Life in darkness: an overview of cave-adapted japygids (Hexapoda, Diplura) Author Sendra, Alberto 11636BAE-AE66-4898-A7C8-35B329E7E3A8 Colecciones Entomológicas Torres-Sala, Servei de Patrimoni Històric, Ajuntament de València, Passeig de la Petxina 15, 46008 València, Spain & Departament de Didàctica de les Ciències Experimentals i Socials, Facultat de Magisteri, Universitat de València, Avda. Tarongers 4, 46022 València, Spain & Universidad de Alcalá, Research Team on Soil Biology and Subterranean Ecosystems, Department of Life Sciences, Faculty of Science, Campus Universitario Crta. A- 2 Km 33.6, 28805 Alcalá de Henares, Madrid, Spain Author Sánchez-García, Alba 86DFDA66-BEC1-428A-A7B0-E90FCFFABCE3 Instituto Geológico y Minero de España-CSIC, València, Spain alba.sanchez@igme.es Author Hoch, Hannelore 38BAFA8B-2C1B-4C84-9107-27D257570B5B Leibniz Institute for Evolution and Biodiversity Science, Museum f ̧ r Naturkunde, Humboldt-University, Invalidenstr. 43, D- 10115 Berlin, Germany Hannelore.Hoch@mfn.berlin Author Jiménez-Valverde, Alberto E31ADC69-98EE-46CB-87E2-5B27B53FF107 Universidad de Alcalá, Research Team on Soil Biology and Subterranean Ecosystems, Department of Life Sciences, Faculty of Science, Campus Universitario Crta. A- 2 Km 33.6, 28805 Alcalá de Henares, Madrid, Spain alberto.jimenezv@uah.es Author Selfa, Jesús C01B4FA6-6C5C-4DDF-A114-2B06D8FE4D20 Laboratori d’Investigació d’Entomologia, Departament de Zoologia, Universitat de València, C / Dr Moliner 50, 46100 Burjassot, València, Spain jesus.selfa@uv.es Author Moutaouakil, Soumia 3E9AF62C-F23B-4B76-B734-05FDBA87C84D Museum of Natural History of Marrakech, Cadi Ayyad University, Morocco moutaouakil.soumia@gmail.com Author Preez, Gerhard Du Unit for Environmental Sciences and Management, North-West University; Private Bag X 6001, Potchefstroom 2520, South Africa, North-West University, Potchefstroom, South Africa Gerhard.DuPreez@nwu.ac.za Author Millar, Ian Department of Conservation, Nelson, New Zealand two-millars@slingshot.co.nz Author Ferreira, Rodrigo Lopes Centro de Estudos em Biologia Subterrânea, Departamento de Ecologia e Conservação, Universidade Federal de Lavras, Lavras, MG. CEP 37200 - 900, Brazil. drops@ufla.br text European Journal of Taxonomy 2023 2023-09-29 894 1 1 54 https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2287/9877 journal article 272973 10.5852/ejt.2023.894.2287 d3af5432-00a0-4ba8-aea4-a65adb38d00b 2118-9773 8388995 11C1DFE4-02F2-4FEA-BAD1-ACCAEA3590DB Imazighenjapyx marocanus Sendra & Sánchez-García gen. et sp. nov. urn:lsid:zoobank.org:act: 59A02F26-DCDF-49DC-AA64-B0E88526A63C Figs 8–15 Etymology The specific name refers to the country of origin, Morocco . Type material Holotype MAROC • ♀ ; Agadir-Ida Outanane region, Imi Ougoug Cave ; 30°36′44.83″ N , 9°28′1.64″ W ; 26 Feb. 2020 ; Rodrigo Lopes Fereira leg.; labelled “♀-holotype - MHNM -ZAD01 ”; MHNM . Fig. 8. Imazighenjapyx marocanus Sendra & Sánchez-García gen. et sp. nov. , holotype, ♀ (MHNM). A . Antennae. B . Central antennomeres. C . Apical antennomere. D . Lateral side of the apical antennomere. E . Detail of apical antennomere. F . Detail of antennal micro-barbed sM setae. Description BODY . Elongate ( Fig. 15C‒D ), length 26 mm ; maximum width at urotergite VII 2.6 mm . Epicuticle reticulate under optical microscope; with micropores at higher magnifications (urite X with 4 micropores/μm 2 , diameter 0.10‒0.14 μm ( Fig. 11B ). Cuticle unpigmented, with sclerotized