 D). Legs slightly long, hind leg reaching posterior border of third abdominal segment. Femur with 7 – 8 M; tibia with 4 – 7 M and calcars at ventral apex thicker very lightly than M; tarsus with 7 – 10 dorsal M and two ventral rows of 10 – 14 thick setae. Pretarsus with two unequal claws (1.2 – 1.3 anterior to posterior claw) and sharp medial unguiculus (Fig. 1 E). ABDOMEN. Abdominal tergites with scarce s and sM. Prescutum of urotergite I with 1 + 1 M, scutum with 1 + 1 M 5 (Fig. 3 B); urotergite II – VII with 1 + 1 ma setae, 5 + 5 M 1 – 5 and two additional M (Fig. 3 C); tergite VIII with 3 + 3 M 2,4 – 5 and one additional M (Fig. 3 D); urite IX without M on dorsal position 8 + 8. Urite X (Fig. 3 E) 1.1 – 1.3 × as long as wide, with distinctly marked carinae; carinae with subparallel margins slightly converging towards posterior border; dorsal side with 2 + 2 M (D 1 and D 4) intracarinal (Fig. 3 E), acropygium sub-quadrangular with borders slightly rounded (Fig. 3 E); lateral side with 3 – 4 + 3 – 4 M (L) (Fig. 3 E); ventral side with 10 + 10 M setae arranged in 3 + 3 rows from right lateral side to left lateral side (Fig. 5 C). Lateral urotergites I ‒ IV with blunt, slightly rounded posterolateral angles; from urotergites V – VIII angles more and more conspicuous with small point on VIII urotergites (Fig. 4 A – C). Urosternites with scarce ms and ss. Prescutum of urosternite I with 3 + 3 M; scutum with 13 + 13 – 14 + 15 M and posterior field of short slightly thicker setae preceding each lateral subcoxal organ (45 – 90 setae each). Urosternites II ‒ III with 15 – 16 + 15 – 16 M arrange in three horizontal rows A – C (Fig. 5 A); urosternite VIII with about 7 + 7 M between two well-defined carinae plus 3 + 3 M on lateral side of carinae (Fig. 5 B); ventral side of abdominal segment IX with 3 + 3 – 4 M. Median glandular organ with 4 – 9 pseudospori with microsetae, more abundant in females (Fig. 6 C) than in males (Fig. 6 F). Lateral subcoxal organ, in holotype each organ with one row of 52 – 54 GS and one row of 35 – 38 SS, organ occupying 0.58 × of interstylar area; GS / st 1 (stylus of first urosternite) = 0.33; SS / st 1 = 0.22 (Fig. 6 A – B; ♂, paratype TAMU-ENTO X 1832015), each organ occupying 0.58 of the interstylar area have 2 – 3 rows of 124 GS and one row of 26 SS; GS / st 1 (stylus of first urosternite) = 0.25; SS / st 1 = 0.15 (Fig. 6 D – E). In addition to these glandular organs, only males showing impair of central glandular organ located in anterior position of third urosternite constituted by open narrow channel surrounding by long barbed setae (Fig. 7 B). Male and female genital papillae with typical shape as in Japygidae; male papillae show large and short lateral appendages bearing ss and numerous sl (Fig. 7 A). Cerci symmetric, length 1.1 – 1.9 mm (0.08 – 0.1 × as long as body), with one proximal tooth in cercus r (predental length / postdental length) = 0,5 – 0,7. External shape of cerci straight in proximal half and curved in distal half, becoming hook towards apex; heavily sclerotized with dorsal and ventral outer carinae arising from dorsal and ventral acetabular articulations (Fig. 3 E – F). Right cercus dorsally concave and with distal ends upward, less evident in left cercus (Fig. 8 A). Teeth almost symmetrical. Right cercus with proximal pointed tooth; predental margin with two rows of 6 + 5 round denticles (Fig. 3 F), postdeltal crenelated margin by one row of 25 – 30 round denticles (Fig. 8 A, C). Left cercus with proximal tooth, less conspicuous than right one; predental margin with two rows of 5 + 5 round