Distinctive Collembola Communities in the Mesovoid Shallow Substratum: Entomobryomorpha of the Sierra de Guadarrama National Park (Central Spain)

ABSTRACT The material for this study was obtained after intensive sampling in the colluvial mesovoid shallow substratum, or MSS, of the Sierra de Guadarrama National Park using 33 subterranean sampling devices (SSD). The data were obtained from the first extraction of the traps between May and October of 2015. This paper presents the results for the Entomobryomorpha Börner, 1913, which was part of the Collembola captured. Four families and 12 genera have been studied: Isotomidae Schäffer, 1896 (Folsomia Willem, 1902, Tetracanthella Schött, 1891, Uzelia Absolon, 1901, Folsomides Stach, 1922, Isotomurus Börner, 1903, Parisotoma Bagnall, 1940, Pseudisotoma Handschin, 1924 and Pachyotoma Bagnall, 1949), Orchesellidae Börner, 1906 (Orchesella Templeton, 1835 and Heteromurus Wankel, 1860), Entomobryidae Schäffer, 1896 (Entomobrya Rondani, 1861) and Lepidocyrtidae Wahlgren, 1906 (Lepidocyrtus Bourlet, 1839 and Pseudosinella Schäffer, 1897). The species of Orchesella were studied in a previous paper (Baquero et al. 2017). The richness of the habitat sampled is defined by twenty-one species, eight of which are new: Pachyotoma penalarensis Baquero & Jordana n. sp., Entomobrya guadarramensis Jordana & Baquero n. sp., Entomobrya ledesmai Jordana & Baquero n. sp., Lepidocyrtus labyrinthi Baquero & Jordana n. sp., Lepidocyrtus paralignorum Baquero & Jordana n. sp., Lepidocyrtus purgatori Baquero & Jordana n. sp., Pseudosinella valverdei Baquero & Jordana n. sp. and Pseudosinella gonzaloi Baquero & Jordana n. sp. Entomobrya intermedia Brook, 1884 (England) is discussed and a new name Entomobrya katzi Jordana & Baquero n. sp. is proposed for E. intermedia sensu Katz et al. (2015) based on the American specimens.


INTRODUCTION
with an elongated body and conspicuous segmentation, three thoracic and six abdominal segments (some Isotomidae have four or five abdominal segments), prothorax not developed and without tergal chaetae. There are some habitual dwelling inhabitants of the ground and litter, but less specialized than Poduromorpha Börner, 1913. While more than 265 species have been found in the entire Iberian Peninsula since the publication of the catalogue by Jordana et al. (1990), five families with 21 genera and 59 species -seven of which were originally described in the Sierra de Guadarrama -have been found in the study area. This information is shown in 12 publications published between 1929published between and 1995published between (Cassagnau 1954Steiner 1955;Selga 1961Selga , 1962aSelga , b, 1963Selga , 1966aSelga , 1966bSelga , 1971Simón 1971;Simón & Selga 1977 [Somosierra]; Acón 1980). All those researchers worked intensely in the area because of the proximity of the National Museum of Natural Sciences and major university centers in Madrid.
The milieu souterrain superficiel or mesovoid shallow substratum (MSS), consists of a network of interstices and fissures in the subsoil, and harbors diverse epigean species of a stenoic nature, and strictly hypogean species that permanently inhabit this environment (Gers 1992;Ortuño et al. 2013). Previous studies focused on ecology (Juberthie et al. 1980;, while others explored some faunal aspects (Růžička et al. 1995;Nitzu et al. 2010;Jiménez-Valverde et al. 2015). This paper is a continuation of two previously published papers that study the biodiversity of the MSS collembolan fauna of the Sierra de Guadarrama (Baquero et al. 2017;Jordana et al. 2020). All these studies document the importance of the MSS biocenosis, demonstrating the enormous potential of this subterranean habitat as a refuge for fauna, and constitute a good tool for the management of natural spaces.

Site
The sampling was conducted in the Sierra de Guadarrama National Park, which is in the eastern half of the Central System (i.e., the Iberian Peninsula) and consists of an area of 33 960 hectares, surrounded by a peripheral buffer zone of 62 687.26 hectares (MAPAMA 2017). The mountain range on which the Sierra de Guadarrama National Park is located, is formed by three mountainous axes (Siete Picos, La Mujer Muerta, Montes Carpetanos, and Cuerda Larga and associated mountainous complex) that converge at two mountain passes, those of Navacerrada and Los Cotos (Fig. 1A). The lithology is dominated by the presence of orthogneiss (Vialette et al. 1987; PNSG a), a metamorphic rock. In the Sierra de Guadarrama, the fragmentation and accumulation of these rocks originated from glacial (Pedraza & Carrasco 2005) and periglacial events (Sanz 1986). Almost the entire study area has numerous scree slopes that allow the development of the MSS. The climate is Mediterranean, with marked continentality. As a general rule, summers are cool and dry, and winters are cold. However, the diverse topography of the mountains belts favors a considerable variety of microclimates (PNSG b;Salazar Rincón & Vía García 2003;JCL & CAM 2010;Palomo Segovia 2012). The study area is divided into three bioclimatic zones: supra-Mediterranean, oro-Mediterranean and cryo-Mediterranean (Rivas-Martínez 1984;Rivas-Martínez et al. 1987). The most outstanding char-

Mots clÉs
Milieu souterrain superficiel (MSS), trappes d'échantillonnage souterrain, écologie, espèces nouvelles. acteristics of these bioclimatic zones in the Sierra de Guadarrama and their most conspicuous vegetation are summarized in Ortuño et al. (2019). On the scree slopes, the rupicolous plant and lichen species communities acquire special relevance (JCL & CAM 2010). Also of importance is precipitation in the form of snow, which is more intense in the cryo-Mediterranean and oro-Mediterranean scrub supra-forest zones, especially in areas that conserve snowfields for many months.