areas on mandible tips, femoral and tibial condyles, ventral apodemes of abdominal segment VIII, distal part of styles, abdominal segment X, and cerci. Body and appendages covered with very abundant ms setae as well as s, sM, and M, in addition to abundant mbsM on antennae. HEAD . Antenna length 8.6 mm , 0.33× length of body, with 41 antennomeres; basal antennomere short, followed by three slightly longer antennomeres with reinforced borders visible on ventral side; medial antennomeres as long as wide ( Fig. 8A–B ). Proximal antennomeres with ms, s, and a few long M; medial and distal antennomeres with a few ms and abundant s setae plus to two whorls of mbsM ( Fig. 8B ); apical antennomere with s and mbsM and about 24 placoid sensilla distributed in 4‒5 groups ( Fig. 8C‒F ). Trichobothria present on antennomeres IV‒VI in a 2/3/3 pattern, with a trichobothria (dorsal one) in proximal position. Head with a few s setae and M and abundant ms setae; on dorsal side 14+14 M: A1, 4, S4, 6, V1−2, V4, M3, M5, I2, L4−5 and P1−2 ( Fig. 9A ); on ventral side: submentum with large 2+2 M in anterior and posterior position plus 5+5 sM, admentum with 3+3 M, mentum at base of labial palps with 2+2 M; external lobes of mentum with abundant sM; the pair of the exertil vesicles of the external lobes visible in the holotype ( Fig. 10A ). Labial palp 4× as long as wide, with one proximal sM and seven medial and distal sM. Lacinia falciform, well sclerotized, with the five laminae pectinate. Fig. 9. Imazighenjapyx marocanus Sendra & Sánchez-García gen. et sp. nov. , holotype, ♀ (MHNM). A . Dorsal side of head and anterior part of pronotum. B . Pronotum and anterior part of mesonotum. C . Mesonotum. D . Metanotum. E . Urotergites I and II. F . Urotergites VII and VIII. Fig. 10. Imazighenjapyx marocanus Sendra & Sánchez-García gen. et sp. nov. , holotype, ♀ (MHNM). A . Ventral side of head. B . Pro-prosternum (pr-prt) with spine (sp), prosternum (pr-st), mesopoststernum (ms-pst), and meso-intersternum (ms-ist). C . Mesosternum (ms-st). D . Meta-poststernum (mt-pst), meta-intersternum (mt-ist), and metasternum (mt-st). E . Urosternite I. F . Urosternites II and III. G . Urosternites IV and V. H . Urosternites VI and VII. I . Urosternites VII and VIII. THORAX . Thoracic segments elongate, with a few s and abundant ms setae. Pronotum with 5+5 M1−5 and 6+6 sM; prescutum of mesonotum with 1+1 M; mesonotum with 5+5 M1−5 and 6+6 sM; prescutum of metanotum with 1+1 M and 3+3 sM; metanotum with 5+5 M1−5 and 11+11 sM ( Fig. 9B‒D ). Thoracic sternites, intersternites, and presternites well-defined with ms, s, and M; sM difficult to distinguish from M so they have all been counted as M ( Fig. 10B‒D ). Pro-presternites and pro-, meso- and metasternites with internal Y-shaped cuticular structures (furcisternites) ( Barlet & Carpentier 1962 ); only in pro-presternites the prolongation of the posterior branch (spine) is visible on the surface ( Denis 1949 ). Pro-presternum with 4+5 M; prosternum with 6 medial anterior M, 7+7 lateral anterior M, and 26+26 lateral posterior M; meso-poststernum with 10+9 M; meso-intersternum with 11+11 M; mesosternum with 12 medial anterior M, 11+11 lateral anterior M, and 25+25 posterior M; meta-poststernum with 14+12 M; meta-intersternum with 11+10 M; metasternum with 19 medial anterior M, 12+14 lateral anterior M, and 20+24 posterior M ( Fig. 10B‒D ). Legs slightly short, hind leg 4.9 mm long, reaching third abdominal