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 Teljapyx aotearoa Sendra & Sánchez-García sp. nov. urn:lsid:zoobank.org:act: 0A203723-553E-402D-9E6C-B8B803157256 Figs 24‒31 Etymology The specific name aotearoa refers to the Māori name for New Zealand , often translated as “land of the long white cloud”. Type material Holotype NEW ZEALAND • Ô; Takaka , Council Cave ; 40°52′35.95″ S , 172°50′46.55″ E , 4 Dec. 2020 ; Rodrigo Lopes Ferreira leg.; labelled “Ô1- holotype-AI.052510 ”; AI . Paratypes NEW ZEALAND • 1 ♀ ; same collection data as for holotype; labelled “ ♀ 1-paratype-MZB ( MCNB ) 2023-0619”; MZB • 1 Ô; same collection data as for holotype; labelled “Ô2-paratype -AI.052511”; MZB . Other studied material NEW ZEALAND • 1 ♀ ; same collection data as for holotype ; AS . Description BODY . Elongate ( Figs 24D , 25B ), length 28 mm in male holotype ( 29 mm in Ô2- paratype and 27.5 mm in ♀ 1- paratype ). Maximum width at urotergite VII of 3 mm in Ô2- paratype , and 2.9 mm in holotype and ♀ 1- paratype . Epicuticle smooth, with numerous micropores in tergites and urosternites, with about 12 micropores/μm 2 , diameter 0.18‒0.26 μm ( Fig. 28B ). Cuticle unpigmented, with slightly sclerotized areas on dorsal frontal head, mandible tips, femoral and tibial condyles, abdominal segments VIII–X, and cerci. Body and appendages covered with ms, s, sM, and M. HEAD . Antenna with 32 antennomeres, 0.50‒0.55× the length of body ( 14 mm long in holotype and ♀ 1- paratype ; 16 mm in Ô2- paratype ); first antennomere short, followed by three longer and wider antennomeres, the third and fourth being the largest and most elongated, twice as long as wide ( Fig. 26C ); medial antennomeres elongated, 1.3× as long as wide, with a large intersegmental ring between them ( Fig. 26B ).All antennomeres with sparse ms setae, plus three apparently whorls of long sM. Trichobothria present on antennomeres IV‒VI in a 2/3/3 pattern, with a trichobothria in distal position ( Fig. 26C ). Apical antennomere with 16‒18 placoid sensilla distributed in three irregular groups ( Fig. 26A ). Head with a few sM and ms; on dorsal side 18+18 M setae: A2−4, S2, 4, 6, M2−4, V2, 4, I1−3, I5, L5 and P1−2 ( Fig. 24A ); on ventral side: submentum with 1+1 large M in posterior position, admentum with 3+3 M, mentum with 1+1 M; external lobes of mentum with abundant sM. Pair of exertil vesicles of the external lobes visible in the holotype . Labial palp elongate, 10× as long as wide, with one proximal sM and four medial and distal sM, plus several ms. Lacinia falciform, well sclerotized, with the five laminae pectinate ( Fig. 25A ). Fig. 24. Teljapyx aotearoa Sendra & Sánchez-García sp. nov. , holotype, Ô (AI). A . Dorsal side of head and pronotum. B . Mesonotum. C . Metanotum. D . Habitus, dorsal side. E . Urotergites I‒II. F . Urotergites III‒V. THORAX . Thoracic segments elongate ( Figs 24A‒C , 27A‒C ). Pronotum with 5+5 M1–5, 4+4 sM, and numerous ms uniformly distributed; prescutum of mesonotum with 1+1 M; mesonotum with 6+6 M1–6 and abundant ms or sM; prescutum of metanotum with 1+1 M, metanotum with 5+5 M1–5 and a few ms; both prescuta with 1+1 M and scattered sM and ms. Thoracic sternites, intersternites, and presternites well-defined, with ms, sM, and M setae ( Fig. 27A‒C ). Pro-presternites and pro-, meso- and metasternites with strong 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 1+1 lateral anterior M and 1+1 nearby sM; prosternum with 1+1 medial anterior M; 3+3 lateral anterior M, 1+1 medial intermediate M and 2+2 lateral posterior M; meso-poststernum with 4+4 M; meso-intersternum with 2+2 M; mesosternum with 1+1 medial anterior M, 3+3 lateral anterior M, 1+1 medial intermediate M, 1 sagittal M, 9+10 medial posterior M and 2+2 lateral posterior M; meta-poststernum with 4+4 M; meta-intersternum with 2+2 M; metasternum with 1+1 medial anterior M, 2+2 lateral anterior M, 1+1 sagittal M, 1+1 medial intermediate M, 10+13 medial posterior M and 2+2 lateral posterior M ( Fig. 27 A−C). Legs elongate, hind leg reaching sixth abdominal segment, 6.8 mm long in holotype , 7.6 mm in ♀ 1- paratype and 8 mm in Ô2- paratype . No frictional setae between trochanter-coxa-femur articulations; femur-tibia-tarsus articulations with a row of long M or sM setae set on large sockets. Coxa with 4 distinct M; femur with 9‒10 M; tibia with more than 30 M, and tarsus with almost 30 M. Calcars not distinguishable from M in shape. Pretarsus with two simple and unequal claws (the shortest ⅔ the longest), and a pointed and short medial unguiculus. Fig. 25. Teljapyx aotearoa Sendra & Sánchez-García sp. nov. , holotype, Ô (AI). A . Anterior part of head, ventral side. B . Habitus, ventral side. C . First urosternite. D . Urosternites VII‒IX. E . Urosternite II. F . Genital papilla. Abbreviations: ap = appendages; o = opening. Fig. 26. Teljapyx aotearoa Sendra & Sánchez-García sp. nov. , ♀1-paratype, ♀ (MZB). A . Distal antennomere. B . Central antennomeres. C . Proximal antennomeres II‒VI.Abbreviation:T = trichobothria. ABDOMEN . Abdominal tergites with a few ms, s, sM and M. Prescutum of urotergite I with 1+1 M, scutum with 1+1 M or sM (ma) and 1+1 M5 ( Fig. 24E ); urotergite II with 1+1 M (ma), 1+1 M1, and 2+2 M4−5; urotergites III‒VII with 1+1 M (ma) and 5+5 M1‒5 ( Fig. 24F ); urotergite VIII with 7+7 M; urite IX with 3+3 ventral M. Urite X 1.5× as long as wide, with marked carinae with subparallel margins slightly converging towards the posterior border; on dorsal side with 2+2 M intracarinal D1, 3, plus 1+1 M or 1 M between D1 (1 M in holotype ); acropygium rounded ( Fig. 29A ); on lateral side with 3+3 M (L1, 3, 5) on carinae; on ventral side with four rows of 3+3 M, 3+3 M, 2+2 M and 2+2 M from anterior to posterior position. Lateral urotergites I‒IV with blunt, slightly rounded to pointed posterolateral angles; angles on urotergites V and VII slightly more conspicuous with small point; on urotergite VIII and urite IX with round end ( Fig. 24E‒F ). Urosternite I ( Figs 25C , 28A ) with ms and s setae; s setae being so abundant as to define a lateral field of up to 150 on each side of the posterior half of scutum, previous to the lateral subcoxal organs; prescutum with 3+3 M and scutum with 11+ 11 M. Medial glandular organ with 2+2 minute setae or no visible setae. Lateral subcoxal organ with one to three rows of glandular setae (GS) with about 140‒180 in males and 60‒70 in females and one row of sensory setae (SS) with about 60‒90 in males and 30 in females; lateral subcoxal organ occupying 0.30× of interstylar area (in males and females); GS/st1 (stylus of first sternite) = 0.2‒0.3 (males) and 0.35‒0.45 (females), SS/st1= 0.12 (males) and 0.2 (females) ( Fig. 28A ). Urosternites II‒VII with a few ms and strong M setae; urosternites II‒VII with three rows of: 7+7 A M, 4+4 B M and 5+5 C M; urosternite VIII with: 2+2 A M, 2+2 B M and 3+3 C M ( Fig. 25D ). Genital papilla in male with large lateral appendages ( Fig. 25F ). Cerci strong, well-developed, elongate, straight along the proximal half and curved in the distal half, becoming a small hook towards apex ( Figs 29–31 ); length ranging from 3 mm in largest specimen (Ô2- paratype ) to 2.9 mm in holotype and ♀ - paratype , always slightly longer than urite X; cerci heavily sclerotized, with dorsal and ventral outer carinae arising from dorsal and ventral acetabular joints; carinae extending almost to apex ventrally and halfway dorsally. Cerci asymmetric with a distinct concave top side with distal end slightly upward. Right cercus with round or pointed medial tooth; predental margin with two rows of 3+3 small, round denticles; postdental margin with a row of 14‒16 small denticles reaching near the hook. Predental margin of left cercus with two rows of denticles apart mostly on the middle; superior row with 14 small round denticles ending before the postmedial denticle smaller than the right cercus; inferior row with 12 small round denticles ending in the postmedial tooth; postdental margin with a row of 12 tiny denticles ending before the hook. Right/left cercus with about 12/15, 15/12, 20/25 dorsal, lateral and ventral long and short M; campaniform sensilla distributed mainly on hook and scarce on inner margins ( Figs 29−31 ). We have included cerci morphology observations in a video production, an animated movie produced with scientific supervision by José Antonio Peñas (see Supp. file 1). Fig. 27. Teljapyx aotearoa Sendra & Sánchez-García sp. nov. , holotype, Ô (AI). A . Pro-prosternum (pr-prt) with spine (sp), prosternum (pr-st), and meso-intersternum (ms-ist). B . Mesosternum (ms-st). C. Meta-intersternum (mt-ist.) and metasternum (mt-st). Fig. 28. Teljapyx aotearoa Sendra & Sánchez-García sp. nov. , ♀1-paratype, ♀ (MZB). A . Lateral posterior portion of the first urosternite. B . Detail of cuticle of first urosternite. Abbreviation: St = stylus. Fig. 29. Teljapyx aotearoa Sendra & Sánchez-García sp. nov. , ♀1-paratype, ♀ (MZB). A . Abdominal segment X and cerci, dorsal side. B . Cuticle detail of cerci, dorsal side. Fig. 30. Teljapyx aotearoa Sendra & Sánchez-García sp. nov. , ♀1-paratype, ♀ (MZB). A . Abdominal segment X and cerci, dorsal view. B . Left cercus, inner side in dorsal view. C . Right cercus, inner side in dorsal view. D . Abdominal segment X and cerci, ventral view. E . Right cercus, inner side in ventral view. F . Left cercus, inner side in ventral view. Taxonomic affinities The taxonomical features of Teljapyx aotearoa Sendra & Sánchez-García sp. nov. in cerci morphology, coxal organs of the first urosternite and pectinate