Methodology
Thirty three sampling points were established (Fig. 1B). The details describing the placement of the traps and the rest of the methodology for sampling the animals have already been described in Baquero et al. (2017). The UTM coordinates (datum WGS84) are given in Table 1, and in the typical localities of the new species described. The authors who performed the sampling included a team that consisted of V. M. Ortuño, E. Ledesma, J. D. Gilgado, A. Jiménez-Valverde, G. Pérez-Suárez and E. Baquero. Permits to collect samples were obtained from the appropriate authorities (General Directorate of Environment of the Community of Madrid and Territorial Service of the Environment of the Junta de Castilla y León). The traps (Table 1) were placed between 20 May 2015 and 9 July 2015, and the first series of samples was obtained between 17 September 2015 and 6 November 2015. The term "activity" is sometimes used in the study instead of "abundance", as it better characterizes a quantitative community parameter obtained by the capture method, pitfall trapping. It means that more active forms of Collembola tend to be caught in traps, thus covering only a part of the species pool that occupies the MSS. After the preliminary triage to separate the Collembola Entomobryomorpha from the sampling fauna within the SSDs, some specimens were selected and mounted in Hoyer's medium for observation under a compound microscope in a phase contrast and DIC. Some specimens were cleared in Nesbitt's fluid. The remaining samples were stored in 70% ethyl alcohol.
In addition to the simplified formula of Jordana & Baquero (2005) as simplification of the dorsal macrochaetotaxy defined originally by Szeptycki (1979), the general color pattern (Katz et al. 2015) and some selected morphological characters (labral papilla shape, claw and empodium form, and mucro shape (Christiansen 1958;Christiansen & Bellinger 1980;Soto-Adames et al. 2008;Jordana 2012) have been used for the identification of the Entomobrya species. The macrochaetotaxy for Pseudosinella follows Gisin & Da Gama (1969), Szeptycki (1979), Mateos (2008 and Soto-Adames (2010). The characters defined by Christiansen et al. (1990)  Each of the 33 sampling points has been analyzed in terms of activity and specific diversity (Fig. 1E, F), and for this purpose, the following correspondence was established: sampling point = SSD (1 m). Data on the presence of a species regis-tered in an SSD (0.5 m) were incorporated in only one case because in the corresponding SSD (1 m) the species did not occur, and given the evidence of presence at the site, it could not be excluded from the calculations of specific diversity.  reMArkS Until now, the distribution of this species was limited to the Pyrenean and pre-Pyrenean area (Potapov 2001), although it has been found in the Cantabrian Mountains (Santamaría et al. 2012). This new record adds a new biotope for the spe-cies, also in the mountains, but in the center of the Iberian Peninsula. The diagnostic characteristics are totally coincident with the original: antenna (Fig. 5A), head (Fig. 5B) and general chaetotaxy ( Fig. 5C-E). Confirmation of the identity of the specimens found in the Sierra de Guadarrama was performed by comparing them with the type series specimens, and based on the presence of the three special long sensilla of the Abd V, which, in this case, are blunt (Fig. 5C, F), and the sensory chaetotaxy given by Potapov (2001), which is identical (Fig. 5F).
Since Potapov (2001) pointed out that this could be a synonym of F. trisetata due to the coincidence of the manubrial chaetotaxy (13 chaetae on F. sexoculata var. iberica Steiner, 1958 and 11-12 F. trisetata), it would be necessary to study Steiner's material (because at least seven Folsomia species have 13 chaetae in the manubrium ventral side).

reMArkS
Originally described from the Sierra de Guadarrama (Steiner 1955), according to Deharveng (1987), its distribution extends from the south of the Ebro River (Iberian Peninsula) to the Atlas Mountains in Morocco. Throughout Europe (de Jong et al. 2014), it is present from Bulgaria, North Africa, Portugal (mainland), Spain (mainland) and Ukraine. The citations of Cassagnau (1959) and Selga (1966aSelga ( , 1971 probably refer to the nearby species T. similis Deharveng, 1987.  reMArkS Already cited in the Sierra de Guadarrama by Acón (1980). Originally described in Germany (Schäffer 1896), it is considered a Holarctic species (Potapov 2001). In this study, only one specimen has been collected and it therefore appears to be occasional in the MSS.
reMArkS It had already been cited in the Sierra de Guadarrama by Cassagnau (1954  etyMology. -The specific epithet 'penalarensis' refers to the presence of this species in the Peñalara massif, which boasts the highest peak of the Sierra de Guadarrama.
deScription Body Size 0.72-0.80. Color dark blue. Integument granulated without reticulation. 6 + 6 to 8 + 8 eyes (sometimes eyes G and H disappear, but it is possible to see the refringent structures below). PAO with four lobes, two times eye A (Fig. 6A). Antenna as in Figure 6B, C; Ant III sensory organ with the central sensilla more or less spherical; Ant IV with seven sensilla, six dorsoexternal and one dorsointernal. Maxillary outer lobe bifurcated and four sublobal hairs (Fig. 6D). Labral formula 4/5,5,4 (labral chaetae papillated). Labium with four basomedial, three proximal and five basolateral chaetae and, as common for the family, with 16 guard chaetae.
ecology So far, this species has only been located in the MSS of the northern slope of the Peñalara massif, in SSD-8, installed in the Canchal de la Majada Aranguez (Figs 1A, C; 3A, B). This site is located at altitudes that exceed 2000 m a.s.l., and is part of the . As a whole, the syntopy of the five species at this site has provided an average relative activity that does not reach a thousand specimens (Fig. 1E). This species only represents 2% of the total Entomobryomorpha studied in this paper (Fig. 1D), but accounts for 65% of the total Isotomidae collected ( Fig. 2A, C).

