The family Onychiuridae (Collembola) from karst caves of the Basque biospeleologic district, with description of four new species Author Beruete, Enrique 0000-0002-3240-8946 University of Navarra, Faculty of Sciences, Department of Environmental Biology, University Campus, 31080 Pamplona, Spain. eberuete @ yahoo. es; https: // orcid. org / 0000 - 0002 - 3240 - 8946 eberuete@yahoo.es Author Arbea, Javier I. c / Ría de Solía 3, ch. 39. 39610 El Astillero (Cantabria). Author Baquero, Enrique 0000-0002-3240-8946 University of Navarra, Faculty of Sciences, Department of Environmental Biology, University Campus, 31080 Pamplona, Spain. eberuete @ yahoo. es; https: // orcid. org / 0000 - 0002 - 3240 - 8946 & University of Navarra, Institute of Biodiversity and Environment (BIOMA), University Campus, 31080, Pamplona, Spain * corresponding author: ebaquero @ unav. es; https: // orcid. org / 0000 - 0002 - 2145 - 8606 eberuete@yahoo.es Author Jordana, Rafael 0000-0002-3240-8946 University of Navarra, Faculty of Sciences, Department of Environmental Biology, University Campus, 31080 Pamplona, Spain. eberuete @ yahoo. es; https: // orcid. org / 0000 - 0002 - 3240 - 8946 eberuete@yahoo.es text Zootaxa 2021 2021-09-21 5040 2 151 194 journal article 10.11646/zootaxa.5040.2.1 1175-5326 5531192 CC1289B1-4FF9-4369-A596-9FF260C8F314 Protaphorura leitzaldeaensis Arbea, Beruete & Jordana sp. nov. Figs 35−39 , Tables 8 and 14 Type locality. Lezegalde cave , Sierra de Aralar , Larraun, Iribas, Aralar N , Navarra , 611 m asl , coordinates 43.07937063 , -1.74705964 . Type Material. Holotype female, slide MZNA714610, Lezegalde cave ( Loc. 26 ), Sierra de Aralar , Larraun , Iribas , Aralar N , Navarra , 4.VIII.1987 , E. Beruete leg. Paratypes : 1 female (MZNA716391) and 1 juvenile (MZNA714609) with same locality and date as holotype ; 1 female (MZNA716390) with same locality and 7.IV.1982 . Other studied material . Leitza , Aralar N , Navarra , 900 m asl , oak forest, 11.III.1993 , R. Jordana et al . leg. 5 females on slides MZNA272676, MZNA272677 and MZNA272678 . Etymology. The specific name derives from the name of the type locality of this species, Norte de Aralar/Leitzaldea. Description. Color in alcohol yellowish white. Length (without antennae) 2.15–2.35mm in females; holotype 2.15 mm. Cylindrical body, with anal spines on papillae. Pso formula: 4,3/0,2,2/3,3,3,4,3 dorsally (female paratype with asymmetry in the pso of the Th III tergite: 2 on one side and 3 on the other), 1/0,0,0/0,0,0,0,0 ventrally ( Figs 35–36 ); ventrally with 1/0,0,0/1,1,1,1 psx and 1,1,1 psp in Th I–III; subcoxa 1 of legs I–III with 1,1,1 pso; subcoxa 2 of the legs with 1,1,1 psx, respectively. Head. Antennae approximately the same length as the head. Well-marked antenna bases. Ant IV with subapical organite and a lateral-external ms located 28–38 % of length from the base. AIIIO with 5 papillae, 5 guard chaetae, 2 small internal sensilla, 2 granulated sensory clubs and one lateral ms. Ant I−III with 11−12,19−22,18 ordinary chaetae, respectively. PAO compossed by 28–34 simple vesicles ( Fig. 38 ). Head with 4+ 4 p-chaetae between the postero-internal pso (p 1’ , P 1 , p 2 , P 3 ; Fig. 35 ). Mandible with a strong molar plate and 4 apical teeth. Maxilla with 3 teeth and 6 lamellae. Maxillary palp simple, with 1 basal and 2 sublobals chaetae. Labral chaeta formula 4/342. Labium type A, with 6 proximal, 4 basomedial (E, F, G and f) and 6 basolateral chaetae. 5−7+5−7 postlabial chaetae along the ventral groove ( Fig. 36 ). Dorsal body chaetotaxy as in Table 8 and Fig. 35 , generally with asymmetries, well differentiated in micro-/mesochaetae and macrochaetae; it is slightly plurichaetotic: multiplication of the number of microchaetae in row m and on lateral areas of the Abd I –IV tergites. Sensory chaetae little different from mesochetae, with blunt apex. Terguites Th II –III with lateral ms. Th I tergite with 19–20 pairs of chaetae (chaetotaxy type 1–2i, 3–4, 1– 2m ). Abd I –III and V tergites without chaeta s’ (s’ on Abd I is present in two females ) ( Fig. 35 ). Abd IV and V tergites with axial p 0 , AbdVI tergite with axial p 0 , a 0 generally present and m 0 rarely present ( Fig. 35 ). Ratio of chaetae M/sp lengths on AbdV 2.5–3.3. Ratio of chaeta M/ AS lengths 2.4–2.7. Lines (straight) passing through the bases of the prespinal chaetae a 1 and m 1 , parallel ( Fig. 35 ). AS short, on well-differentiated papillae, 0.5–0.7 times as long as the inner edge of the claw III . TABLE 8. Protaphorura leitzaldeaensis sp. nov. Dorsal chaetotaxy. Abbreviations: AS, anal spine; me, meso-; mi, micro-; M, macrocheta; s, chaeta-s (sensillum); “–“, absent; “()”, present or absent.

