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Pérez-González, Sergio, Zaballos, Juan P., Ghannem, Samir (2013): Intraspecific variation in Typhlocharis Dieck, 1869 (Coleoptera, Carabidae, Anillini): the case of two new species of the baetica group. Zootaxa 3710 (1), DOI: http://dx.doi.org/10.11646/zootaxa.3710.1.3

Typhlocharis mixta n. sp. Figs 4, 5 Type series. Holotype, ♂, SPAIN, Badajoz, Puerto de los Reventones (11 km NE Fregenal de la Sierra), 22 - 11- 2012, 500 m (38 �� 15 ��� N, 06�� 34 ��� W ), J. P. Zaballos, S. P��rez & S. Ghannem leg. (Coll. J.P. Zaballos, UCM). Paratypes: 659 ♂♂, 616 ♀♀ same data as holotype; 1 ♂ same locality as holotype, 24 -04- 2012, J. P. Zaballos & S. P��rez leg.; 10 ♂♂, 11 ♀♀ SPAIN, Badajoz, Valverde de Burguillos (4.4 km NE), 22 - 11-2012, 488m (38 �� 21 ��� N, 06�� 31 ��� W), J. P. Zaballos, S. P��rez & S. Ghannem leg. (Coll. J.P. Zaballos and Coll. S. P��rez-Gonz��lez, UCM). 4 ♂♂, 4 ♀♀ same data as holotype (MNCN, Madrid, Catalogue number: 11612). Diagnosis. Small endogean anophthalmous beetle, with narrow and subrectangular body covered by microreticulate integument and scattered pubescence. Vertex with pars stridens. Clypeus with a distinctive median tooth. Pronotum with a wide medial hiatus. Elytra faintly serrated in lateral margins and apical margin with three to six pairs of small denticles, highly irregular. Umbilicate series with six setae (4 + 2). Abdomen with ventral foveae only in females. Pseudotetramerous tarsi in all the legs. Aedeagus as in Fig. 5, ���plough-shaped��� endophallic sclerite and broad parameres. Female genitalia as in Fig. 5, with subcylindrical spermatheca. Description. Length 1.21���1.49 mm (males), 1.27���1.60 mm (females) (Fig. 4). Head: wider (0.24���0.34 mm) than long (0.22���0.30 mm), labrum subrectangular to slightly notched, rounded edges and triangular sclerotized area with middle button. Clypeus with a median tooth in the anterior margin. Right mandible with a prominent, pointy, terebral tooth on inner margin. Left mandible without teeth, but inner margin with sharp edge. Ligula with very prominent middle lobe and long paraglossae. Wide gula, approximately twice as long as wide, the lateral sutures may appear fainter. Rest of cephalic characters and cephalic chaetotaxy, including antennae, as described for T. acutangula n. sp. Pronotum: subrectangular, longer (0.32���0.45 mm) than wide (0.3���0.4 mm), slightly narrowed in posterior region. Proportions of pronotum present a wide range of variation (Fig. 6). Wide medial hiatus (moderately wider than three adjacent intersetae spaces). Lateral margins with denticles near posterior angles highly variable in shape and number, from two or three, irregular and faint to four or five, well marked and serrated. The posterior angles are also variable from pointy and protruding to blunt and low (Fig. 6). Other characters and chaetotaxy are the same as for T. acutangula n. sp., except a pattern of three pairs of setae, parallel to posterior margin [s-l-l / l-l-s], the row of small, thin setae of anterior and posterior margins (filiform) and the short discal pubescence, arranged in four pairs of longitudinal rows. Prosternum as described for T. acutangula n. sp. Elytra: approximately two times longer (0.64���0.85 mm) than wide (0.32���0.41 mm), subparallel. Lateral margins with 18���21 subtriangular denticles, faint and progressively smoother towards posterior, but still marked in last third. Apical margin smoothly rounded, with three to six pairs of small denticles (showing a notable range of individual variation, but mostly combinations of four and five pairs) (Fig. 7 A, B). Transverse scutellar organ with straight to slightly subtriangular margin and pair of ���buttonholes��� present near base of elytra. Chaetotaxy: umbilicate series formed by anterior group with four setae and posterior group with two setae (Fig. 4). One pair of scutellar setae. No apparent discal setae, but the population of Puerto de los Reventones is prone to have one pair of anterior ���pseudodiscals��� (i.e. pubescent setae longer than the surrounding