reMArkS
Considering the group of characters for the family, the 6 + 6 ocelli and a very specific PAO are enough to establish the specimens found as a new species, assigned to the Pachyotominae subfamily due to the absence of anal spines, the presence of furca, fewer than four chaetae in the anterior part of the manubrium, granulation of the body, abundant sensory chaetotaxy, dens with teeth and absence of Mc. The new species really has an extraordinary shape of PAO compared with congeners and Isotomidae as a whole. The characters that are used for the separation of the different genera of the Proisotominae s.l. seem to have a low diagnostic value. It is probable that all this taxonomy is artificial and it will take further work to update definitions of the genera and probably also the subfamilies of all Isotomidae. The separation of the specimens of this sampling into a new species according to the number of eyes, PAO and presence or not of the tenent hair on the tibiotarsus can be seen in Table 2, which includes the eye number, PAO shape, PAO/eye ratio, empodial terminal filament presence and shape, anterior manubrium chaetae number, posterior manubrium chaetae number, anterior dens chaetae number, posterior dens chaetae number, tenaculum teeth number, tenaculum chaetae number and mucro teeth number and shape. This table is an example of the absence of differential generic characters for the subfamily Proisotominae s.l.

Head
Eight eyes, GH smaller than EF. Antennae length 1.36 mm, 2.08-2.92 times the length of the head (n = 4; the antennae have suffered the sampling method by the time the specimens have been in the polyethylene glycol); Ant IV with simple apical vesicle and pin chaeta present; sensory organ of Ant III with the special rod-like sensilla, and three additional guard sensilla (Fig. 9C); relative length of Ant I/II/III/IV = 1/2.08/1.95/1.89 (n = 3). Prelabral chaetae ciliated. Labral papillae multispinose. Lateral process of labial papilla E 1/3 shorter than the papilla, not reaching their apex. Maxillary palp bifurcated, with three sublobal chaetae (Fig. 9B).

Body and legs
Length ratio of Abd IV/III = 5.30 (between 3.85-6.43; n = 10). Trochanteral organ with approximately 35 chaetae (Fig. 9H). Tibiotarsus not sub-segmented, without smooth chaetae, except for smooth terminal chaeta on legs III. Claw with four teeth: paired at 50% and first unpaired at 75% from base; dorsal teeth not basal (Fig. 9I). Empodium lanceolate, with smooth external lamella (pe) in leg III. Tenent hair clavate. Length of manubrium and dens 0.53 and 0.66 mm, respectively (average for n = 9). Manubrial plate with four or five chaetae and two pseudopores. Mucro with teeth similar in size, mucronal spine reaching the tip of the subapical tooth (Fig. 9J); non-crenulated area of dens two times the length of mucro. Body chaetae as in Figure 9G. ecology Species widely distributed in the three mountain ranges (Fig. 1B, C). It is present in the MSS of the three bioclimatic zones and its overwhelming implantation in the subsoil, and the fact that it has never been registered as epigeous, suggests that it is a regular inhabitant of this habitat. It was extraordinarily abundant in SSD-27 of the Canchal de Bailanderos (Figs 1B, C; 3C, D), located in the cryo-Mediterranean zone. At this site 1034 specimens of E. guadarramensis Jordana & Baquero n. sp. were collected; thus, almost all of the Entomobryomorpha (1039 not including Orchesella) found from SSD-27 (Fig. 1E) evidence the dominance of this species over the other three species of syntopic Entomobryomorpha (Figs 1F; 3D).
deScription Size and color Body length: 2.12 mm, up to 2.33 mm (n = 7), excluding antennae. Ground color white or very pale yellow, with pigment as small patches as in Figure 8B; head with pigment between eyes and vertex, and on antennae on internal Ant I and from Ant II to IV.

Head
Eight eyes, GH smaller than EF. Antennae length 1.10-1.43 mm, 3.20 times the length of the head; relative length of Ant I/II/III/IV = 1/2.64/2.40/2.37 (n = 7); sensory organ of Ant III with the special rod-like sensilla, and three additional guard sensilla (Fig. 10B); Ant IV with apical vesicle bilobed. Prelabral chaetae ciliated. Labral papillae multispinose (Fig. 10C). Lateral process of labial papilla E not reaching the apex of the papilla. Labial chaetae ciliated: only one M, and R half of a M.