segment. Femur-tibia-tarsus articulations with a row of sM; coxa and trochanter with 9 ventral M; femur with 6 ventral and 5 dorsal M; tibia with 6 ventral and 3 dorsal M; tarsus with 3 dorsal M and abundant sM plus two ventral rows of seven and six thick setae and a calcar at ventral apex thicker than other M. Pretarsus with two short, thick, unequal claws and a sharp medial unguiculus. Fig. 11. Imazighenjapyx marocanus Sendra & Sánchez-García gen. et sp. nov. , holotype, ♀ (MHNM). A . Lateral posterior part of first urosternite. B . Median glandular organ. Abbreviations: GS = glandular setae; Ps = pseudospores; St = stylus. ABDOMEN . Abdominal tergites with scarce s and sM. Prescutum of urotergite I with 1+1, scutum with 1+1 sM (ma), 1+1 M5 and 2+1 sM; urotergite II with 1+1 sM (ma), 2+2 M1−2, and 2+2 M4−5; urotergites III‒VII with 1+1 ma and 5+5 M1‒5; tergite VIII with 5+5 M1−5; urite IX with 8+8 ventral M ( Fig. 9E‒F ). Urite X ( Fig. 12A‒D ) 1.8× as long as wide, with distinctly marked carinae; carinae with subparallel margins slightly converging towards posterior border; dorsal side with 6+6 M intracarinal (D1−6 M), 1+1 M between D2, one sagittal M between D4, 1+1 M between D6 ( Fig. 12A ), acropygium rounded ( Fig. 12A ); lateral side with 7+7−8+8 M (L), and several sM ( Fig. 12C ); ventral side with 52 M setae apparently arranged in 6 rows from right lateral side to left lateral side ( Fig. 12D ). Urosternites with scarce ms and s setae and abundant sM and short M. Prescutum of urosternite I with 12+13 M; scutum with up to 120 M plus 200 sM on posterior position ( Fig. 10E ). Median glandular organ with abundant pseudospores (Ps), more than 30 ( Fig. 11B ). Lateral subcoxal organ with one row of about 120 glandular setae (GS) and one row of 140 sensory setae (SS); lateral subcoxal organ occupying 0.38× of interstylar area; GS/st1 (stylus of first sternite) = 0.3; SS/st1= 0.08 ( Fig. 11A ). Urosternites II‒III with about 140 M; urosternites IV‒VII with about 160 M; urosternite VIII with about 50 M between two well-defined carinae plus 5+5 M on lateral side of carinae ( Fig. 10F‒I ). Cerci asymmetric, strong, length 2.3 mm , straight in the proximal half and curved in the distal half, becoming a large hook towards apex; heavily sclerotized with dorsal and ventral outer carinae arising from dorsal and ventral acetabular articulations; carinae extending to apex ventrally and almost reaching the apex dorsally ( Figs 13A , 14A‒B ). Cerci dorsally concave and with the distal ends upward ( Fig. 12C ). Teeth subsymmetrical. Right cercus with medial tooth pointed; predental margin with two rows of 3+3 small round denticles; postdental margin scraper-like, very protruding, with a row of 15 denticles terminating near the hook. Predental margin of left cercus with three rows of 8+5+7 denticles (superior and intermediate rows with round denticles; inferior row with pointed denticles) terminating at medial large tooth; postdental margin with 11 small round denticles terminating before the hook. Right cercus with 18, 32, 30 M (dorsal, lateral, and ventral); left cercus with 19, 30, 30 M (dorsal, lateral, and ventral). Campaniform sensilla present on hook and inner margins of cerci ( Figs 13‒14 ). Fig. 12. Imazighenjapyx marocanus Sendra & Sánchez-García gen. et sp. nov. , holotype, ♀ (MHNM). A . Abdominal segment X, dorsal view. B . Detail of abdominal segment X, dorsal view. C . Lateral view of abdominal segment X and cerci. D . Abdominal segment