pubescence). Rest of characters as in T. acutangula n. sp. Legs: similar in both sexes. Intermetacoxal space not widened. Metacoxae with smoothly rounded ���flap���. Rounded metatrochanters, metatibiae with moderately dilated distal region and a short perpendicular seta in the inner side. Inner side of pro- and mesofemora covered with small protuberances, absent in metafemora. Pseudotetramerous tarsi in all the legs, tarsomeres 4 �� and 5 �� retain a faint articular membrane, but are partially fused and not capable of a normal articulation. Abdomen: as described for T. acutangula n. sp., but with deep ventral foveae in the first and second ventrite of females, absent in males. Last ventrite with a sexually dimorphic pattern of five to six pairs of setae in posterior margin: l-s-s-l-s-m / m-(s)-l-s-s-l (Fig. 4). Aedeagus: sickle-shaped median lobe (length: 0.22 mm) and ���bottleneck-shaped���, smoothly rounded apex, slightly bent to right (anatomically oriented) in dorsal view (Fig. 5 A,B). Endophallus with ���plough-shaped��� sclerites, as drawn in Fig. 5 B. Short and broad parameres, both with two medium-sized apical setae (Fig. 5 B). Female genitalia adjusts to the description of Vigna-Taglianti (1972). Long tubular gonocoxites, with double apical setae. Lateral setae absent, but lateral pores in the gonocoxites. Short gonoduct with two regions, thinner proximally (diameter 0.003 mm) and thicker distally (diameter 0.008 mm). Spermatheca subcylindrical and short, ���barrel-shaped��� (length: 0.011 mm). Conical spermathecal gland (length: 0.014 mm), sclerotized in distal region (Fig. 5 C). Etymology. The name of the species plays a double meaning, referring to the high intraspecific variation that gives a mixture of body shapes and sizes to the populations. On the other hand, it refers to the presence of certain morphologic traits (such as the tooth in clypeus, the trend to diffuse gula or the pseudotetramery) unexpected for members of baetica group, but known in other species groups, like mixed pieces in different jigsaw puzzles. Habitat. Typhlocharis mixta n. sp. is found in two localities separated 13 km (Fig. 1): 1) Puerto de los Reventones, in open prairie environment with low hills and valleys crossed by seasonal streams. Vegetation mainly composed by low herbaceous plants, grasses and thistles, with scattered broom bushes (Retam Raf.). Higher grasses and rushes (Juncus L.) appears near the stream. Dark, brownish soil, with scarce large boulders of gneisses and shales (Proterozoic, Valverde Formation). This location was sampled four times (March, April, June and November), with the best humidity range in the last sampling. 2) Valverde de Burguillos, in a small meadow surrounded by cultures and open holm oak forest. Vegetation is akin to the former locality, with the exception of holm oaks. Brown to reddish soil, rich in clays, with embedded small to medium sized granite boulders (Proterozoic). Variability. There are not significant differences between the populations of Puerto de los Reventones (1285 specimens) and Valverde de Burguillos (21 specimens), both have the same range of intraspecific variation. The species is morphologically very diverse in proportions and size, from 1.2 to 1.6 mm in length, but the presence of a tooth in clypeus, umbilicate series 4 + 2, the perpendicular setae on distal end of metatibiae, pseudotetramerous tarsi, ventral foveae in females, genitalia and the wide hiatus are virtually constant within the population. The terebral tooth is usually very prominent but in some specimens is blunt, low or almost absent. The tooth of clypeus ranges from blunt to very pointy (but always present). The lateral sutures of the gula are very faint in about 20 % of the series, resembling the ���difusse gula��� morphology observed in quadridentata group (P��rez-Gonz��lez & Zaballos 2013 c). There is slight variation in the pronouncement of epilobes. Two specimens have a widened pars stridens, two times wider than the average. Pronotum and elytra have the higher rates of variability. Pronotum proportions can be certainly different, with