The high complexity of Micronetinae Hull, 1920 (Araneae, Linyphiidae) evidenced through ten new cave-dweller species from the Morocco

ABSTRACT The Museum of Natural Sciences of Barcelona is the repository of a set of samples of spiders captured during a prolonged period of time in various caves of the Moroccan Atlas. Seventeen species of Lepthyphantes Menge, 1866 “sensu lato” were found within this material, ten of them new to science: Lepthyphantes almoravidus Barrientos n. sp., Lepthyphantes biospeleologorum Barrientos n. sp., Lepthyphantes ensiferus Barrientos n. sp., Lepthyphantes fadriquei Barrientos n. sp., Lepthyphantes imazigheni Barrientos n. sp., Lepthyphantes lamellatus Barrientos n. sp., Lepthyphantes leknizii Barrientos n. sp., Lepthyphantes sasi Barrientos n. sp., Palliduphantes banderolatus Barrientos n. sp. and Palliduphantes megascapus Barrientos n. sp. We offer the description of the aforementioned species, as well as the description of the unknown male of Lepthyphantes taza Tanasevitch, 2014. We also discuss the affinities found between these and other related Mediterranean species, as well as their possible location within the Micronetinae genera currently described.


INTroDUCTIoN
Morocco is a mountainous country located in the Maghreb region of North Africa. Geographically, it is characterised by a lengthy Atlantic ocean coastline in the west, a Mediterranean Sea coastline in the north, and large tracts of desert and a rugged mountainous interior dominated by the Atlas mountain range (Fig. 1). The Atlas mountain range extends eastward, crossing Algeria and Tunisia, with some summits reaching 4000 meters high. The northeastern Atlas Mountains connect also with the rif mountain range in an east-west orientation. Within this mountainous complex, numerous karstic accidents occur, including caves that are largely unexplored and/or biologically understudied. However, a few caves have been the subject of attention by various biospeleologists' teams and researchers, especially since the second half of the 20th century. one of them is the Associació Catalana de Bioespeleologia (BIoSP), which has been carrying out expeditions to Morocco since 1970 with special attention given to the Central Atlas area. overcoming numerous difficulties, BIoSP campaigns to Morocco have continued to this day, with more than 200 Atlas mountain caves explored. Samples have been deposited in the Museu de Ciències Naturals de Barcelona and have remained unpublished until now.

The genus Lepthyphantes "sensu Lato"
The genus Lepthyphantes was created by Menge in 1866. For many years and still today, the genus has served as a "catchall" for many species of Micronetinae Hull, 1920 that share a similar morphology, the phylogeny then available being too unresolved to include them in other recently described genera. The current catalogue consists of 167 Holarctic and Ethiopian species (World Spider Catalog 2018), with the majority Palaearctic. Many of these species are waiting for a new analysis that would allow their taxonomic relocation.

The Lepthyphantes in Morocco
The countries of the Maghreb (and Morocco in particular) have been explored in an unbalanced manner. Many of the available data are due to biospeleological expeditions, ignoring the external fauna. Simon (1884), Machado (1940), Denis & Dresco (1957), Brignoli (1978) and Tanasevitch (2014) provide Moroccan data of this type. Exceptions to these circumstances are the works of Wunderlich (1980), Bosmans (1985Bosmans ( , 1991 and Bosmans & Bouragba (1992),

MATErIAL AND METHoDS
Among the material captured in the caves visited by BIoSP 751 specimens belong to Linyphiidae Blackwall, 1859. An important part of this material can be assigned to the genus Lepthyphantes "sensu lato" (60♂, 151♀, and 162 inmatures). All the material is deposited in the Museu de Ciències Naturals de Barcelona with the references that are detailed in Appendix I and in the section "studied material" for each species. The samples were obtained by direct capture. The material is preserved in 70% ethanol and was analysed according to normal methodology carried out through observation by binocular microscope (Wild M10).
For the new species and a few previously identified species a more exhaustive study protocol was followed, with individualization and clarification by means of 10% KoH of the genitalia. The separated pieces are conserved in microvials associated with the main sample or in permanent preparations with identical labelling. These manipulations were considered essential for the morphological analysis and illustrations. The interpretation of the structures follows the terminology proposed by Saaristo & Tanasevitch (1996) and assist by Hormiga (2000) for the whole Micronetinae; for a better understanding we will use the same abbreviations.