0	1	2	3	4	5	6	7
Head
a	me+M
d	–	mi	mi	M	3mi	–	–
sd	–	mi	mi	mi	me	mi	–
sd’	–	mi	mi	mi	mi	me	–
v	–	mi	me	mi	M	–	–
ca	–	me	–	me	–	–	–
cm	–	–	–	mi	mi	me	–
cb	–	mi	me	–	M	mi	–
cp	–	mi	–	–	mi	mi	–
p	–	mi+me	mi	M	mi(z)	M	mi
Th I
m	2mi	2mi	2mi	2mi	2mi	2mi	–
p	M	me	1–2me	me	M	1–2me	M
Th II−III
a	me	mi	me+(me)	mi	2me	mi	2me
m	mi	M	mi	me	me	M	2-4me+ms
ca	mi	mi	–	mi	mi	mi	–
cp	mi	mi	–	mi	me	–	–
p	mi	M	M	2mi	M	2–3me	M
Abd I
a	me	me	mi	mi	mi	2mi	me
m	2mi	M	–	me	2mi	2mi	M
ca	mi	mi	–	2mi	2mi	–	me
p	2mi	me	2s	M	2me	me	M
Abd II–III
a	me	mi	mi	2–3mi	2–4mi	2–4mi	3–4mi
m	2mi	M	–	M	2mi	2mi	M
ca	mi	mi	–	2mi	2mi	–	me
p	2mi	me	mi	M	2me	me+(me)	M
Abd IV
a	me	mi	2mi	4me	2mi	2me	6mi
m	1–2mi	M	2mi	2me	2me	M	3mi
ca	2mi	me	2mi	mi	mi	mi	12mi
cp	mi	mi	–	mi	2mi	2me	M
p	me	mi	M	s	me	me	M	2me
Abd V
a	me	M	me	me	me	me	M
m	me	me	mi	mi	mi	mi	mi
p	me	mi	M	s	M	2me	2me	M
Abd VI
a	me	me	me	s
m	(me)	mi	M
p	me	AS	M

FIGURES 35−36. Protaphorura leitzaldeaensis sp. nov. 35 , dorsal view; 36 , ventral view. Scale: 0.2 mm. FIGURES 37−39. Protaphorura leitzaldeaensis sp. nov. 37 , tibiotarsus and claw of leg III; 38 , PAO; 39 , anal valves. Scales: 0.05 mm (37−38), 0.1 mm (39). Ventral body chaetotaxy according to Fig. 36 . Th I–III, between the legs, with 1,2,2 pairs of chaetae, respectively. VT with 9−11+ 9−11 distal and 2 +2 basal chaetae. Furca reduced to a cuticular fold with 2+2 dental micro- chaetae: 1+ 1 located in the fold and 1+ 1 clearly located behind; three rows of posterior manubrial chaetae: rows ma, mm and mp with 3 +3 chaetae, of which the outer ones are macrochetes ( Fig. 36 ). Female genital plate with up to 47 chaetae plus two posterior microchaetae. Unknown males. Anal valves with numerous acuminate chaetae; each lateral valve with chaeta a 0 , 2a 1 and 2a 2 ; posterior leaflet with a 0 , 2a 1 , 2a 2 , 2b 1 , 2b 2 , c 0 , 2c 1 and 2c 2 ( Fig. 39 ). Legs. Subcoxae 1 of legs I−III with 5−7,6−8,5−7 chaetae, subcoxae 2 with 1,5−6,6, coxae with 4,12−13,13−16, trochanters with 10−11,11,11−13 and femurs with 19,21−23,20−22 chaetae, respectively. Tibiotarsi of legs I−III with 22−23 (11,8,3−4), 24 (11,8,5), 24−25 (11,8,5−6) chaetae, respectively, of which 2,2,2 are macrochetae with blunt apex located in the basal row (R chaetae) ( Fig. 37 ). Claw with inner tooth. Empodium narrow and pointed, without basal lamella, as long as the inner edge of the claw ( Fig. 37 ). Discussion. P. leitzaldeaensis sp. nov. belongs to the group of P. octopunctata species defined by Pomorski & Kaprus’ (2007) and Kaprus’ & Pomorski (2008) because it has 4 pso at the base of the antennae. Within this group, it is close to a set of Palearctic species that present 0,2,2/3,3,3 pairs of pseudocelli from the tergite Th I to Abd III and 1,1,1 pso in the subcoxa 1 of the legs I–III ( P. eichhorni ( Gisin, 1954 ) ; P. ianstachi ( Yosii, 1972 ) ; P. licheniphila Kaprus’ & Pomorski, 2008 ; P. lucensis Arbea, 2015 ; P. nutak ( Yosii, 1972 ) ; P. quadriocellata ( Gisin, 1947 ) ; P. altuaria (Pomorski & Kaprus’, 2007); P. suboctopunctata ( Khanislamova, 1986 ) ; P. tetragrammata ( Gisin, 1964 ) ; and P. valsainensis ( Acón, 1981 ) , whose main diagnostic differences are summarised in Table 14 . The new species is close to P. lucensis and P. tetragrammata for presenting the same number of dorsal pso, but it differs from them by the number of ventral psx (1,1,1,1 in the new species and in P. tetragrammata compared with 0,0,0,0 in P. lucensis ), the arrangement of the prespinal chaetae of AbdVI (parallel in the new species and in P. lucensis compared with convergent in P. tetragrammata ), a more pronounced plurichaetosis (6,6,6,5 pairs of axial chaetae in the Abd I–IV tergites in the new species compared with 4,4, 4,4 in the other two species), the short AS (ratio of AS /claw lengths 0.5−0.7 in the new species compared with 0.8−1.0 in the other two species) and its larger size (up to 2.4mm in the new species compared with 1.7mm in the other two species).