denticles; postdental margin with two crenelated rows of tiny scale shape denticles (Fig. 8 B, D). Right cercus with 16 dorsal M and 14 ventral M position; left cercus with 12 dorsal M and 6 ventral M position (Fig. 9 A – D). Campaniform sensilla present on hook and inner margins of cerci (Fig. 8 C – D). Taxonomic affinities of the genus and species The species of Japygidae started to be described in the second half of the 19 th century and the beginning of the 20 th (Cook 1899; Grassi 1886, 1889; Haliday 1864; Péringuey 1901; Verhoeff 1904, 1923; Westwood 1842), including for North American caves (Packard 1874). Nevertheless, roughly speaking, it is not until Filipo Silvestri’s contributions that the taxonomy of the family started to be worked out more comprehensively – see for instance Silvestri (1928, 1929 a, 1929 b, 1930 a, 1934, 1948 a, 1948 b, 1948 c, 1948 d). Yet, only a few taxonomic characters have been used for the classification of genera and species within the family. These include the cercal ornamentation, the presence of lateral and central glands on the first urosternite, and the number of pectinate laminae on the lacinia, among a few others. Nonetheless, Juraj Paclt (1957) paid special attention to the morphology of the cerci as a primary criterion of classification, which led him to reduce the number of genera – most of them originally described by Silvestri (1949) ‒ from 42 to 23. In contrast, and in later years, Jean Pagés (1995) downplayed Paclt’s criteria, which he considered overly simplistic. Instead, he emphasized chaetotaxy features such as the arrangement of macrosetae, sensorial and glandular setae, and so reinstated several of the former genera, and proposed new ones (Pagés 1953, 1955, 1980, 1981, 1993, 1994, 1995, 2000). If we followed Paclt’s (1957) criterion, the specimens studied from caves in Arkansas (North America) would have led us to propose the new species within the genus Indjapyx Silvestri, 1930, which includes 32 species widely distributed across tropical and subtropical regions from Oriental Asia to the Pacific Islands (i. e., Silvestri 1930 b, 1948 d; Pagés 1995, 1978, 2002). However, we have considered additional chaetotaxy characters, such as the glandular setae of the lateral and central organs and certain ornamentation on the cerci, to propose a new taxon at the genus level: Quapawjapyx Sendra gen. nov. Quapawjapyx Sendra gen. nov. shares a notable similarity in cercal morphology with Indjapyx: species of both genera have subsymmetrical cerci, each bearing a main tooth and two rows of predental tubercles along the inner margin (Silvestri 1930 b; Pagés 1978, 1994). This general cercal structure is also shared by eighth other genera within the Japygidae: Abjapyx Silvestri, 1949, Catajapyx Silvestri, 1933, Imazighenjapyx Sendra & Sánchez-García, 2023, Isojapyx Silvestri, 1948, Japyx Haliday, 1864, Monojapyx Paclt, 1957, Parindjapyx Silvestri, 1933, and Pauperojapyx Pagés, 1995 (Silvestri 1933 a; Silvestri 1948 d, 1948 e; Paclt 1957; Pagés 1995; Sendra et al. 2023). However, only four of these genera (Imazighenjapyx, Indjapyx, Paurojapyx, and Parindjapyx) share with Quapawjapyx two additional key taxonomic characters: a pectinate first lamina of the maxillae, and the presence of pseudosporis (glandular micro-pores, with or without microsetae) in the median glandular organ of the first urosternite. Nevertheless, each of these genera is characterized by several distinctive features that clearly differentiate them from one another. On the one hand, Imazighenjapapyx is characterized by an unusual high number