MeAsureMenTs
The measures of the legs are given in the form of a polynomial, whose terms correspond to the different pieces , i.e., (femur + patella + tibia + metatarsus + tarsus diAgnosis. -As in Bosmans (2006: figs 6-10) the males are easily recognized by the shape of their lamella characteristica; it is long and "C" curved, laminar, and with subtriangular and slightly toothed distal part. In its posterior median part the paracymbium presents a thick and sharp denticle and its anterior branch is irregular and digitiform. The females have an elongated epigyne, which narrows characteristically in its basal third and expands again in the distal part; the distal portion of the scape protrudes in the central area of the latter. According to our observations and the data provided here, L. aelleni could be related to other species of the Moroccan Atlas, such as L. maurusius, L. taza and L. almoravidus Barrientos n. sp.
disTribuTion. -Lepthyphantes aelleni is known from the chasm of Kaf el Bouk, from which comes the holotype (Denis & Dresco 1957) and from the cave of Friouato (Bosmans 2006). With the new data, the number of caves in which it is found is expanded; so that in addition to the Taza region, the species is also in the Eastern region. diAgnosis. -The epigyne is broad and small, protruding on the ventral side of the epigastrum. The bursa copulatrix has a very short front edge; the lateral edges expand and separate to accommodate the scape; the posterior wall differentiates a median plate, narrow in the center and rounded on the sides. The proscape is wide and flattened, with rounded contour and a notch in the anterior part; a short and rounded strecher in the distal part of the scape, but flanked by two rounded expansions. By the morphology of its epigyne (and without information on the male copulatory bulb) this species has some affinities with L. brevihamatus, L. longihamatus and, in general, with the entire "afer group", in which probably it should be placed.

Lepthyphantes almoravidus
eTyMoLogy. -This species is dedicated, in a general way, to all the biospeleologists. Anonymously they use part of their time to develop an exciting and hard (sometimes risky) activity, in pursuit of the forms that inhabit the caverns. Without their generous dedication, this small job would not have been possible.
disTribuTion. -This species is exclusively known from the type locality. Its basal part showing a wide bursa copulatrix, with a very short anterior wall; lateral walls expanding and separate to accommodate the scape; posterior wall wide and reinforced by a sclerotized median plate, narrowed in the center and rounded on the sides. Anterior and lateral walls with a thick, sparse and well-arched hairiness. Scape starting from the anterior part of the bursa, and with a wide and flat proscape, rounded on the sides and with a notch on its distal part (connecting with the middle part); fertilization ducts visible in submarginal position. Median part of scape inflected and hidden under the proscape; the distal part, tourning towards it again, dilating and showing below the proscape; strecher short and rounded, but flanked by two rounded expansions. Distal part usually curved outwards, so that its distal part approaches the proscape.

Male
Unknown. diAgnosis. -The characters indicated by Bosmans (1985) are especially useful. The male, although very close to that of L. longihamatus, can be easily distinguished by the shape of its suprategular process (simple and non-bifid); the paracymbium is strongly curled in its median part. The females can be distinguished more easily; the proscape is clearly rounded, without a median notch distally, and the strecher (distal part of the scape) is short and protrudes very little below. This species is clearly included in the "afer group".