three main morphs: subrectangular, smoothly curved lateral margins and 3-4 well marked posterolateral denticles (Fig 6: 1,2), the most common; rectangular, straight lateral margins and 4-5 (even 6 in one specimen) well marked denticles (Fig 6: 3,4) and subsquare, with 2-3 blunt and irregular denticles (Fig 6: 5,6). Asymmetric pronotum are abundant, and they affect the number of denticles or the pronouncement of the posterior angle (Fig 6: 7). Five specimens have aberrant pronotums with widened, notched or asymmetric lateral margins (Fig 6: 8���10). Transverse scutellar organ varies from straight to subtriangular and the margins can be irregular or smooth. Lateral denticles of elytra can be more or less marked. Apical denticles of elytra appear in a wide range of combinations and shapes (Fig. 7): the most common are 4 / 4 (17 % of the series), 4 / 5 (15 %), 5 / 4 (15 %), and 5 / 5 (22 %), but they range from 2 / 3 to 7 / 7. All the observed combinations and their abundance within the population are expressed in Fig. 7 B. The inner pair, closer to the elytral suture is especially developed and bigger than the other denticles (Fig. 7 A: 3) in some specimens, resembling those of Typhlocharis tertia P��rez-Gonz��lez & Zaballos, 2013. About 14 % of the series (145 specimens) have bifid or double denticles and 7 specimens show triple denticles (Fig. 7 A: 4-6). Asymmetries (Fig. 7 A: 7���9) and damaged elytra are also common. Variations in patterns of chaetotaxy occur in labium, prebasilar, last ventrite and sensilla coeloconica of the 11 th antennomere. It is remarkable that the ventral sensilla coeloconica in 5 �� and 6 �� antennomeres also varies: 7 specimens have only sensilla coeloconica in the 5 �� antennomere and 3 specimens have them in antennomeres 5 ��, 6 �� and 7 ��. It is the first evidence of variation in this feature within the same species, and the presence of sensilla coeloconica in 5 �� antennomere but not in 6 �� has been reported only in Typhlocharis tetramera P��rez-Gonz��lez & Zaballos, 2013. Finally, there are several specimens with teratologies, like antennomere fusion (5 specimens, with fused 3 ��- 4 ��, 5 ��- 6 ��, 9 ��- 10 �� and 10 ��- 11 �� antennomeres respectively), or malformed mesotibiae (1 specimen, associated with a malformed pronotum, possible result of a failed moult). Affinities. Typhlocharis mixta n. sp. presents the diagnostic characters of baetica group, defined by the presence of several pairs of apical denticles. Affinities with other species of the group are the umbilicate series of 4 + 2, shared with T. pacensis Zaballos & Jeanne, 1987; T. furnayulensis Zaballos & Banda, 2001; T. secunda Perez- Gonz��lez & Zaballos, 2013 and T. acutangula n. sp. The protuberances in femora also appears in T. baetica Ehlers, 1883; T. tertia Perez-Gonz��lez & Zaballos, 2013; T. quarta Perez-Gonz��lez & Zaballos, 2013 and T. acutangula n. sp.; the high number and serrated aspect of posterolateral denticles of pronotum are also typical of T. baetica and T. quarta, but many specimens have pronotum with few and less defined denticles akin to that of T. tertia and the other species of baetica group. The broad parameres are unusual within the group and strongly resembles those of T. tertia. T. secunda Perez-Gonz��lez & Zaballos, 2013 is very similar: both species shares the same umbilicate series, size range, overall shape of pronotum posterolateral denticles and presence (not always in T. mixta) of anterior ���pseudodiscal��� setae. However, the new species is well differentiated by its diagnostic features. The tooth of clypeus, subcylindrical spermatheca, metatibial perpendicular short seta and the pseudotetramerous tarsi found in T. mixta n. sp. are unique of this species within baetica group, but appear in other Typhlocharis species. The tooth of clypeus is the definitory trait of the six species of gomezi group (Zaballos & Farin��s 1995; Zaballos & Ruiz-Tapiador 1997) which also have a short perpendicular seta in metatibiae. The trend of diffuse gula in some specimens, the variability of ventral sensilla coeloconica in antennomeres 5 �� and 6 �� and the pseudotetramerous tarsi are features previously observed only in species of quadridentata group (P��rez-Gonz��lez & Zaballos 2013 c). In both cases the possible homology of these characters remains unclear. Key to species of baetica group (update of P��rez - Gonz��lez & Zaballos 2013 a) 1. Elytra with umbilicate series 4 + 4...................................... T. quarta P��rez-Gonz��lez & Zaballos, 2013 - Elytra with umbilicate series different to 4 + 4.............................................................. 