of macrosetae on the thoracic sternal sclerites (e. g., prosternum with 72 M, mesosternum with 84 M and metasternum 89 M) (Sendra et al. 2023). In contrast, Quapawjapyx Sendra gen. nov. presents much lower counts: prosternum with 16 M, mesosternum with 17 M and metasternum with up to 18 M. On the other hand, Indjapyx, Paurojapyx, and Parindjapyx possess lateral organs on the first urosterntite bearing few GS setae, generally about a dozen, usually similar in size to or longer than the adjacent styli (GS / st 1 about 1) (Silvestri 1930 b, 1933 a; Pagés 1995). This condition is markedly different in Quapawjapyx, which exhibits up to 3 rows with as many as 128 GS, all distinctly shorter than the nearby styli. Additionally, the cerci of Quapawjapyx have a very proximal tooth (r = 0,5 – 0,7), clearly separated from the central tooth, unlike in Indjapyx, Paurojapyx, and Parindjapyx, where the ratio is higher (r around 0.8 – 1.3) (Silvestri 1930 b, 1933 a; Pagés 1995). All these distinguishing features support the recognition of Quapawjapyx and characterize its sole species, Q. osage Sendra gen. et spec. nov. Quapawjapyx osage Sendra gen. et spec. nov. shows moderate cave-adapted features (Pages 1964; Sendra et al. 2023), starting with its slender body, particularly in the thoracic segments, and a relatively large overall body size of 13 – 22 mm in adults. The last antennomere bears 8 placoid sensillae; the labial palp is 3 × as long as wide; and the hind leg reaches the posterior border of the third abdominal segment.	en	Sendra, Alberto, Jiménez-Valverde, Alberto, Selfa, Jesús, Cupello, Mario (2026): New cave japygids (Diplura: Japygidae) from North America. European Journal of Taxonomy 1043: 166-198, DOI: 10.5852/ejt.2026.1043.3217, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3217/14261
03B7D164FFE9FFDCF532FD28FB4531A3.taxon	description	urn: lsid: zoobank. org: act: E 4 C 20253 - 2250 - 431 A- 9 AFD- 8 DED 270 A 90 A 7 Figs 10 – 19	en	Sendra, Alberto, Jiménez-Valverde, Alberto, Selfa, Jesús, Cupello, Mario (2026): New cave japygids (Diplura: Japygidae) from North America. European Journal of Taxonomy 1043: 166-198, DOI: 10.5852/ejt.2026.1043.3217, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3217/14261
03B7D164FFE9FFDCF532FD28FB4531A3.taxon	diagnosis	Diagnosis Slightly elongate body, length 18 – 20 mm in adults. Epicuticle smooth visible under optical microscope with cuticle unpigmented, with small sclerotized areas. Dorsal head side about 9 + 9 M (A 1, 4; S 1, 4, 6; M 2, 3, 5; I 2 – 3, 5, P 2). Submentum with 1 + 1 M, admentum with 3 + 3 M; mentum with 1 + 1 M. Labial palp 3.3 × as long as wide. Lacinia falciform with the five laminae pectinate; besides third lamina pectinate insertion there are two thin spines. Antenna 0.37 × as long as body, with 30 antennomeres; last antennomere with more than 10 placoid sensilla unevenly distributed. Thoracic segments slightly elongated; scutum of pronotum, mesonotum and metanotum with 5 + 5 M; prescutum of mesonotum and metanotum with 1 + 1 M; propresternum with 1 + 1 M; prosternum with 3 + 3 la, 2 + 2 ma and mp, one short sagM intermediate and 3 + 3 lp M; meso-poststernum with 5 + 5 M; meso-intersternum with 3 + 3 M; mesosternum with 3 + 3 la, 2 + 2 ma and two sagM intermediate and 8 + 8 lp; meta-poststernum with 5 + 5 M; meta-intersternum with 3 + 3 M; metasternum with 3 + 3 la, 2 + 2 ma and mp and two sagM. Hind leg reaching the posterior border of fifth abdominal segment. Prescutum of urotergite I with 1 + 1 M, scutum without macrosetae; urotergites II – V with 1 + 1 ma setae, 5 + 5 M 1 – 5 