Lepthyphantes brevihamatus
disTribuTion. -The holotype comes from the cave Ifri Bernat, Ait Mohammed, Azilal (Bosmans 1985) and has also been collected in the Ifri el Kaid cave (Tanasevitch 2014 diAgnosis. -The paracymbium is very broad and subquadrangular, due to the development of its middle part; it presents a pair of sharp denticles in its proximal area; the apical part is very short. In the bulb, the development and shape of the lamella characteristica stand out. It is a well-sclerotized process that is oriented forward and outward; it is clearly curved distally and shows a laminar structure in a semicircle like a sickle. The epigyne is very developed and outstanding on the ventral side, almost perpendicular to the surface of the body, due to lengthening of the basal area. The scape is large, but there is no special development of the proscape; the whole set describes a very characteristic "S" path that fills the back of the epigyne. If we stick to the structural features of the epigyne, L. ensiferus Barrientos n. sp. would be a species close to L. fagei Machado, 1939; but if we look at male genital structures (clearly illustrated for this second species in Hormiga & ribera 1990), the differences are evident. Consequently, L. ensiferus Barrientos n. sp. remains in an uncertain systematic position.
eTyMoLogy. -The name refers to the falciform aspect of the embolic division of the lamella characteristica, in the copulatory bulb; its "sickle" shape is especially visible in a lateral observation.
disTribuTion. -The samples come from a single cavity, located in the province of Bou Izakarn, south of Agadir.   Palp and copulatory bulb (Figs 4A-C; 13J). Paracymbium very wide, subquadrangular, due to the great development of its proximal part, very broad at base, subtriangular distally and without basal pilosity; middle part with a pair of acute denticles, quite close, in the posterior area, and little excavated in its anterior part; the apical part very short and narrowing in a straight fingering. Bulb ventrally and laterally emphasizing the development of the lamella characteristica; a consistent process, starting from the back of the radix, orienting forward and outward; slightly curved, and showing distally, a clearly laminar structure describing a semicircle, like a broad-leaved sickle (arrangement particularly clear in a retrolateral observation). on ventral side, the suprategular apophysis curved, sharp and well sclerotized visible at the anterior end. radix ( Fig. 4C) developping, in front of the lamella characteristica, another long and acute process, well sclerotized, and slightly curved, the terminal apophysis. Embolus located in front of the radix, forming three translucent and sublaminar points; one of them, embolus proper, allowing to glimpse the route of the canal, starting from Fickert's gland (fgl).
Female genitalia. Epigyne (Figs 4D, E; 13H, I) very developed and protruding in the ventral side of epigastrum, almost perpendicular to the surface of opisthosoma. Basal zone elongate, formed by the anterior and lateral walls of the bursa copulatrix, opening backward; these walls with a long and scattered hairiness. Posterior wall developing the median plate, semicircular and sclerotized, on which the distal part of the scape rests. Lateral walls not forming "wings", although they distally expand in two triangular points closing the flanks of the bursa copulatrix. Scape relatively large; starting from the anterior wall of the bursa, so that lateral walls with a notch, rounded on each side, delimit the beginning. No specific development of the proscape, because it is limited to change direction, leaning towards the opisthosoma wall (only in a posterior observation appearing a little more dilated and rounded). Middle part of the scape oriented towards the inside of the bursa copulatrix, but maintaining a curvature in "C" so that the distal part is oriented outwards; scape route easily observable laterally. Entire escape greatly sclerotised and looking as a rigid structure, but allowing to observe by transparency the route of the fertilization ducts. diAgnosis. -In its posterior middle part, the paracymbium presents a very developed elongation, acute and curved towards the external side; its apical branch is narrow, straight and lanceolate.
In the radix a sharp, spiny tip develops at its posterior end. The lamella characteristica consists of a long and narrow projection, which gradually sharpens towards the tip, maintaining a progressive curvature. The terminal apophysis, equally long and narrow, describes a curved path, parallel to the lamella characteristica, and somewhat longer. The epigyne is wide at its base and perpendicular to the surface; it narrows progressively distally where a proscape with an elliptical contour, flanked by the lateral walls, stands out. The set of the scape describes a structure in "S" (easily seen in side view). The morphology of the epigyne resembles that of L. ensiferus Barrientos n. sp.; however, the structure of the male genitalia separates the two species, which places L. fadriquei Barrientos n. sp. in an uncertain position in the group of Micronetinae.
eTyMoLogy. -We dedicate this species to Floren Fadrique, the main promoter of the latest biospeleological surveys in Morocco; without his generosity, initiative and dedication on the ground (before, during and after each expedition) we would not have many of the interesting samples of this small study.
disTribuTion. -L. fadriquei Barrientos n. sp. is known only from the type locality, a cavity of the province of Meknès, in the high Moroccan Atlas. Palp and copulatory bulb (Figs 5A-C; 13G). In the paracymbium, posterior middle part particularly well developed; prolonged by a long sharp projection and curved outwards (Fig. 5A, B); proximal part wide and trapezoidal, with some hairs in its basal area; middle part marked by two protrusions or dilatations: one just mentioned in the posterior part and one in the anterior part, thick and blunt, delimiting the anterior pocket; from the anterior pocket, the apical branch narrow, straight, somewhat spatulate, and relatively short. Ventrally bulb showing on its prolateral side the basal structure of the embolic division (Fig. 5C), the radix. radix with a sharp and spiny process in its posterior part, and immediately afterwards, on its internal side, the lamella characteristica. Lamella consisting of a long and narrow projection, progressively sharpened toward the tip, maintaining a progressive curvature. Terminal apophysis parallel to the lamella characteristica, equally long and narrow, and describing a curved path of somewhat greater length; these two similar structures running parallel. Embolus complex, arising from the anterior part of the radix; with a laminar expansion, thin and transparent, as protective element of the embolus proper. Suprategular apophysis as a sharp process in front and below the terminal apophysis and lamelle characteristica; sharp, sclerotized, thicker and curved (Fig. 5A).  Female genitalia. Epigyne (Figs 5D, E; 13E, F). Epigyne very developed and outstanding, in relation to the size of the opisthosoma; almost perpendicular to epigastrum surface. Its basal part narrowing progressively and formed by the anterior wall and the lateral walls, covered with long and scattered hairs. Distally, scape voluminous; proscape drawing a smooth surface with an elliptical contour; proscape framed between the extensions of the lateral walls.
In the margins of the proscape, the fertilization ducts well visible. Scape: middle part located practically below the proscape; distal part oriented outward again, forming a narrow point (stretcher) protruding below. Entire set describing in a small space a double directional change in the shape of an "S", which can be seen in a lateral view (Fig. 5E). By transparency through the anterior wall of the epigyne, the receptacula observed in the pedunculated area, near the scape base. diAgnosis. -The epigyne is well developed on the ventral side of the epigastrum. Its basal part lengthens and narrows distally, while curving backwards (forming a right angle). The side walls form a pair of short "wings" that protect the scape. The proscape is narrow and rounded, somewhat domed; the distal part of the scape forms a projection (stretcher) protruding below the proscape. The structure of the epigyne resembles that of L. taza, a species with which it could have some relationship. However, the curvature of the entire structure makes L. imazigheni Barrientos n. sp. a different species.
eTyMoLogy. -Imazighen is the name that designates the first settlers of North Africa. Throughout their history these peoples have  Female genitalia. Epigyne (Figs 6B, C; 14A, B) very developed and protruding in the ventral side of epigastrum. Its basal part elongate and slightly narrowed distally, while curving backwards (forming a right angle); opening of the bursa copulatrix thus displaced distally. Pilosity of epigyne walls short but clearly distinct; hairs long and well areolated at their base. Posterior wall with a median plate, little visible by the arrangement of the structure itself. Lateral walls elongate forming a pair of short "wings" protecting the scape. Scape originating from the anterior part of the bursa copulatrix, from which it is separated by a furrow. Proscape narrow with a rounded contour, somewhat domed on its external face; flexing forward, hidden by the wings of the lateral walls (middle part of the scape) and backwards (distal part), narrowing and lengthening by a fingering (stretcher) that protrudes below the proscape.