Body and legs
Length ratio of Abd IV/III = 4.27 (n = 7). Microchaetae on body relatively broadened (Fig. 10G). Tibiotarsus sub-segmented, without smooth chaetae, except for smooth terminal chaeta on legs III. Claw with four teeth: paired at 50% and first unpaired at 70% from base; dorsal teeth not basal, in an intermediate position between base and paired internal teeth (Fig. 10H). Empodium lanceolate, with serrate external lamella (pe) in leg III. Tenent hair clavate. Trochanteral organ with approximately 22 chaetae (Fig. 10I). Length of manubrium and dens 0.44 and 0.55 mm, respectively. Manubrial plate with four chaetae and two pseudopores. Non-ringed part of dens two times the length of mucro; mucro with teeth similar in size, mucronal spine reaching the tip of the subapical tooth.
Macrochaetotaxy (Fig. 10A, D-F Katz et al. (2015) found an Entomobrya captured in Chester (USA) and ascribed the specimen to E. intermedia. Some specimens of E. intermedia from England were studied for the review of the palearctic Entomobryinae (specimens from different parts of the United Kingdom sent to Rafael Jordana by Peter Shaw) (Jordana 2012). The coloration of both populations is similar but the macrochaetotaxy is different: H4 (Jordana & Baquero 2005) has three chaetae in the specimens from England, one (sometimes an additional mes) in the American specimen; A2 has four Mc in the English form (m 3 , m 3ep , m 3e and m 3ei ), two in the American form (m 3 and m 3e ); A7 has more Mc in the English form than in the American form; in addition, the labral papillae are smooth in the English specimens and multispinate in the American specimen. Given these differences and the geographical origin of the specimens, we consider E. intermedia sensu Katz et al. (2015) to be a new species of Entomobrya from the USA, denominated Entomobrya katzi Jordana & Baquero n. sp., with an abbreviated formula 3-2-0-1-2/3-5/2-2/1-0-1/0-4-0-2-2. Table 4 shows that the new species differs by multiple characters from the species with which it shares the simplified formula of Abd II-III.

Head
Antennal head ratio 1.58 (n = 6). Ant IV without apical bulb, apical organite and accessory sensilla as in Figure 11B. Ant III sense organ with two curved and expanded sensilla, one of them bigger than the other (Fig. 11C) three spiny guard sensilla, one of them blunt; on Ant II one distal similar but straight to Ant III expanded sensilla; Head Mc P a5 present, A 0 , A 2 and A 3 as ciliated Mc, A 2a as mes; t, s and p chaetae present on ocular well (p as mes, bigger than the other), three scales in the area; head dorsal chaetotaxy (Fig. 11A) with 5-8 antennal (An) ciliated Mc basomedian labial fields chaetae smooth. Four prelabral ciliated chaetae; labrum with three rows, 'a' row with four apically ciliated chaetae, 'm' and 'p' with five smooth chaetae (Fig. 11D). Four labral papillae, conical or with a spinelike chaeta. Maxillary palp bifurcated with three smooth appendages (Fig. 11E). Labial papilla (l.p.) E with finger-shaped process not reaching at base of apical appendage (Fig. 11F). Labial row with M 1 , M 2 , R*, E, L 1 and L 2 ciliated Mc (R half to two-thirds of M; M 1 sometimes absent and usually asymmetric) (Fig. 11G). Postlabial chaetotaxy with 3 + 1 ciliated central Mc along the groove. 12 + 12 spinelike chaetae on posterior dorsal head.
Thorax chaetotaxy (Fig. 12) Th II and Th III without Mc; Th II with 's' and 'ms' in posterolateral position at level of m row; Th III with two 'a' mic before psp, a 2 , a 3 , a 5 , a 6 , m 2 (above psp), m 3 , m 4 , p 2 -p 6 , and on lateral tergite a mes with the lateral sensilla (sl) interiorly.
Abdomen chaetotaxy (Figs 12; 13) Abd I with a 1 before psp; m 1 beside psp; a 3 , a 6 , m 3 , m 4 , m 6 and p 5 (with the 'ms' near a 3 ). Abd II, mi and ml chaetae present over bothriotrichum (m 2 ); a 2p (p) present as smooth mic; a 2 (a) as smooth mic; m 3 (B) present as ciliated Mc; 'as' over m 3 and a 3 upside over a 2 (two times its length); m 3e and p 4 (q 1 and q 2 ) present as smooth mic; lm and ll present as pointed ciliated mic over bothriotrichum (a 5 ); a 6 , m 4 , m 6 and p 5 as smooth mic; m 5 as Mc. Abd III, mi, ml and a 2 as pointed ciliated mic over bothriotrichum (m 2 ); 'as' between a 2 and m 3 ; m 3 as smooth mic; a 3 very up; p 3 below m 3 , and m 4 as smooth mic; lm, li, ll and a 6 as ciliated pointed mic surrounding bothriotrichum (a 5 ); im, em and am 6 as small ciliated mic over m 5 bothriotrichum; pm 6 and p 6 as ciliated Mc with d 3 between them; 'ms' near p 5 as smooth mic; p 8p as ciliated mes; a 7 , a 8 , m 7 , m 8 , p 7 and p 8 as smooth mic. Abd IV with four median mac (C 1 , B 4-6 ; ratio between C 1 -B 4 /B 4 -B 6 0.60-0.74, n = 3), and 6 lateral Mc (E 2-4 , F 1-3 ); T 5 as mic, D 3 , T 6 and T 7 as mes (D 3 as Mc in some specimens); before T 2 bothriotrichum, usually, three pointed ciliated mic (a, m and D 1 ), with a supplementary 's' chaeta present in only one specimen (♂) and asymmetric (Fig. 13).
ecology Species widely distributed in the three mountain ranges, found in the MSS of more than half of the sampling points ( Fig. 1A-C). Although it is present in the three bioclimatic zones, given the average catch and its frequent occurrence, it is more common with increasing altitude. Nevertheless, its greatest activity was recorded in SSD-6 of Canchal La Pedriza (Fig. 3G, H), located in the oro Mediterranean forest zone, and accounts for 32% of the 234 Entomobryomorpha (not including Orchesella) collected there (Fig. 1E). At this site, L. labyrinthi Baquero & Jordana n. sp. is syntopic with seven other species (Figs 1F; 3H) of the group analyzed in this study.
The wide distribution of L. labyrinthi Baquero & Jordana n. sp. does not correspond to the activity records, since it represents only 1% of the Entomobryomorpha and Entomobryidae, studied in this work, (Figs 1D; 2A, B).  deScription Size and color Body length up to 2.40 mm including head (mean 1.57 mm, n = 27 adults), excluding antennae (holotype: 2.10 mm). Color white with blue pigment on Ant III-IV and tip of Ant II; blue pigment on vertex of head and ocular patch (Fig. 8F). Scales present on Ant I-II, ventral and dorsal head, thorax and abdomen dorsally, coxae II-III and femora-tibiotarsus I-III, and furcula dorsally and ventrally.