and one additional M; urotergites VI – VII with 1 + 1 ma, 5 + 5 M 1 – 5 and two additional M; urotergite VIII with 1 + 1 ma, 4 + 4 M 1 – 2,4 – 5. Urite X 1.6 – 1.7 × as long as wide, with distinctly marked subparallel carinae converging towards posterior border; dorsal side with M (D 1 – 2, 4) intracarinal, acropygium sub-quadrangular with borders slightly rounded; lateral side with 3 + 3 M (L); ventral side with 10 + 10 M setae arranged in 3 + 3 rows from right lateral side to left lateral side each with 5 + 5 M. Lateral urotergites I ‒ VII with blunt, slightly rounded posterolateral angles; urotergites VIII with an apparent lateral angle. Prescutum of urosternite I with 4 + 4 M; scutum with 13 + 13 M arranged in three rows (A – C) and posterior field of short slightly thicker setae preceding each lateral subcoxal organ. Urosternites II ‒ III with 16 – 16 + 17 – 17 M arranged in three horizontal rows; urosternite VIII with central portion with 5 + 1 + 5 M and one sagittal M, lateral sides separated by well-defined carinae, with 4 + 4 M (A and B); ventral side of abdominal segment IX with 3 + 3 M. Median glandular organ with 4 – 6 pseudospori with microsetae. Lateral subcoxal organ in females each organ with two rows of about 110 GS and one row of about 50 SS, organ occupying 0.60 × of interstylar area, being their setae relatively short with GS / st 1; in males each organ with three rows of about 220 SG and one row of about 12 SS, organ occupying 0.64 × of interstylar area, being their setae relatively shorter than female. In males an impair of central glandular organ in the anterior position of the third and fourth urosternites. Cerci asymmetric, strong, 0.08 – 0.1 × as long as body; right cercus with medial pointed tooth; predental margin with two large pointed denticles, postdental margin crenelles by 9 – 10 tiny denticles. Left cercus with distal tooth, less conspicuous than right one; predental margin with two rows of 11 dorsal denticles and 7 ventral denticles, one of them larger than others; postdental margin smooth.	en	Sendra, Alberto, Jiménez-Valverde, Alberto, Selfa, Jesús, Cupello, Mario (2026): New cave japygids (Diplura: Japygidae) from North America. European Journal of Taxonomy 1043: 166-198, DOI: 10.5852/ejt.2026.1043.3217, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3217/14261
03B7D164FFE9FFDCF532FD28FB4531A3.taxon	etymology	Etymology ‘ Nimiipuu’ (‘ we, the people’) is the name by which the Nez Perce Native American tribe refers to themselves. They have thrived in the Pacific Northwest of North-America for more than ten thousand years and today live primarily on their tribal reservation in Idaho. We dedicate this new species to the Nimiipuu people and to their enduring struggle for survival.	en	Sendra, Alberto, Jiménez-Valverde, Alberto, Selfa, Jesús, Cupello, Mario (2026): New cave japygids (Diplura: Japygidae) from North America. European Journal of Taxonomy 1043: 166-198, DOI: 10.5852/ejt.2026.1043.3217, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3217/14261
03B7D164FFE9FFDCF532FD28FB4531A3.taxon	materials_examined	Type material Holotype USA – Idaho • ♀; Lincoln County, Little Arch Cav.; 16 Jul. 1999; D. Hubbard Jr leg.; TAMUIC, TAMU-ENTO X 1831140. Paratypes USA – Idaho • 1 ♀; Blaine County, Chalk Cave; 11 Jul. 2000; D. Hubbard Jr leg.; MUVHN, MUVHN-ZE 8001 • 1 ♂; Blaine County, Wagon Butte Cave; 13 Jul. 2000; D. Hubbard Jr leg.; TAMUIC, TAMU-ENTO X 1856622 • 2 incomplete specs; Government Cave, Blaine County; 16 Jul. 2000; D. Hubbard Jr leg.; TAMUIC, TAMU-ENTO X 1856559, TAMU-ENTO X 1835908. Other material examined USA – Idaho • 1 incomplete ♂; Lincoln County, Tee Cave; 12 July 2000; D. Hubbard Jr leg.; ASPC.	en	Sendra, Alberto, Jiménez-Valverde, Alberto, Selfa, Jesús, Cupello, Mario (2026): New cave japygids (Diplura: Japygidae) from North America. European Journal of Taxonomy 1043: 166-198, DOI: 10.5852/ejt.2026.1043.3217, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3217/14261