Male
Unknown. diAgnosis. -The paracymbium is especially wide in its proximal part; at the junction with the median part it extends through a small spiny projection; the apical part is digitiform and curved in inverted "C" . The bulb has a lamella characteristica flattened, curved 90° distally, where it is finely serrated and fluted. The terminal apophysis and the basal lobe of the embolus are in the form of submembranous and transparent sheet, with a denticulate border. The epigyne is small and simplified. The proscape is flattened and rounded, wider than long, and with a posterior notch in the folding area. The middle and distal parts of the scape cannot be observed, as they are practically hidden under the proscape; only the distal part of the stretcher is visible. In general, the genital structures are simplified, but sufficiently different to support the definition of a new species of the afer group, particularly diversified in the Maghreb. By the shape of its proscape, the new species seems close to L. longihamatus and perhaps also to L. longipedis.

Lepthyphantes lamellatus
eTyMoLogy. -The chosen name highlights the structure of the lamella characteristica; it has a flattened and laminar appearance, with its distal edge finely serrated. It is easily observable in the bulb at rest.
disTribuTion. -This species is exclusively known from the type locality, a cave in the province of Azilal, in the high Atlas of Morocco.  (Figs 7A-c; 14F). Tibia and tarsus (cymbium) without notable elements, although the latter presents laterally the paracymbium and lodges a voluminous bulb in its ventral face. Paracymbium wide and trapezoidal in its proximal zone; in the junction with the middle part an acute spinous protrusion delimiting the posterior pocket. Apical part considerably narrowed, having distally the shape of a curved and blunt fingering; the whole set curved in a regular "C". At rest, bulb with on its retrolateral side, a lamella characteristica wide and laminar, D-shaped, toothed and finely fluted distally distally; this structure characteristic and easily observed. From the radix a submembranous terminal apophysis and the embolus; embolus apical lobe with a transparent membrane with with a denticulate border. Main lobe of embolus protected by this membranous lobe; more sclerotised and terminated by a sharp tooth. In the anterior part of the resting bulb, distal part of suprategular apophysis visible, having the shape of an acute and sclerotised projection. diAgnosis. -The epigyne is small and perpendicular to the surface of the epigastrum; it has a wide bursa copulatrix in its basal part, closed behind by a subquadrangular median plate. The scape shows a rigid proscape with a rounded and flat contour (the semicircular path of the fertilization ducts can be reached at its edges); the middle and distal parts of the scape are reduced; the latter presents small, distal projection that protrudes slightly below the proscape. Following the characterization made by Bosmans (2006), this species is very close to L. afer (Simon, 1913); it differs from this species by presenting a practically round proscape (instead of oval) and being completely blind.   Lateral walls and posterior wall well developed, with a medial plate. Scape originating from the anterior part of the bursa copulatrix, not separated from the bursa copulatrix by a furrow; proscape rigid with a rounded and flat contour (its tegument well sclerotized but showing the fertilization ducts, which follow a semicircular route next to the edges of the proscape); median part of the scape quite small, so that it is immediately linked to the distal part; distinctly more narrow than the proscape and quite sclerotised; stretcher with a small blunt finger-shaped projection, without lateral wings, at the base of which begin the fertilization ducts.