X, ventral view. Taxonomic affinities Following Paclt (1957a) Imazighenjapyx marocanus Sendra & Sánchez-García gen. et sp. nov. resembles Indjapyx in the morphology of the cerci with two predental rows on the right and left margins, and in simple lateral subcoxal organ with pseudospores on the medial organ. The genus Indjapyx has 27 described species, all of which are from southern mainland Asia and islands ( Pagés 1984 , 1994, 2002). As pointed out by Silvestri (1930a) and Pagés (1984) , a distinctive feature of Indjapyx is the proximal position of the a trichobothria (dorsal one), which is also present in Imazighenjapyx marocanus gen. et sp. nov. However, I. marocanus lacks the typical abundant ms setae on dorsal head and shows three characters unknown in other japygid taxa: the right cercus has a highly protruding, scraper-like postdental margin; the numerous thoracic macrosetae (prosternum with 72 M, mesosternum with 84 M and metasternum 89 M); and the singular micro-barbed sM on antennomeres. Fig. 13. Imazighenjapyx marocanus Sendra & Sánchez-García gen. et sp. nov. , holotype, ♀ (MHNM). A . Terminal part of abdominal segment X and cerci in dorsal view. B . Tip of right cercus. C . Medial part of inner side of right cercus. D . Proximal to medial part of inner side of right cercus. Habitat Imi Ougoug is a limestone cave located 43 kilometres northeast of Agadir City, at the bottom of a cliff overlooking the Talmat River. The cave is 1097 meters long. It has a low entrance leading to a narrow zig-zag passage that bifurcates after 10 meters; on the left, the largest upstream segment leads to a large chamber ( 32 m long and 9 m high in the middle). This part is separated into two sections that end with siphons. On the right, a sinuous path with casings and basins continues alternately and terminates 435 m from the entrance, at - 65 m , on a little siphoning lake ( Fig. 15 ). Forty meters from the entrance, a single specimen of Imazighenjapx marocanus gen. et sp. nov. was discovered in the right branch. The specimen was found on the cave wall in a lateral overflow of the meander between a temporary waterfall and the first lake. The entrance’s morphology, which is completely rounded, and all other morphological elements clearly indicate a losing stream function for the downstream part (right): pebbles and a giant’s kettle indicate a free flow. Many ancient stalagmitic masses that eroded and polished with a marble appearance stand noticeably. However, it appears that in the event of a severe flood, the cavity will almost completely fill up. Fig. 14. Imazighenjapyx marocanus Sendra & Sánchez-García gen. et sp. nov. , holotype, ♀ (MHNM). A . Cerci, ventral view. B . Detail of right cercus in ventro-lateral view. C . Right cercus, ventral view. D . Leaft cercus, ventral view. Fig. 15. Location and habitat of Imazighenjapyx marocanus Sendra & Sánchez-García gen. et sp. nov. A . External area in the surroundings of the Imi Ougoug cave the arrow indicates the location of the cave entrance). B . Imi Ougoug cave interior general aspect. C–D . Living holotype specimen. After “Win-Timdouine” Cave, this is the region’s second most touristic cave. It has several names: “Ifri Ouado”, a Berber term that means “the blowing cave” due to the air current that comes out of it. The other name for the cave is “grotte des araignées,” which means “cave of spiders” referring to the large quantity of opilions that can be found there behind the entrance.