03B7D164FFE9FFDCF532FD28FB4531A3.taxon	description	Description BODY. Slightly elongate body, length 18 – 20 mm in adults; maximum width at urotergite IV 1.8 mm (Figs 10 A, 11 A). Epicuticle smooth visible under optical microscope. Cuticle unpigmented, with sclerotized areas on mandible tips, claws, abdominal stylus ending, lateral apodemes of abdominal segments VII – X and cerci; and femoral, tibial and mostly cercal condyles. Body and appendages covered with normal short setae (ss) and few long setae (ls) and microsetae (ms) mostly acting as proprioceptors in some body parts articulations, plus the typical pattern of macrosetae (M) in thoracic and abdominal sclerites and appendages in Japygidae; sclerites of V – X abdominal segments covered by micropores and ms, also present scares on other segments (Fig. 11 D). HEAD. Dorsal side about 9 + 9 M: A 1, 4; S 1, 4, 6; M 2, 3, 5; I 2 – 3, 5 and P 2 (on Idaho _ 1 _ holotype). On ventral side submentum with 1 + 1 M in posterior position; admentum with 3 + 3 M; mentum at base of labial palps with 1 + 1 M; external lobes of mentum with abundant thick setae. Labial palp 3.3 × as long as wide, with two apical M, one subapical M and several sl. Labrum short with abundant short thick setae. Lacinia falciform, well sclerotized, with five laminae pectinate, first one modestly pectinate with distal fringe. Besides third lamina pectinate insertion there are two thin spines (Fig. 11 B – C). Mandible with four distensible teeth and one tiny tooth in interior position. ANTENNAE. Antenna 0.37 × length of body, with 30 antennomeres; first antennomere short than wide, second to fourth antennomeres slightly longer than wide with tiny reinforced borders; medial and distal antennomeres slight longer than wide: all antennomeres covered with ss, ls and one whorl of M, but two whorls of M in penultimate and ultimate antennomere; this last antennomere also included more than 10 placoid sensilla uneven distributed (Fig. 10 B). Thirteen trichobothria present on antennomeres IV ‒ VI: 2 dorsal / 1 ventral on IV, 3 / 2 on V and VI; with dorsal trichobothria (a trichobothria of the IV antennomere overpassing the whorl of M. THORAX. Thoracic segments slightly elongated, with few ls, ss and typical pattern of M: scutum of pronotum, mesonotum and metanotum with 5 + 5 M (1 + 1 ma, 1 + 1 la, 3 + 3 lp 1 - 3); prescutum of mesonotum and metanotum with 1 + 1 M (Fig. 12 A). Thoracic sternites, intersternites, and presternites well-defined with ss, sl, and several M; border of leg articulations with fields of ms as proprioceptors (Figs 12 B, 13 A – B). Pro-presternites and pro-, meso- and metasternites with internal Y-shaped cuticular structures (furcisternites) (Barlet & Carpentier 1962); only in propresternites prolongation of the posterior branch (spine) is visible on surface (Denis 1949). Propresternum with 1 + 1 M; prosternum with 3 + 3 la, 2 + 2 medial anterior and posterior M, one short sagittal M intermediate and 3 + 3 lp; meso-poststernum with 5 + 5 M; meso-intersternum with 3 + 3 M; mesosternum with 3 + 3 la M, 2 + 2 ma and two sagittal mi and 8 + 8 lp; meta-poststernum with 5 + 5 M; meta-intersternum with 3 + 3 M; metasternum with 3 + 3 la, 2 + 