Male
Unknown.  diAgnosis. -In his description, Bosmans (1985) indicates that it is a species very close to Lepthyphantes afer (Simon, 1913), Lepthyphantes venereus Simon, 1913 and Lepthyphantes emarginatus Fage, 1931, but it is distinguished from them by its copulatory organs, especially the spine of the patella, the paracymbium with a single posterior tooth, a small lamella characteristica, the clearly trimmed proscape, the very long stretcher and the deep cleft of the median plate.
disTribuTion. -The holotype comes from the cave Akiam Ins Er rebi, Imilchil (Bosmans 1985); to this locality is added the one indicated by Tanasevitch (2014), Ifri el Caid, Ait Mohammed. The first belongs to the province of ouarzazate and the second to Azilal; both are located in the high Atlas of Morocco. our data confirm its presence in the type locality and add a new one, the Ifri Bouyzem cave, in Azilal.

Lepthyphantes maurusius Brignoli, 1978
MATeriAL exAMined. diAgnosis. -According to Brignoli (1978) the epigyne is characteristic, although typical of the species close to the "pinicola group". It will be necessary to confirm this relationship. There is obviously a group of taxa comprising "Spanish-Moroccan forms", such as L. aelleni, L. taza, L. imazigheni Barrientos n. sp., L. phallifer Fage, 1931, L. fagei Machado, 1939 and even L. lorifer Simon, 1907, which could be related. Among them L. maurusius can be easily distinguished by its epigyne and the opening of the bursa copulatrix, which is oriented outwards, instead of backwards or towards the posterior walls.
disTribuTion. -L. maurusius is only known from the type locality (Brignoli 1978), the cave of Sidi Mejbeur, in the Taza region (high Atlas of Morocco). The data of Tanasevitch (2014) and those given in the present work confirm its presence in the above-mentioned cave. diAgnosis. -The paracymbium is domed in its middle part, with a pair of short denticles on its lower edge; the apical part is narrow, with a sinuous and finger-shaped apex. The lamella characteristica is bifurcated basally: inner branch narrow, laminar and ending in tip; outer branch also laminar, widening progressively and with a series of sclerotized tips in its distal edge. The anterior wall of the epigyne is strongly curved backwards; the lateral walls lengthen on the sides forming a pair of subtriangular "wings", which protect the proscape. The proscape expands after a brief basal narrowing, making a well sclerotized subcircular structure. Because of its female genital traits, L. sasi Barrientos n. sp. recalls L. djazairi Bosmans, 1985, always within the characteristics of the "afer group". The structures shown by the male copulatory bulb are more complex to relate; the basal fusion of the lamella characteristica and terminal apophysis constitutes a characteristic feature; but as a whole it ressembles those presented by L. aelleni.
eTyMoLogy. -This species is dedicated to the speleologist Francisco Sas Planas, recently deceased, one of the main promoters of the Moroccan expeditions in its beginnings (years 1968-1970); this little tribute and our gratitude for him.
disTribuTion. -The samples studied come from two nearby caves located in Merhraoua, in the high Atlas of Morocco (Taza region). Palp and copulatory bulb (Figs 9A-C; 14I). Paracymbium thick and clearly domed in its middle zone (Fig. 9A), whose lower edge is delimited by a pair of short denticles; proximal part wide; upper end rounded, with a group of short hairs at its base; middle part domed, making the two pockets, one posterior and one anterior (this specially developed); apical part of paracymbium narrowing progressively, so that its end is sinuous and fingershaped. Bulb as on Fig. 9A, B: with the lamella characteristica attached at its base to the terminal apothysis. Terminal apophysis straight, narrow, laminar, and distally pointed; outer branch (lamella characteristica, located next to the paracymbium) also laminar but widening and progressively curved accompanying the surface of the bulb. Distal part of the lamella characteristica of subtriangular contour, presenting a series of sclerotized tips related to a clear striation at least in its distal half; apex darker with denticulate margins. radix (Fig. 9C) distally differentiating a spiny and acute prolongation. Embolus complex protruding from the midle zone of the latter prolongation. Main part of embolus simple, narrow at base and widened at its distal area that is laminar, except at the black point where the duct is opened; accompanied by a laminar structure, narrow at base, which also dilates at its apex making a translucent sheet, protecting the distal part of the main part of the embolus. Fickert's gland clearly visible. Suprategular apophysis sclerotized and with several acute tips, protruding in the anterior part of the bulb, under the complex of the embolic division. , H) standing out on the ventral side of the epigastrum; almost perpendicular to the surface of the opisthosoma, with the opening of the bursa copulatrix looking back. Basal part bulging and elongate in front and on the sides. Anterior wall, with a long and scattered hairiness, curved back giving a rounded profile. Lateral walls also elongate forming on each side a pair of subtriangular "wings", protecting the flanks of the proscape; in the basal zone, receptacula seen by transparency. Posterior wall short and with a narrow median plate. Proscape originating from the anterior wall of the bursa copulatrix, between the two expansions of the lateral walls. Near its base, proscape widening after a brief basal narrowing, giving a well-sclerotized subcircular contour structure. Middle part of the scape curved and placed under the proscape, hidden and facing forward; slightly sclerotized and curved again to give the distal part, very short, which also does not protrude from the proscape; strecher observed laterally below it. Fertilization ducts observed by transparency. diAgnosis. -The paracymbium is narrow; it presents an acute denticle in the proximal part of its middle zone, and a laminated prolongation with three points in its anterior part; the apical part is finger-shaped and bent distally. In the bulb, the lamella characteristica is fused at the base with the terminal apophysis, making a bifurcated structure: the terminal apophysis is short, narrow and sinuous; the lamella characteristica is narrow and extends forward on the outer side of the bulb, apically denticulated at on the margins. The female is characterized by the peculiar conformation of the epigyne: the walls of the bursa copulatrix protrude, moving its opening away from the epigastric wall. The lateral walls differentiate an elongated process ("wings") on each side, flanking a reduced proscape, as well as the rest of the scape, which is not very apparent. Lepthyphantes taza could be related to a set of species from the Maghreb, L. aelleni, L. maurusius, L. imazigheni Barrientos n. sp. and L. sasi Barrientos n. sp. (although in the latter there are features in the female genitalia that bring it closer to the "afer group").