2 2. Elytra with umbilicate series 4 + 3........................................................................ 3 - Elytra with umbilicate series 4 + 2 or 4 + 1.................................................................. 6 3. Inner side of femora with protuberances, warty surface. Length> 1.4 mm....................................... 4 - Inner side of femora without protuberances, smooth surface. Length T. baetica Ehlers, 1883 - Apical margin of elytra with 3 pairs of large denticles. Length 1.7 mm.......... T. tertia P��rez-Gonz��lez & Zaballos, 2013 5. Elytra with lateral denticles marked in all their length. Very faint or absent ventral foveae. Length 1.1 mm........................................................................................ T. aguirrei Zaballos & Banda, 2001 - Elytra with lateral denticles marked only in the first third of their length. Ventral foveae well developed in both sexes. Length 1.3 mm T. matiasi Zaballos & Banda, 2001 6. Elytra with umbilicate series 4 + 2........................................................................ 7 - Elytra with umbilicate series 4 + 1....................................................................... 11 7. Clypeus with a median tooth. Ventral foveae in females. Length 1.2���1.6 mm........................... T. mixta n. sp. - Clypeus with straight margin, without median tooth........................................................... 8 8. Metatibiae with a pointy dentiform projection in the distal end. Ventral foveae absent. Length 1.1���1.3 mm..................................................................................................... T. acutangula n. sp. - Metatibiae without dentiform projection.................................................................... 9 9. Elytra with 1 pair of discal setae in the anterior third. Length 1.4 mm.......... T. secunda P��rez-Gonz��lez & Zaballos, 2013 - Elytra without discal setae. Length T. pacensis Zaballos & Jeanne, 1987 - Apical margin of elytra with 5���7 pairs of small apical denticles. ���Y-shaped��� endophallus sclerites. Length 1.1 mm............................................................................. T. furnayulensis Zaballos & Banda, 2001 11. Elytra with lateral denticles marked in all their length. Thin, ���Y-shaped��� endophallus sclerites. Subspheric-irregular spermath- eca. Length 1.1 mm T. millenaria Zaballos & Banda, 2001 - Elytra with lateral denticles marked only in the first third of their length. Thick, stick-shaped endophallus. Reniform spermath- eca. Length 1.1 mm................................................. T. prima P��rez-Gonz��lez & Zaballos, 2013, Published as part of P��rez-Gonz��lez, Sergio, Zaballos, Juan P. & Ghannem, Samir, 2013, Intraspecific variation in Typhlocharis Dieck, 1869 (Coleoptera, Carabidae, Anillini): the case of two new species of the baetica group in Zootaxa 3710 (1), DOI: 10.11646/zootaxa.3710.1.3, http://zenodo.org/record/223802