Head
Antennal head ratio 1.5 (n = 4). Ant IV without apical bulb, four types of sensilla (Fig. 14A), and apical organite and accessory sensilla as in Figure 14B; Ant III sense organ with two expanded sensilla, three spiny guard sensilla, s-blunt sens, ciliated and weakly ciliated chaetae (Fig. 14C); on Ant II two distal similar to Ant III expanded sensilla. Head Mc Pa 5 present; A 0 , A 2 and A 3 as Mc, A 2a as ciliated mes; 5-8 antennal (An) ciliated Mc; s, t and p chaetae present on ocular well (p as mes) (Fig. 14D); basomedian labial fields chaetae smooth. Four prelabral ciliated chaetae (only one specimen among 42 observed has smooth prelabral chaetae); labrum with three rows, 'a' row with four bifurcate chaetae, 'm' and 'p' with five smooth chaetae. Four labral papillae, mono to three spinulated (small projection, not a relatively large chaetalike projection). Maxillary palp bifurcated with three smooth appendages. Labial papilla (l.p.) E with fingershaped process not reaching base of apical appendage. Labial row with M 1 , M 2 , R*, E, L 1 and L 2 ciliated Mc (R half to two thirds of M; sometimes M 1a or M 1p present and usually asymmetric). Postlabial chaetotaxy with 3 + 1 ciliated central Mc along the groove.
Thorax chaetotaxy (Fig. 15A, B) Th II and Th III without Mc; Th II with s and ms in posterolateral position at level of m row; Th III with a1 before psp, a 3 , a 4 , a 6 , m 2 , m 4 , m 5 and m 6 , p 2 , p 3 , p 4 , p 5 and p 6 , two lateral mes with the lateral sensilla (s) between them, and four Mc in front of the sensilla.