disTribuTion. -This species was only known from the type locality (Tanasevitch 2014 Palp and copulatory bulb ( Fig. 10A-C). Paracymbium narrow; proximal part wide, with rounded edges and small hairs; middle part with an acute denticle at the posterior end (which closes the posterior pocket) and a laminated three-pronged denticle at the anterior end (which in turn delimit the posterior pocket); apical part finger-shaped, bent distally. With the bulb at rest: radix well visible ventrally on prolateral side. Lamella characteristica fusing at the base with the terminal apophysis, protruding from the back, so that it appears as a bifurcated structure. Anterior branch short, narrow and sinuous; posterior branch laminar, narrow, longer and extending forward on the outer side of the bulb; distal part darker, with denticulate margins. Embolus emerging from the middle part of the radix; quite complex: main part simple, laminar, widened in its middle part and finished in a black bifurcated tip; accompanied, from the base, by a sheet becoming translucent and protecting distally the main part. Fickert's gland visible in the radix. Suprategular apophysis sclerotised, pointed and curved, protruding below the embolus and facing forward. diAgnosis. -The proximal part of the paracymbium is narrow; the middle part is wider and has an anterior denticle, long and sharp; the anterior part is wide and has rounded contours. In the bulb, the lamella characteristica is elongated and narrow, ending in a sharp point; in a subterminal position, it presents a flat and laminar denticle terminated in two points, like a banderole. The epigyne protrudes on the ventral side of the epigastrum; the posterior wall bears a middle plate, with two juxtaposed rounded areas; the lateral walls protect the scape. The scape is narrow, elongated and almost completely hidden; the proscape, barely individualized, is oriented towards the back, and can be clearly observed in side view; middle and distal parts of the scape describing an inflection in "S" near the median plate. Although the shape of the lamella characteristica individualizes P. banderolatus Barrientos n. sp., its structures relate it to other species of the genus, originated from caves of the Iberian Peninsula, such as P. cortesi ribera & De Mas, 2003, P. lorifer (Simon, 1907 or P. cadiziensis (Wunderlich, 1980), among others.
eTyMoLogy. -The name refers to the shape of the lamella characteristica, narrow and elongated at its base, and dilated in an expansion that recalls a "banderole", in its distal part.
disTribuTion. -The localities in which this new species has been captured are located in Ait Abdallàh, province of Taroudant, Antiatlas, in southern Morocco. Palp and copulatory bulb (Figs 11A-C; 15C). Proximal part of paracymbium narrow and with parallel sides, with some short hairs; middle part noticeably wider and flattened, with a long, sharp denticle in the anterior area, and only one more pronounced ridge on its posterior margin; so that the "pockets" are poorly marked; anterior part also wide, with rounded contours, and extended backwards by a laminar curved structure. Bulb as on Figure 11A, B: lamella characteristica clearly visible elongated and narrow, ending in a sharp, slightly curved tip; in a subterminal position, with a flat and laminar denticle, finished in two points, like a banderole. radix revealing the Fickert's gland and distally developing a small sharp tip (only visible with the expanding bulb). Terminal apophysis emerging from radix middle part, sublaminar and rounded. Embolus in more distal position, with two sclerotized tips and protected by a laminar protrusion. Below the radix, forward and clearly visible at the anterior end, the suprategular apophysis, curved and pointed, protruding. Female genitalia. Epigyne (Figs 11D-F; 15A, B) standing out on the ventral side of the epigastrum, leaning backwards on the surface of the postepigastrum. This arrangement resulting from the lengthening of the anterior wall of the bursa copulatrix (which thereby opens backwards). Anterior wall with a long and scattered hairiness, easy to see, as well as by transparency the two receptacula in the basal area.
Posterior wall with the medial plate, consisting of two juxtaposed rounded areas. Lateral walls not forming "wings", but protecting the scape and with an abundant pilosity, like the anterior wall. Scape narrow, elongated and almost completely hidden, since inflected inmediately and oriented backwards. In this way, proscape slightly differentiated and oriented backwards, narrowing and elongated, being easily observed laterally. Middle part of the scape and distal part ZOOSYSTEMA • 2020 • 42 (1) small, forming an inflection in "S" that is close to the median plate of the posterior wall.
Palliduphantes cadiziensis (Wunderlich, 1980)  disTribuTion. -Although the type locality (Algeciras, Cádiz) is located in the south of Spain (Wunderlich 1980), this species has been mentioned later in several localities of Morocco (Bosmans 2006), including in the region of Taza, in the two cavities that we have previously indicated.   of the scape widening and curved again, describing a "C. The whole structure forming a rigid and narrow structure revealing the fertilization ducts by transparency. The two receptacles also observed, at the base of the structure.

