Quapawjapyx osage Sendra gen. et sp. nov.

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Figs 1–9, 19

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 (A1, 3; S2, 4, 6; M2, 3, 5; I1–3, 5; L3 and P1–2). 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 M5; urotergite II–VII with 1+1 ma setae, 5+5 M1–5 and two additional M; urotergite VIII with 3+3 M2,4–5 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 D1 and D4 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/st1 (stylus of first urosternite) = 0.33; SS/st1 = 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/st1 (stylus of first urosternite) = 0.25; SS/st1= 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.

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.

Type material

Holotype USA – Arkansas • ♀; Marion County, Dogman Cave; 26 Jul. 2002; M.E. Slay leg.; TAMUIC, TAMU-ENTO X1831147.

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 X1832015 • 1 ♂; Newton County, Pretty Clean cave; 7 Jul. 2001; M.E. Slay and C. Bitting leg.; MUVHN, MUVHN-ZE8000 • 1 ♂; Newton County, Toney Bend Mine #2; 27 May 2004; M.E. Slay and C. Bitting leg.; ASPC.

Description

BODY. Slightly elongate and large body; length 13–22 mm in adults; maximum width at urotergite II 2.2 mm (Fig. 1A). 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. 1D).

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: A1, 3; S2, 4, 6; M2, 3, 5; I1–3, 5; L3 and P1–2 (♂, paratype TAMU-ENTO X1832015) (Fig. 2A). 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. 1C). Mandible with four distensible teeth and one tiny tooth in interior position (Fig. 1C).

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. 1D), 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. 1B). 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. 2B); 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. 2C); 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. 3A). Thoracic sternites, intersternites, and presternites well defined with few ss, sl, and several M; border of leg articulations with ms (Fig. 2D). 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. 2D). 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. 1E).

ABDOMEN. Abdominal tergites with scarce s and sM. Prescutum of urotergite I with 1+1 M, scutum with 1+1 M5 (Fig. 3B); urotergite II–VII with 1+1 ma setae, 5+5 M1–5 and two additional M (Fig. 3C); tergite VIII with 3+3 M2,4–5 and one additional M (Fig. 3D); urite IX without M on dorsal position 8+8. Urite X (Fig. 3E) 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 (D1 and D4) intracarinal (Fig. 3E), acropygium sub-quadrangular with borders slightly rounded (Fig. 3E); lateral side with 3–4+3–4 M (L) (Fig. 3E); ventral side with 10+10 M setae arranged in 3+3 rows from right lateral side to left lateral side (Fig. 5C). 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. 4A–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. 5A); urosternite VIII with about 7+7 M between two well-defined carinae plus 3+3 M on lateral side of carinae (Fig. 5B); 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. 6C) than in males (Fig. 6F). 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/st1 (stylus of first urosternite) = 0.33; SS/st1 = 0.22 (Fig. 6A–B; ♂, paratype TAMU-ENTO X1832015), each organ occupying 0.58 of the interstylar area have 2–3 rows of 124 GS and one row of 26 SS; GS/st1 (stylus of first urosternite) = 0.25; SS/st1= 0.15 (Fig. 6D–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. 7B). 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. 7A). 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. 3E–F). Right cercus dorsally concave and with distal ends upward, less evident in left cercus (Fig. 8A). Teeth almost symmetrical. Right cercus with proximal pointed tooth; predental margin with two rows of 6+5 round denticles (Fig. 3F), postdeltal crenelated margin by one row of 25–30 round denticles (Fig. 8A, 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. 8B, D). Right cercus with 16 dorsal M and 14 ventral M position; left cercus with 12 dorsal M and 6 ventral M position (Fig. 9A–D). Campaniform sensilla present on hook and inner margins of cerci (Fig. 8C–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, 1929a, 1929b, 1930a, 1934, 1948a, 1948b, 1948c, 1948d). 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 1930b, 1948d; 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 1930b; 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 1933a; Silvestri 1948d, 1948e; 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/st1 about 1) (Silvestri 1930b, 1933a; 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 1930b, 1933a; 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.