Abdomen chaetotaxy (Figs 15C-E; 16)
Abd I with a 1 before psp, and a 2 ; a 5 as; m 2 , m 3 , m 4 , m 5 and m 6 ; p 5 and p 6 , a sensilla in front of p 6 and m 6 , and three lateral mes. Abd II, mi and ml chaetae present over bothriotrichum (m 2 ); a 2p (p) present as smooth mic; a 2 (a) as smooth mic; m 3 (B) present as Mc; 'as' over m 3 and a 3 very up; m 3e and p 4 (q 1 and q 2 ) present as slightly ciliated mic; li, lm and ll present as pointed ciliated mic over bothriotrichum (a 5 ); a 6 , m 6 and p 5 as smooth mic; m 4 as slightly ciliated mic; m 5 as Mc. Abd III, mi, ml and a 2 as pointed ciliated mic over bothriotrichum (m 2 ); m 3 and m 4 as slightly ciliated pointed mic; 'as' before m 3 ; a 3 very up; p 3 below m 3 as smooth mic; lm, li, ll and a 6 as ciliated pointed mic surrounded bothriotrichum (a 5 ); im and em as small ciliated mic under a 5 bothriotrichum; am 6 as ciliated pointed mic over bothriotrichum (m 5 ); pm 6 and p 6 as ciliated Mc with d 3 between them; 'ms' near p 5 smooth mic; m 7a and p 8p as ciliated mes; m 7 , m 8 , p 7 and p 8 as smooth mic. Abd IV with four median mac (C 1 , B 4-6 ; ratio between C 1 -B 4 /B 4 -B 6 0.79, n = 32), and 7 lateral mac (D 3 , E 2-4 , F 1-3 ); T 5 as mic, T 6 and T 7 as ciliated mes; before T 2 bothriotrichum, there are usually three pointed ciliated mic (a, m and D 1 ); in a 20% is present the supplementary 's' chaeta. Abd V as in Figure 15E.
ecology Species widely distributed in the three mountain ranges (Fig. 1A-C), and present in the three bioclimatic zones. Only surpassed in distribution by E. guadarramensis Jordana & Baquero n. sp., and almost at the same time as H. major. From an altitudinal perspective, the average of collections per bioclimatic zone shows that the activity of L. paralignorum Baquero & Jordana n. sp. increases with altitude. However, this is due to the bias provided by two sampling points: 1812 specimens (SSD-22) and 3708 specimens (SSD-30), respectively, for this new species. La Loma de Pandasco (SSD-30), in the cryo-Mediterranean zone, is one of the places with the most extreme environmental conditions (Fig. 4A,  B). At this site L. paralignorum Baquero & Jordana n. sp. represents 99% of the total collected specimens (Fig. 1E), being syntopic with other four Entomobryomorpha species (excluding Orchesella) ( Fig. 1F; 4B) that were poorly represented.
reMArkS Lepidocyrtus paralignorum belongs to the L. lignorum group as the previous species. With regard to the shape of the labral papillae, it is separated from L. peisonis and L. ruber by smooth papillae, and from L. traseri, L. tellecheae and L. uzeli because they have a chaeta-like projection. Lepidocyrtus barbulus is separated from the remaining species by having the labral chaetae of row 'a' pointed instead of bifurcated, and, in addition to the chaetae, M 1 , M 2 and R are duplicated or triplicated, something not found in any other species of the group. Lepidocyrtus instratus and L. violaceus have only three teeth on the inner border of the claw (the last unpaired tooth missing). Lepidocyrtus juliae has a particular color pattern, with four dorsal spots on Abd II and Abd IV; it also has intraocular 'q' chaeta, and four scales in the area; Lepidocyrtus juliae does not have the chaeta d 3 in Abd III, and does not have ml in Abd II. Lepidocyrtus lignorum has all chaetae over bothriotricha fan-shaped and external lamella of empodium smooth. See Table 5.
It is the most abundant species, with capture records that account for 29% of the Entomobryomorpha studied here (Fig. 1D), and 30% of the Entomobryidae (not including Orchesella) (Fig. 2A, B).   deScription Body Body length up to 1.25 mm (holotype), head included (mean 1.05 mm, n = 6 adults), excluding antennae. Body pale violet blue, ocular spot black, antennae partially bluish from distal part of Ant I to tip, dorsal head slightly pigmented, Th II-Abd III with bluish bands (darker on Abd II-III), and an oval spot with a pale interior area on lateral Abd V (Fig. 8E). Scales absent on antennae, present on coxa, ventral and lateral manubrium, ventral dens, thorax, and abdomen; manubrium and dens similar in length (0.27 mm, n = 5); not annulated part of dens 4-5 times the length of mucro. Microchaetae on body with a particular aspect (Fig. 17J).

Head
Antennal head ratio 2.40 (n = 5). Ant IV with simple apical bulb, apical organite not capitate and accessory sensilla as in Figure 17A; Ant III sense organ with two curved and expanded sensilla (Fig. 17B) three spiny guard sensilla, one of them blunt. Four prelabral chaetae, lateral ciliated and central bifurcated and ciliated; labrum with three rows, 'a' row with four apically bifurcated chaetae, 'm' and 'p' with five smooth chaetae (Fig. 17C). Four labral papillae not visible or absent. Maxillary palp bifurcate with three smooth appendages. Labial papilla (l.p.) E as in Figure 17D with finger-shaped process reaching toward base of apical appendage. Labial row with M 2 , R*, E, L 1 and L 2 ciliated Mc (R half to two thirds of M). Postlabial chaetotaxy with one ciliated and two smooth central Mc along the groove (Fig. 17F). Head dorsal chaetotaxy with four antennal (An) ciliated Mc. A 0 , A 2 , A 3 , M 2 , S 3 and Pa 5 as Mc; R 1s (A 2a ) as mes; 4-5 Mc on series An (Fig. 17E); interocular chaetotaxy not seen.
Abdomen chaetotaxy (Figs 18,19) Abd I with a 1 before psp; a 2 -a 3 , a 5 -a 6 ('ms' near and external to a 6 ); m 2 (next to psp), m 3 -m 6 ; p 5 -p 6 . Abd II, mi and ml chaetae present over bothriotrichum (m 2 ); a 2p (p) absent; a 2 (a) and m 3 (B) present as ciliated Mc; 'as' over m 3 and a 3 upside over a 2 (1.5 times the length of as); m 3e and p 4 (q 1 and q 2 ) present as smooth mic; lm and ll present as pointed ciliated mic over bothriotrichum (a 5 ); a 6 , m 4 , m 6 and p 5 as smooth mic; m 5 as a very big ciliated Mc. Abd III, mi, ml and a 2 as pointed ciliated mic over bothriotrichum (m 2 ); 'as' between a 2 and m 3 , next to m 3 ; m 3 as ciliated mic; a 3 , m 3 and p 3 equidistant; p 3 below m 3, and m 4 as mes; lm, ll and a 6 apparently as ciliated mic surrounding bothriotrichum (a 5 ); im, em and am 6 as small ciliated mic over m 5 bothriotrichum; pm 6 and p 6 as very long and pointed ciliated Mc; d 3 absent; 'ms' near p 5 as smooth mic; m 8 as ciliated mes; m 7 and p 7 as smooth mic. Abd IV with four median mac (C 1 , B 4-6 ; ratio between C 1 -B 4 /B 4 -B 6 0.46, n = 5), and 7 lateral mac (D 3 , E 1-4 , F 2-3 ; E 2 missing in a ♂); T 5 as mic, T 6 and T 7 as mes; before T 2 bothriotrichum, usually, there are three pointed ciliated mic (a, m and D 1 ) (Fig. 19).