Male
Unknown.
Genus diAgnosis. -The medial part of the paracymbium has a pair of separated teeth of similar length. The lamella characteristica of the male bulb has a short and wide distal denticle. In the epigyne, the visible part of the scape has the shape of an anchor in its posterior end, in the middle separating two lateral semicircular projections. Frequently, the opisthosoma presents black spots in the form of circumflex accents, especially marked in its posterior half.

DISCUSSIoN
The data provided here represent direct and not methodical catches. These catches, when analysing species by species, are relatively scarce. However, as a whole the data provided here are a unique addition to Micronetinae literature and in some cases they confirm some species previously recorded from Moroccan caves. The number of novelties captured by BIoSP is surprising, but it is also clear that among the taxa there seem to be some morphological relationships that are strengthened when positioned in their geographical context. First, we must highlight the presence of ten taxonomic novelties in a set of seventeen different species that fit into the genus Lepthyphantes "sensu lato" and Palliduphantes. This situation is partly explained by the fact that BIoSP has visited a large number of caves over an extended period of time, many of them previously unsurveyed. It is also necessary to note there is both considerable and unexamined diversity in the Atlas Mountains. Thus, the descriptions in the present work of ten new species are potentially only a fraction of an even greater possible diversity.
It is premature to speculate on how, when, and from which species the diversification of these indisputably related forms took place. But perhaps it is now worth risking a discussion of possible relationships in an attempt to clarify the Lepthyphantes "sensu lato" phylogeny.
The so-cALLed "afer group" Fage (1931;1945) highlighted the existence of considerable morphological affinity between some species of the Mediterranean basin, without defining or delimiting any group. Later, Brignoli (1971) dedicated an extensive discussion to this question by speaking clearly of an "afer group" by placing at the head of Lepthyphantes afer Simon, 1913. Brignoli insisted that the most obvious character of relationship in this group is the morphology of the epigyne in females and the paracymbium and lamella characteristica in males. Bosmans (1985;2006) described several species that he placed without discussion in Brignoli's "afer group". In a similar manner Hormiga & ribera (1990) took advantage of the description of Lepthyphantes bidentatus Hormiga & ribera, 1990 to compile the Mediterranean species that can be included in this group. Identical information is found in the work of Saaristo & Tanasevitch (1993). We do not find any discussion of the "afer group" in this 1993 work, just an implicit acceptance and the list of species included within the group; an attitude that Tanasevitch (2014) repeated when studying samples deposited in the Geneva Museum. Now, when studying the same taxa collected in Morocco, we find previously known species (L. brevihamatus, L. longihamatus) as well as new ones (L. sasi Barrientos n. sp., L. biospeleologorum Barrientos n. sp., L. lamellatus Barrientos n. sp. and L. leknizii Barrientos n. sp.) which are clearly related at least by the structure of the epigyne. In all of these taxa it is possible to recognize the basic morphological features enunciated by Brignoli. However, in the species for which we have adult males (L. sasi Barrientos n. sp. and L. lamellatus Barrientos n. sp.) the morphology of the paracymbium and lamella characteristica differs from the pattern enunciated for the group. It is therefore extremely complex to ensure a close relationship between all these species, requiring a more detailed analysis of all these structures.