2 ma, mp and two sagM (Figs 12 B, 13 B). Legs slightly long, hind leg reaching the posterior border of the fifth abdominal segment. Coxa with 3 sternal M; trochanter with 4 sternal M and one interior M; femur with 10 M mostly in ventral position and row of thick setae in apical position; tibia with 4 long M on dorsal position and 5 thick M on ventral in addition to calcars; tarsus with 5 dorsal long M and two ventral rows of 12 thick setae. Pretarsus with two unequal claws (1.4 – 1.5 anterior to posterior claw) and sharp medial small unguiculus (Fig. 11 E). ABDOMEN. Abdominal tergites with scarce ss and sl. Prescutum of urotergite I with 1 + 1 M, scutum without macrosetae (Fig. 14 A); urotergites II – V with 1 + 1 ma setae, 5 + 5 M 1 – 5 and one additional M in lateral position (Fig. 14 B – C); urotergites VI – VII with 1 + 1 ma, 5 + 5 M 1 – 5 and two additional M in lateral position (Figs 14 D, 15 A); urotergite VIII with 1 + 1 ma, 4 + 4 M 1 – 2,4 – 5 (Fig. 15 B); urite IX without M on dorsal position. Urite X (Fig. 18 A – B) 1.6 – 1.7 × as long as wide, with distinctly marked carinae; carinae with subparallel margins slightly converging towards posterior border; dorsal side with M (D 1 – 2, 4) intracarinal (Fig. 18 A), acropygium sub-quadrangular with borders slightly rounded (Fig. 18 A); lateral side with 3 + 3 M (L) (Fig. 18 A); ventral side with 10 + 10 M setae arranged in 3 + 3 rows from right lateral side to left lateral side with each with 5 + 5 M (Fig. 18 B). Lateral urotergites I ‒ VII with blunt, slightly rounded posterolateral angles; urotergite VIII with apparent lateral angle conspicuous (Fig. 15 B). UROSTERNITES. With ss and few sl. Prescutum of urosternite I with 4 + 4 M; scutum with 13 + 13 M arranged in three rows (A 1, 3 – 5, 7, B 1 – 3,5, C 1, 4, 7, 9) and posterior field of short slightly thicker setae preceding each lateral subcoxal organ (98 – 104 setae each) (Fig. 16 A). Urosternites II ‒ III with 16 – 16 + 17 – 17 M arrange in three horizontal rows (A 1 – 7) 1 – 9, B 1 – 3, 5, C 2 – 3, 6, 8 – 9 (Fig. 16 B); urosternite VIII with central portion with 5 + 1 + 5 M (A 1 – 2, B 1 – 2, C 1 – 3 and one sagittal M, lateral sides separated by well-defined carinae, with 4 + 4 M (A 3 – 4, B 3 – 4) (Fig. 16 C); ventral side of abdominal segment IX with 3 + 3 M (Fig. 16 C). MEDIAN GLANDULAR ORGAN. Of 4 – 6 pseudospori with a microsetae (Fig. 17 B). Lateral subcoxal organ: in female (paratype MUVHN-ZE 8001), each organ with two row of about 110 GS and one row of about 50 SS, organ occupying 0.60 × of interstylar area, being their setae relatively short with GS / st 1 (stylus of first urosternite) = 0.19; SS / st 1 = 0.16 (Fig. 17 A); in male (paratype TAMU-ENTO X 1856622), each organ with three rows of about 220 SG and one row of about 12 SS, organ occupying 0.64 × of interstylar area, being their setae relatively shorter than female: GS / st 1 (stylus of first urosternite) = 0.16; SS / st 1 = 0.11 (Fig. 17 C). In addition to these glandular organs, in males impair of central glandular organ located in anterior position of third and fourth urosternites constituted by open narrow channel surrounding by few long setae (Fig. 17 D). GENITAL PAPILLAE (male and female). With typical Japygidae shape; male papilla with conical appendages, 1.3 × as long as wide, bearing ss and numerous sl and long and large row of posterior setae on posterior border of genital opening (Fig. 11 F – G). CERCI. Asymmetric, strong, length 1.4 – 1.8 mm (0.08 – 0.1 × as long as body), slightly curved in proximal and medial exterior shape becoming hook towards