Legs
Scales only on coxae, not on rest of appendage. Trochanteral organ V-shaped with about 7 spine-like chaetae (n = 5) (Fig. 17G). Claw with four teeth on inner edge: basal pair at 50%, a unpaired median at 65% (highly developed), and one minute unpaired subapical; two big lateral teeth intermedial to base and paired, and dorsal at level of lateral. Empodium acuminate, 0.66 times the length of claw, with pe lamella serrated and other lamellae smooth (ae, ai, pi). Tibiotarsus III distally with one inner smooth chaeta reaching the tip of empodium and same size than claw; tenent hair spatulated, smooth, similar in size than claw (Fig. 17H).
deScription Body Body length up to 2.40 mm, head included (mean 2.05 mm, n = 11 adults), excluding antennae (holotype: 2.05 mm). Color blue dark, especially on Ant I-IV (except tip of IV), anterior part of the head, and posterior area of the tergites Th II-Abd VI), coxae, and basal manubrium; Th II darker in front area (Fig. 8G). Scales absent on antennae and legs, present on ventral and dorsal head, thorax and abdomen dorsally, and furcula only ventrally.
Thorax chaetotaxy (Fig. 21) Th II and Th III without Mc; Th II with s and ms in anterolateral position; Th III with a1 before psp, a 3 , a 4 , a 6 , m 2 , m 4 , m 5 and m 6 , p 2 , p 3 , p 4 , p 5 and p 6 , two lateral mes with the lateral sensilla (s) between them, and four Mc in front of the sensilla.

Abdomen chaetotaxy (Figs 21; 22)
Abd I with a 1 , a 2 and p 1 before psp; a 3 , a 4 , a 6 , m 2 , m 4 -m 6 ; p 1 -p 6 , a sensilla in front of m 6 , and some lateral mes. Abd II, mi and ml chaetae present over bothriotrichum (m 2 ) (sometimes an additional mic between ml and a 2 ); a 2p (p) present as slightly ciliated mic; a 2 (A) as small Mc or mes, but not mic; m 3 (B) present as Mc; 'as' over m 3 and a 2 , and a 3 a little above 'as'; m 3e and p 4 (q 1 and q 2 ) present as slightly ciliated mic; lm and ll present as slightly broadened at tip ciliated mic over bothriotrichum (a 5 ); m 4 as slightly ciliated chaeta; m 5 as mes. a 6 (smooth), a 7-8 , m 6 , p 5 -p 8 (slightly ciliated) as mic; Abd III, mi, ml and a 2 as slightly broadened ciliated mic over bothriotrichum (m 2 ); 'as' before m 3 that is apparently smooth; a 3 , m 4 and p 3 as slightly ciliated pointed mic; a 3 very up; im, li, lm and a 6 as ciliated pointed mic surrounding bothriotrichum (a 5 ); em, am 6 and a 7 as small ciliated mic under a 5 bothriotrichum (sometimes an additional mic near a 7 ); pm 6 and p 6 as Mc with d 3 between them (d 3 not always present, and duplicated in one specimen); 'ms' (d 2 ) near p 5 as smooth mic; m 7a and p 8 as mes; m 7 -m 9 , p 7 and p 9 as smooth mic. Abd IV with three median mac (C 1 , B 5-6 ; ratio between C 1 -B 5 /B 5-6 1.00, n = 9), and 7 lateral mac (D 3 , E 2-4 , F 1-3 ); T 5 as mic, D 2 , De 3 , E 4p , F 3p , T 6 and T 7 as mes; before T 2 bothriotrichum four ciliated mic (a, m, s and D 1 ) as in Figure 22; pi and pe as ciliated fan-shape mic.

Legs
Legs without scales. Trochanteral organ with near 40 spinelike chaetae. Claw with four teeth on inner edge: basal pair at 40% and 50% with respect to the internal claw edge length, respectively, first unpaired median at 70%, and one minute (sometimes imperceptible) unpaired subapical at 90%; two lateral teeth at 20%, and one more basal dorsal tooth. Empodium acuminate, all with non-serrated pe lamella (but with a small tooth on first third of all legs, and a minute serration ⦰ on legs 1 and 2), other lamellae smooth (ae, ai, pi); claw : empodium ratio = 1 : 0.65. Tibiotarsus III distally with one inner smooth chaeta 0.50 longer than claw; tenent hairs capitate, smooth, and 0.90 shorter than claw (Fig. 20E).

Furcula
Manubrium and dens with scales only ventrally, and with the same length; manubrial plate (dorsally) with three internal, approximately thirteen external ciliated Mc, and 2 psp (Fig. 20D). Non-ringed area of dens 2-3.5 times the length of mucro, with subapical tooth a little smaller than apical tooth (Fig. 20F).
ecology Species widely distributed in MSS of Montes Carpetanos and Siete Picos-La Mujer Muerta, not detected in Cuerda Larga ( Fig. 1A-C). According to the available data (presence and activity), it appears to show a preference for the subsoil of the oro-Mediterranean zone, with dominance in the forest strip. Its presence in the cryo-Mediterranean zone has not been verified and the upper level of this species is 2049 m a.s.l., which corresponds to SSD-10 installed in the Canchal Hoya de la Laguna Grande (supraforestal strip of the oro-Mediterranean zone). Its greatest activity was recorded in SSD-11 in the Canchal Cerro Ventoso (Fig. 4E, F), exceeding a thousand specimens (more than half of the Entomobryomorpha, excluding Orchesella, collected there (Fig. 1E). The MSS of this site, under the narrow influence of the pine forest (Pinus sylvestris), has revealed itself as one of the most diverse in Collembola, as it contains eight species of Entomobryomorpha (excluding Orchesella), with P. valverdei Baquero & Jordana n. sp. as the dominant species (Figs 1F; 4F).