Typhlocharis mixta n. sp. Figs 4, 5 Type series. Holotype, ♂, SPAIN, Badajoz, Puerto de los Reventones (11 km NE Fregenal de la Sierra), 22 - 11- 2012, 500 m (38 º 15 ’ N, 06º 34 ’ W ), J. P. Zaballos, S. Pérez & S. Ghannem leg. (Coll. J.P. Zaballos, UCM). Paratypes: 659 ♂♂, 616 ♀♀ same data as holotype; 1 ♂ same locality as holotype, 24 -04- 2012, J. P. Zaballos & S. Pérez leg.; 10 ♂♂, 11 ♀♀ SPAIN, Badajoz, Valverde de Burguillos (4.4 km NE), 22 - 11-2012, 488m (38 º 21 ’ N, 06º 31 ’ W), J. P. Zaballos, S. Pérez & S. Ghannem leg. (Coll. J.P. Zaballos and Coll. S. Pérez-González, UCM). 4 ♂♂, 4 ♀♀ same data as holotype (MNCN, Madrid, Catalogue number: 11612). Diagnosis. Small endogean anophthalmous beetle, with narrow and subrectangular body covered by microreticulate integument and scattered pubescence. Vertex with pars stridens. Clypeus with a distinctive median tooth. Pronotum with a wide medial hiatus. Elytra faintly serrated in lateral margins and apical margin with three to six pairs of small denticles, highly irregular. Umbilicate series with six setae (4 + 2). Abdomen with ventral foveae only in females. Pseudotetramerous tarsi in all the legs. Aedeagus as in Fig. 5, ‘plough-shaped’ endophallic sclerite and broad parameres. Female genitalia as in Fig. 5, with subcylindrical spermatheca. Description. Length 1.21–1.49 mm (males), 1.27–1.60 mm (females) (Fig. 4). Head: wider (0.24–0.34 mm) than long (0.22–0.30 mm), labrum subrectangular to slightly notched, rounded edges and triangular sclerotized area with middle button. Clypeus with a median tooth in the anterior margin. Right mandible with a prominent, pointy, terebral tooth on inner margin. Left mandible without teeth, but inner margin with sharp edge. Ligula with very prominent middle lobe and long paraglossae. Wide gula, approximately twice as long as wide, the lateral sutures may appear fainter. Rest of cephalic characters and cephalic chaetotaxy, including antennae, as described for T. acutangula n. sp. Pronotum: subrectangular, longer (0.32–0.45 mm) than wide (0.3–0.4 mm), slightly narrowed in posterior region. Proportions of pronotum present a wide range of variation (Fig. 6). Wide medial hiatus (moderately wider than three adjacent intersetae spaces). Lateral margins with denticles near posterior angles highly variable in shape and number, from two or three, irregular and faint to four or five, well marked and serrated. The posterior angles are also variable from pointy and protruding to blunt and low (Fig. 6). Other characters and chaetotaxy are the same as for T. acutangula n. sp., except a pattern of three pairs of setae, parallel to posterior margin [s-l-l / l-l-s], the row of small, thin setae of anterior and posterior margins (filiform) and the short discal pubescence, arranged in four pairs of longitudinal rows. Prosternum as described for T. acutangula n. sp. Elytra: approximately two times longer (0.64–0.85 mm) than wide (0.32–0.41 mm), subparallel. Lateral margins with 18–21 subtriangular denticles, faint and progressively smoother towards posterior, but still marked in last third. Apical margin smoothly rounded, with three to six pairs of small denticles (showing a notable range of individual variation, but mostly combinations of four and five pairs) (Fig. 7 A, B). Transverse scutellar organ with straight to slightly subtriangular margin and pair of ‘buttonholes’ present near base of elytra. Chaetotaxy: umbilicate series formed by anterior group with four setae and posterior group with two setae (Fig. 4). One pair of scutellar setae. No apparent discal setae, but the population of Puerto de los Reventones is prone to have one pair of anterior “pseudodiscals” (i.e. pubescent setae longer than the surrounding pubescence). Rest of characters as in T. acutangula n. sp. Legs: similar in both sexes. Intermetacoxal space not widened. Metacoxae with smoothly rounded “flap”. Rounded metatrochanters, metatibiae with moderately dilated distal region and a short perpendicular seta in the inner side. Inner side of pro- and mesofemora covered with small protuberances, absent in metafemora. Pseudotetramerous tarsi in all the legs, tarsomeres 4 º and 5 º retain a faint articular membrane, but are partially fused and not capable of a normal articulation. Abdomen: as described for T. acutangula n. sp., but with deep ventral foveae in the first and second ventrite of females, absent in males. Last ventrite with a sexually dimorphic pattern of five to six pairs of setae in posterior margin: l-s-s-l-s-m / m-(s)-l-s-s-l (Fig. 4). Aedeagus: sickle-shaped median lobe (length: 0.22 mm) and “bottleneck-shaped”, smoothly rounded apex, slightly bent to right (anatomically oriented) in dorsal view (Fig. 5 A,B). Endophallus with ‘plough-shaped’ sclerites, as drawn in Fig. 5 B. Short and broad parameres, both with two medium-sized apical setae (Fig. 5 B). Female genitalia adjusts to the description of Vigna-Taglianti (1972). Long tubular gonocoxites, with double apical setae. Lateral setae absent, but lateral pores in the gonocoxites. Short gonoduct with two regions, thinner proximally (diameter 0.003 mm) and thicker distally (diameter 0.008 mm). Spermatheca subcylindrical and short, “barrel-shaped” (length: 0.011 mm). Conical spermathecal gland (length: 0.014 mm), sclerotized in distal region (Fig. 5 C). Etymology. The name of the species plays a double meaning, referring to the high intraspecific variation that gives a mixture of body shapes and sizes to the populations. On the other hand, it refers to the presence of certain morphologic traits (such as the tooth in clypeus, the trend to diffuse gula or the pseudotetramery) unexpected for members of baetica group, but known in other species groups, like mixed pieces in different jigsaw puzzles. Habitat. Typhlocharis mixta n. sp. is found in two localities separated 13 km (Fig. 1): 1) Puerto de los Reventones, in open prairie environment with low hills and valleys crossed by seasonal streams. Vegetation mainly composed by low herbaceous plants, grasses and thistles, with scattered broom bushes (Retam Raf.). Higher grasses and rushes (Juncus L.) appears near the stream. Dark, brownish soil, with scarce large boulders of gneisses and shales (Proterozoic, Valverde Formation). This location was sampled four times (March, April, June and November), with the best humidity range in the last sampling. 2) Valverde de Burguillos, in a small meadow surrounded by cultures and open holm oak forest. Vegetation is akin to the former locality, with the exception of holm oaks. Brown to reddish soil, rich in clays, with embedded small to medium sized granite boulders (Proterozoic). Variability. There are not significant differences between the populations of Puerto de los Reventones (1285 specimens) and Valverde de Burguillos (21 specimens), both have the same range of intraspecific variation. The species is morphologically very diverse in proportions and size, from 1.2 to 1.6 mm in length, but the presence of a tooth in clypeus, umbilicate series 4 + 2, the perpendicular setae on distal end of metatibiae, pseudotetramerous tarsi, ventral foveae in females, genitalia and the wide hiatus are virtually constant within the population. The terebral tooth is usually very prominent but in some specimens is blunt, low or almost absent. The tooth of clypeus ranges from blunt to very pointy (but always present). The lateral sutures of the gula are very faint in about 20 % of the series, resembling the “difusse gula” morphology observed in quadridentata group (Pérez-González & Zaballos 2013 c). There is slight variation in the pronouncement of epilobes. Two specimens have a widened pars stridens, two times wider than the average. Pronotum and elytra have the higher rates of variability. Pronotum proportions can be certainly different, with three main morphs: subrectangular, smoothly curved lateral margins and 3-4 well marked posterolateral denticles (Fig 6: 1,2), the most common; rectangular, straight lateral margins and 4-5 (even 6 in one specimen) well marked denticles (Fig 6: 3,4) and subsquare, with 2-3 blunt and irregular denticles (Fig 6: 5,6). Asymmetric pronotum are abundant, and they affect the number of denticles or the pronouncement of the posterior angle (Fig 6: 7). Five specimens have aberrant pronotums with widened, notched or asymmetric lateral margins (Fig 6: 8–10). Transverse scutellar organ varies from straight to subtriangular and the margins can be irregular or smooth. Lateral denticles of elytra can be more or less marked. Apical denticles of elytra appear in a wide range of combinations and shapes (Fig. 7): the most common are 4 / 4 (17 % of the series), 4 / 5 (15 %), 5 / 4 (15 %), and 5 / 5 (22 %), but they range from 2 / 3 to 7 / 7. All the observed combinations and their abundance within the population