apex; borders sclerotized with dorsal and ventral outer carinae arising from dorsal and ventral acetabular articulations (Fig. 10 C). Right cercus with medial pointed tooth; predental margin with two large pointed denticles, postdental margin crenelles by 9 – 10 tiny denticles (Fig. 10 C, E). Left cercus with distal tooth, less conspicuous than right one; predental margin with two rows of 11 dorsal denticles and 7 ventral denticles, one of them larger than others; postdental margin smooth (Fig. 10 C – D). Right cercus with 8 D dorsal, 10 L and 12 V; left cercus with 6 D M, 16 L and 15 V M. Taxonomic affinities Taking into account the simplistic taxonomic perspective adopted by Paclt (1957), the specimens studied from caves in Idaho would be assigned to Burmjapyx Silvestri, 1930. Paclt (1957) grouped under Burmjapyx eighth previously described genera: Holjapyx Silvestri, 1948, Protjapyx Silvestri, 1948, Austrjapyx Silvestri, 1949, Hapljapyx Silvestri, 1949, Merojapyx 1949, Henicjapyx, 1949, Xenjapyx Silvestri; 1949, and Homojapyx Pagés, 1953. In Paclt’s redefinition, Burmjapyx is characterized by a medial organ lacking disculis, five pectinate laminae on the maxillae, and asymmetric cerci bearing two predental denticles on the left cercus and only one on the right cercus. However, following the approach of Pagés (1995), we have adopted a more detailed chaetotaxy criterion for distinguishing genera within Japygidae. Accordingly, we retain the original genera described prior to Paclt’s revision and, based on these criteria, we assign the Idaho cave japygids to the genus Holjapyx. Silvestri (1948 a) established the genus Holjapyx to accommodate a previously described species, Holjapyx diversiunguis (Silvestri, 1911) [= Japyx diversiunguis Silvestri, 1911], originally found in Yosemite Park (California). In his initial description, Silvestri (1911) highlighted the unequal proportions between the two main claws and the presence of a small unguiculus in between. Later, in his diagnosis of the genus, Silvestri (1948 a) emphasized several key characters, in addition to those adopted for Burmjapyx by Paclt (1957), such as the presence of 26 antennomers and more notable, a specialized glandular organ or sac in males, located in the anterior medial region of the third and fourth urosternites. Four years later, Pagés (1952) described three species from New Zealand, which he tentatively assigned to Holdajpyx, using the provisional designation “ Holjapyx? ”. He noted that males of these species lack the medial sac on any urosternite and possess a greater number of antennomeres. However, he also highlighted their similarity to H. diversiunguis in two features not mentioned by Silvestri (1948 a): (1) the presence of fine spines in the middle of the pectinated laminae on the internal lobe of the maxilla, and; (2) the number of additional macrosetae on urosternites III – VII, which display the typical 5 + 5 macrosetae (M 1 – 5) plus two extra macrosetae (Pagés 1952). In the same decade, Leslie M. Smith (1959) made the last major contribution to the taxonomy of Holjapyx with the description of nine new species, all from California State. These species were distinguished based on minor morphological differences, and Smith (1959) also published a taxonomic key for the species of Holjapyx, excluding the species of “ Holjapyx? ” described by Pagés (1952). In the same work, Smith (1959 b) redefined the genus Holjapyx, highlighting the diagnostic value of the position of the main tooth on the left cercus. He introduced the ratio R, defined as the distance from the base of