piniphantes?
Many species remain catalogued (World Spider Catalog 2018) in the genus Lepthyphantes (sensu Menge, 1866). Among them are several spiders captured in caves and hollows in the mountainous areas of Morocco and Algeria (Denis & Dresco 1957;Brignoli 1978;Tanasevitch 2014). Some authors (Bosmans 1985) have attempted to bring them within some morphological proximity to Piniphantes pinicola (Simon, 1884) so they would be candidates for inclusion in the genus Piniphantes Saaristo & Tanasevitch, 1996. However, these confounding groups of Moroccan and Algerian spiders also have common characteristics, suggesting a close relationship (Bosmans 2006;Tanasevitch 2014). These comments apply mainly to L. aelleni and other Atlas species such as L. maurusius and L. taza. The males of these two former species were not known (the one of the first remains unpublished). However, the morphology of the epigyne displays a few traits that can be valued as key elements of a possible relationship, such as the length of the basal structure, the elongation of its walls (anterior, lateral and posterior) which carry the bursa copulatrix, the location of the scape to a distal position, and finally the considerable Among the material that we have analysed and consider novel is a male of L. taza and females of L. almoravidus Barrientos n. sp. and L. imazigheni Barrientos n. sp. (albeit unfortunately unaccompanied by corresponding males). The comparative morphology of L. aelleni and L. taza males give support to a close relationship in both the structure of the paracymbium, In our opinion L. aelleni, L. maurusius, L. taza, L. almoravidus Barrientos n. sp. and L. imazigheni Barrientos n. sp. are possibly related due to the structure of their male and female genitalia. Therefore, these species could be integrated into a natural group, incorporated geographically in the high Mo-roccan Atlas. Lepthyphantes ajoti Bosmans, 1991 andL. exvaginatus Deeleman, 1984, perhaps could be added to this group (Bosmans 2006) although we were unable to examine specimens of these two species.
paLLiduphantes in norTh AfricA Saaristo & Tanasevitch (2001) included 47 species in the new genus Palliduphantes (in addition to other 16 that are synonymized). of these, nine are found in the Iberian fauna,  (Wunderlich, 1980). Subsequently several other species have been described: three for the Iberian fauna (ribera et al. 2003), two for the Algerian fauna (Bosmans 2006) and one for Tunisian fauna (Bosmans 2003). Most of the 58 species currently considered in the genus Palliduphantes (World Spider Catalog 2018) are found in the western Palearctic zone and primarily in Mediterranean countries. Therefore, considering the size, characteristics and position of Morocco, the description of only two species of Palliduphantes is probably an underrepresentation. The morphology of the paracymbium and the lamella characteristica of the male of P. banderolatus Barrientos n. sp. conform well to the diagnostic characters of Palliduphantes (Saaristo & Tanasevitch, 2001). on the contrary, the female has an elongated epigyne that is loosely consistent with the one described by Wunderlich (1987) for P. longiscapus (Wunderlich, 1987) of Gran Canaria, or the one presented by P. longiseta (Simon, 1884) of Corsica (Muller & Heimer 1991;Bosmans & Colombo 2015). In any case, the consistency and layout of the scape (together with the characters of the male) make P. banderolatus Barrientos n. sp. a clearly independent species. Such a greater length of the female genital structures is also seen in a second species from the south of Morocco, Palliduphantes megascapus Barrientos n. sp., of which the male remains unknown. Therefore the assignment of P. megascapus Barrientos n. sp. to Palliduphantes is still provisional. However, taking into account the morphology of its epigyne, the relationship between P. banderolatus Barrientos n. sp. and P. megascapus Barrientos n. sp. seems to be beyond doubt.
uncerTAin posiTion of L. fadriquei bArrienTos n. sp. proxiMiTy beTween L. ensiferus bArrienTos n. sp. And L. fagei MAchAdo, 1939? Although the above groupings of possible morphological affinities are debatable, the comments made could be useful in later considerations. However, two of our ten new species (L. ensiferus Barrientos n. sp. and L. fadriquei Barrientos n. sp.) are difficult to place. When considering the conformation of female genitalia, Lepthyphantes ensiferus Barrientos n. sp. seems to have an especially close morphological relationship with an Iberian species, Lepthyphantes fagei, although there is a significant difference in the male structures of both species. It is possible, albeit less probable, that L. fadriquei Barrientos n. sp. could be grouped with these two forms. In this scenario however the male genitalia of L. fadriquei Barrientos n. sp. seems to be even more dissimilar from the features presented by L. ensiferus Barrientos n. sp. and L. fagei. It is therefore appropriate to include L. ensiferus Barrientos n. sp. and L. fadriquei Barrientos n. sp. within the vaguer grouping of Lepthyphantes "sensu lato".