are expressed in Fig. 7 B. The inner pair, closer to the elytral suture is especially developed and bigger than the other denticles (Fig. 7 A: 3) in some specimens, resembling those of Typhlocharis tertia Pérez-González & Zaballos, 2013. About 14 % of the series (145 specimens) have bifid or double denticles and 7 specimens show triple denticles (Fig. 7 A: 4-6). Asymmetries (Fig. 7 A: 7–9) and damaged elytra are also common. Variations in patterns of chaetotaxy occur in labium, prebasilar, last ventrite and sensilla coeloconica of the 11 th antennomere. It is remarkable that the ventral sensilla coeloconica in 5 º and 6 º antennomeres also varies: 7 specimens have only sensilla coeloconica in the 5 º antennomere and 3 specimens have them in antennomeres 5 º, 6 º and 7 º. It is the first evidence of variation in this feature within the same species, and the presence of sensilla coeloconica in 5 º antennomere but not in 6 º has been reported only in Typhlocharis tetramera Pérez-González & Zaballos, 2013. Finally, there are several specimens with teratologies, like antennomere fusion (5 specimens, with fused 3 º- 4 º, 5 º- 6 º, 9 º- 10 º and 10 º- 11 º antennomeres respectively), or malformed mesotibiae (1 specimen, associated with a malformed pronotum, possible result of a failed moult). Affinities. Typhlocharis mixta n. sp. presents the diagnostic characters of baetica group, defined by the presence of several pairs of apical denticles. Affinities with other species of the group are the umbilicate series of 4 + 2, shared with T. pacensis Zaballos & Jeanne, 1987; T. furnayulensis Zaballos & Banda, 2001; T. secunda Perez- González & Zaballos, 2013 and T. acutangula n. sp. The protuberances in femora also appears in T. baetica Ehlers, 1883; T. tertia Perez-González & Zaballos, 2013; T. quarta Perez-González & Zaballos, 2013 and T. acutangula n. sp.; the high number and serrated aspect of posterolateral denticles of pronotum are also typical of T. baetica and T. quarta, but many specimens have pronotum with few and less defined denticles akin to that of T. tertia and the other species of baetica group. The broad parameres are unusual within the group and strongly resembles those of T. tertia. T. secunda Perez-González & Zaballos, 2013 is very similar: both species shares the same umbilicate series, size range, overall shape of pronotum posterolateral denticles and presence (not always in T. mixta) of anterior “pseudodiscal” setae. However, the new species is well differentiated by its diagnostic features. The tooth of clypeus, subcylindrical spermatheca, metatibial perpendicular short seta and the pseudotetramerous tarsi found in T. mixta n. sp. are unique of this species within baetica group, but appear in other Typhlocharis species. The tooth of clypeus is the definitory trait of the six species of gomezi group (Zaballos & Farinós 1995; Zaballos & Ruiz-Tapiador 1997) which also have a short perpendicular seta in metatibiae. The trend of diffuse gula in some specimens, the variability of ventral sensilla coeloconica in antennomeres 5 º and 6 º and the pseudotetramerous tarsi are features previously observed only in species of quadridentata group (Pérez-González & Zaballos 2013 c). In both cases the possible homology of these characters remains unclear. Key to species of baetica group (update of Pérez - González & Zaballos 2013 a) 1. Elytra with umbilicate series 4 + 4...................................... T. quarta Pérez-González & Zaballos, 2013 - Elytra with umbilicate series different to 4 + 4.............................................................. 2 2. Elytra with umbilicate series 4 + 3........................................................................ 3 - Elytra with umbilicate series 4 + 2 or 4 + 1.................................................................. 