the left cercus to the main tooth divided by the total length of the cercus (see Fig. 10 D – F), and reported R values ranging from 0.371 to 0.541 for the Californian species of Holjapyx. We proposed Holjapyx nimiipuu Sendra sp. nov., which exhibits all generic features previously described along with several exclusive characters that justify its recognition as a new species. Three of these distinguishing characters are related to cave adaptations: (1) the larger body size, reaching up to 20 mm in H. nimiipuu, which is more than twice the size of soil-dwelling species of Holjapyx; (2) the longer antennae, with 30 antennomeres instead of the usual 26; and (3) the significantly higher number of placoid sensilla, with at least ten (Fig. 10 B), compared to the only three in other species of Holjapyx. Additionally, H nimiipuu is characterized by the presence of two conspicuously large predental denticles on the right cercus (Fig. 10 D – F). As previously noted regarding the antennae, H. nimiipuu Sendra sp. nov. shows clear cave-adapted features (Pagés 1964; Sendra et al. 2023), which are also evident in its slightly elongated body – particularly the thoracic segments –, its labial palp, which is 3.3 × as long as wide, the hind leg, which reaches the posterior margin of the fifth abdominal segment, and the urite X, which is 1.6 – 1.7 × as long as wide. Other studied material from the Texas University Collection Alongside these two new cave-adapted japygid species, the Texas University collection includes several non-cave-adapted japygids listed below and included in the distributional map (Fig. 19).	en	Sendra, Alberto, Jiménez-Valverde, Alberto, Selfa, Jesús, Cupello, Mario (2026): New cave japygids (Diplura: Japygidae) from North America. European Journal of Taxonomy 1043: 166-198, DOI: 10.5852/ejt.2026.1043.3217, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3217/14261
03B7D164FFFDFFDCF526FDA2FC8E3393.taxon	materials_examined	Material examined USA – California • 1 ♂; Tulare County, Lost Soldier’s Cave; 21 Jul. 2003; TAMUIC, TAMUIC _ JKK 54 - A- 5 • 1 ♀; Tulare County, Lost Soldier’s Cave; 23 Jul. 2003; TAMUIC, TAMUIC _ JKK 79 - A- 2 • 1 ♀; Tulare County, Lost Soldier’s Cave; 6 Jul. 2004; TAMUIC, TAMUIC _ JKK 12 - D- 3 • 2 ♂♂; Tulare County, Lost Soldier’s Cave; 7 May 2004; TAMUIC, AMUIC _ JKK 102 - C- 2, TAMUIC _ JKK 101 - C- 9 • 1 ♂; Tulare County, Lost Soldier’s Cave; 9 Jun. 2003; TAMUIC, TAMUIC _ JKK 103 - C- 7.	en	Sendra, Alberto, Jiménez-Valverde, Alberto, Selfa, Jesús, Cupello, Mario (2026): New cave japygids (Diplura: Japygidae) from North America. European Journal of Taxonomy 1043: 166-198, DOI: 10.5852/ejt.2026.1043.3217, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3217/14261
03B7D164FFFDFFDCF526FDA2FC8E3393.taxon	description	Description Body length: 2 ♀♀ 14 – 22 mm; 4 ♂♂ 15 – 19 mm.	en	Sendra, Alberto, Jiménez-Valverde, Alberto, Selfa, Jesús, Cupello, Mario (2026): New cave japygids (Diplura: Japygidae) from North America. European Journal of Taxonomy 1043: 166-198, DOI: 10.5852/ejt.2026.1043.3217, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3217/14261
03B7D164FFFDFFDCF557FBD2FE7D348B.taxon	materials_examined	Material examined USA – Oklahoma • 1 ♀; Delaware County, McGee Cave; 31 Sep. 2001; S. Mc Ginnis and D. Fenolio leg.; TAMUIC, TAMUIC.	en	Sendra, Alberto, Jiménez-Valverde, Alberto, Selfa, Jesús, Cupello, Mario (2026): New cave japygids (Diplura: Japygidae) from North America. European Journal of Taxonomy 1043: 166-198, DOI: 10.5852/ejt.2026.1043.3217, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3217/14261