reMArkS
The species that share the traditional formula of Gisin (1965Gisin ( , 1967a are, in addition to P. gonzaloi Baquero & Jordana n. sp., P. styriaca Neuherz &Nosek, 1975, P. subcentralis Gama, 1985 andP. valverdei Baquero & Jordana n. sp. Table 6 shows the differences between these four species. In terms of activity, P. valverdei Baquero & Jordana n. sp. is the fourth best represented species of Entomobryomorpha (excluding Orchesella) in the MSS, with 11% (Fig. 1D), and Entomobryidae with 12% ( Fig. 2A, B). diAgnoSiS. -Body with blue pigment, including antennae and first leg segments, as in Figure 8H. Head with 6 + 6 eyes (A-F); Mc A 0 , A 1 and A 2 present, A 1a present; t and p chaetae present (p as Mc); basomedian labial fields chaetae smooth; posterior labial row with M 1 , M 2 , R*, E, L 1 and L 2 Mc (R half to two thirds of M); two or three anterior postlabial chaetae as ciliated Mc. Th II-III without Mc; Abd II with chaeta a 2p present, a 3 forward from 'as' sensilla; a 2 as mes or short Mc, and m 3 as ciliated Mc; Abd IV with three median mac (C 1 , B 5-6 ), four ciliated (some fan-shaped) mic behind anterior bothriotrichum and bothriothrichal complex mic D 1p present; claw with four internal teeth: two basal and two unpaired; empodium acuminate; manubrial plate with 3 internal and 6-9 external chaetae.

Pseudosinella gonzaloi
deScription Body Body length up to 1.80 mm, head included (mean 1.55 mm, n = 8 adults), excluding antennae (holotype: 1.40 mm). Color blue dark, especially on Ant I-IV, head vertex, and posterior area of the tergites Abd I-Abd VI), coxae, dorsal and basal manubrium; Th II darker at front area. Scales absent on antennae and legs, present on ventral and dorsal head, thorax and abdomen dorsally, and furcula only ventrally.

Thorax chaetotaxy
Th II and Th III without Mc.
Abdomen chaetotaxy (Figs 24;25) Abd II, mi and ml chaetae present over bothriotrichum (m 2 ) (sometimes an additional mic externally to mi); a 2p (p) present as slightly ciliated mic; a 2 (a) as small Mc or mes, but not mic; m 3 (B) present as Mc; 'as' over m 3 , and a 3 above a 2 and m 2 ; m 2e and p 4 (q 1 and q 2 ) present as slightly ciliated mic; lm and ll present as slightly broadened at tip ciliated mic over bothriotrichum (a 5 ) (additional mic interior to a 5 bothriotrichum; m 4 as slightly ciliated chaeta; m 5 as mes; a 6 (smooth), a 7 -a 8 , m 6 , p 5 -p 8 (slightly ciliated) as mic; Abd III, mi, ml and a 2 as slightly broadened ciliated mic over bothriotrichum (m 2 ); a 3 , m 3 , m 4 and p 3 as smooth mic; li, lm and a 6 as ciliated mic above bothriotrichum (a 5 ); im and am 6 as small ciliated mic under a 5 bothriotrichum; pm 6 and p 6 as Mc with d 3 as slightly ciliated mic between them; 'ms' (d 2 ) near p 5 as smooth mic; m 7a and p 8p as mes; p 8 as small ciliated mic; a 8 , m 7 -m 9 and p 9 as smooth mic. Abd IV with three median mac (C 1 , B 5-6 ; ratio between C 1 -B 5 /B 5-6 109/99 = 1.10), and 7 lateral mac (D 3 , E 2-4 , F 1-3 ); T 5 and D 2 as mic; De 3 , E 4p2 , F 3p , Fe 5 , T 6 and T 7 as mes; before T 2 bothriotrichum four ciliated mic (a, m, s and D 1 ), some fan-shaped; pi and pe as ciliated fanshape mic.
ecology Species present in the MSS of the three mountainous ranges (Fig. 1A-C) but only verified from the supra-Mediterranean and oro-Mediterranean bioclimatic zones (in the forest strip), in the latter with records that suggest a more stable population. The largest number of specimens was collected with SSD-16, installed in the Canchal Las Revueltas-Los Horcos ( Fig. 4G-H), site with strong influence of the pine forest (Pinus sylvestris), and where P. gonzaloi Baquero & Jordana n. sp. accounted for 67% of the 895 Entomobryomorpha (excluding Orchesella) collected there (Fig. 1E). At this site, where this species is dominant, it is syntopic with four other Entomobryomorpha species (Figs 1F; 4H).

reMArkS
The species that share the traditional formula of Gisin (1965, 1967a, b) with this species are, in addition to P. valverdei Baquero & Jordana n. sp., P. styriaca Neuherz & Nosek, 1975 and P. subcentralis. Table 6 shows the differences between these four species.