maxillae match the original description of the genus Teljapyx Silvestri, 1949 . Teljapyx was proposed for two species from South America ( Silvestri 1949 ): Teljapyx riestrai Silvestri, 1949 and Teljapyx megalocerus , an already described species ( Silvestri 1905 ). Paclt (1957a) in his monography of the group proposed to include within the genus Teljapyx a total of 19 species from five other genera: Japyx Haliday, 1864 ; Catajapyx Silvestri, 1933 ; Sinjapyx Silvestri, 1949 ; Proncojapyx Silvestri, 1949 and Congjapyx Pagés, 1954 . Furthermore, Smith (1959) included another species but using different criteria from those presented by Silvestri (1949) and Paclt (1957a) , that is, the presence of a pair of predental denticles in both cerci. Teljapyx aotearoa has close morphological and geographical similarities to Teljapyx leai (Silvestri, 1930) , described from Tasmania and later found in mainland Australia ( Womersley 1939 ). Several differences between T. leai and T. aotearoa can be observed. First, body size and appendages are larger and more elongated in T. aotearoa than in T. leai : T. leai with a body length of up to 13.2 mm and a cerci length of 1.2 mm vs up to 29 mm body and 3 mm cerci in T. aotearoa . Second, T. leai has 30 antennomeres vs 32 antennomeres in T. aotearoa ; the urotergite X in T. leai has 3+3 D1–3 intracarinal dorsal macrosetae and T. aotearoa has 2+2 D1, 3 but 1 or 2 sagittal between D1 macrosetae; and the right cercus has a predental margin with two rows of 1+3 denticles in T. leai vs 3+3 denticles in T. aotearoa . Fig. 31. Teljapyx aotearoa Sendra & Sánchez-García sp. nov. , ♀1-paratype, ♀ (MZB). A . Distal part of left cercus, latero-internal view. B . Central part of left cercus, latero-internal view. Fig. 32. Location and habitat of Teljapyx aotearoa Sendra & Sánchez-García sp. nov. A . External area in the surroundings of the Council cave. B . Council cave entrance. C . General view of the interior of the Council Cave. D . Living specimen of Teljapyx aotearoa . Habitat Council Cave is located near the Motupipi settlement in Golden Bay, on a small limestone outcrop approximately 1 km long and less than 100 m wide exposed by the uplift of the nearby Pikikiruna Fault. The Motupipi limestone is presumed to be continuous under the bed of Dry River with a larger block immediately north of Dry River. It also continues beneath overlying geologies to the west, probably for a considerable distance. Almost the entire Motupipi limestone outcrop, including the section containing Council Cave, is overlain by the Council Cave Scenic Reserve. The reserve still maintains a reasonable amount of native vegetation ( Fig. 32A ) although it is heavily covered by exotic species. Council Cave is the largest known cave in the outcrop. The main entrance ( Fig. 32B ), at the northernmost point of the cave, is located close to a small road lying at about the level of the pasture, which extends from the western side of the limestone. The cave has a main, mostly dry, horizontal passage 170 m long that runs parallel with the edge of the limestone. This passage has suffered much damage in the past, from speleothem breakage to extensive graffiti. A second parallel conduit, which frequently holds a small stream, lies below and west of the main passage and has been explored up to at least 20 m beyond the end of the upper level. This lower conduit is often inaccessible due to water rising to roof level in many low areas. The main conduit of Council Cave presents several speleothems ( Fig. 32C ), although the floor is mainly covered by sediments. Specimens of Teljapyx aotearoa sp. nov. were observed, especially at the deeper regions of the cave. During the samplings, specimens were observed freely walking on the cave floor, notably on areas covered by sediments ( Fig. 32D ). Individuals moved slowly by feeling the substrate with their long antennae. Specimens seem not to react to light, so that these organisms apparently do not show phototaxy. No specimens were observed close to each other. When disturbed (by the touch of a brush) they moved quickly, trying to hide in cracks or spaces under rocks. It is important to note that Council Cave is one of the most important caves in New Zealand for its cave-restricted fauna, which has beetles, harvestmen, diplurans, centipedes, and spiders ( Santos et al. 2019 ). Fortunately, the cave is now protected by its scenic reserve status, and visitors can only access its main cave conduit via a locked gate. The key is held at the local office of the Department of Conservation, thereby ensuring sufficient protection. In this way, the Department is aware of who enters the cave and for what reason. At present, there is no official permit system for entry, although it could be implemented in the future.