6 3. Inner side of femora with protuberances, warty surface. Length> 1.4 mm....................................... 4 - Inner side of femora without protuberances, smooth surface. Length 4. Apical margin of elytra with 6–7 pairs of small denticles. Length 1.5......................... T. baetica Ehlers, 1883 - Apical margin of elytra with 3 pairs of large denticles. Length 1.7 mm.......... T. tertia Pérez-González & Zaballos, 2013 5. Elytra with lateral denticles marked in all their length. Very faint or absent ventral foveae. Length 1.1 mm........................................................................................ T. aguirrei Zaballos & Banda, 2001 - Elytra with lateral denticles marked only in the first third of their length. Ventral foveae well developed in both sexes. Length 1.3 mm T. matiasi Zaballos & Banda, 2001 6. Elytra with umbilicate series 4 + 2........................................................................ 7 - Elytra with umbilicate series 4 + 1....................................................................... 11 7. Clypeus with a median tooth. Ventral foveae in females. Length 1.2–1.6 mm........................... T. mixta n. sp. - Clypeus with straight margin, without median tooth........................................................... 8 8. Metatibiae with a pointy dentiform projection in the distal end. Ventral foveae absent. Length 1.1–1.3 mm..................................................................................................... T. acutangula n. sp. - Metatibiae without dentiform projection.................................................................... 9 9. Elytra with 1 pair of discal setae in the anterior third. Length 1.4 mm.......... T. secunda Pérez-González & Zaballos, 2013 - Elytra without discal setae. Length 10. Apical margin of elytra with 4–5 pairs of small denticles. Diffuse endophallus sclerites. Length 0.9–1 mm........................................................................................ T. pacensis Zaballos & Jeanne, 1987 - Apical margin of elytra with 5–7 pairs of small apical denticles. “Y-shaped” endophallus sclerites. Length 1.1 mm............................................................................. T. furnayulensis Zaballos & Banda, 2001 11. Elytra with lateral denticles marked in all their length. Thin, “Y-shaped” endophallus sclerites. Subspheric-irregular spermath- eca. Length 1.1 mm T. millenaria Zaballos & Banda, 2001 - Elytra with lateral denticles marked only in the first third of their length. Thick, stick-shaped endophallus. Reniform spermath- eca. Length 1.1 mm................................................. T. prima Pérez-González & Zaballos, 2013

Typhlocharis acutangula n. sp. Figs 2, 3 Type series. Holotype, ♂, SPAIN, Huelva, San Silvestre de Guzm��n (0.7 km N), 21 - 11-2012, 175 m (37 �� 23 ��� N, 07�� 21 ��� W), J.P. Zaballos, S. P��rez & S. Ghannem leg. (Coll. J.P. Zaballos, UCM). Paratypes: 18 ♂♂, 13 ♀♀ same data as holotype (Coll. J.P. Zaballos and Coll. S. P��rez-Gonz��lez, UCM). 1 ♂, 1 ♀ same data as holotype (MNCN, Madrid, Catalogue number: 9999). Diagnosis. Small endogean beetle, anophthalmous, with narrow and subrectangular body covered by microreticulate integument and scattered pubescence. Vertex with pars stridens. Elytra well serrated in lateral margins and apical margin with four to six pairs of small denticles. Umbilicate series with six setae (4 + 2). Metatibiae with a distinct dentiform projection at the anterodistal end. Abdomen without ventral foveae. Aedeagus as in Fig. 3, ���stick-shaped��� endophallic sclerite. Female genitalia as in Fig. 3, with ovoid spermatheca. Description. Length 1.14���1.35 mm (males), 1.17���1.35 mm (females) (Fig. 2). Head: wider (0.25���0.30 mm) than long (0.24���0.27 mm), with subhexagonal microsculpture and stridulatory organ (pars stridens) in vertex region, in both sexes. Posterolateral semilunar notch in both sides of cephalic capsule. Subrectangular labrum, with rounded edges and triangular area with middle button of thicker cuticle. Clypeus with straight anterior margin. Moniliform antennae with 11 antennomeres progressively more quadrate in lateral view (morph 1 sensu P��rez- Gonz��lez & Zaballos 2013 b), last one pyriform. Pedicel of third antennomere not elongate. Sensilla coeloconica of last antennomere distributed in a three anterodorsal and one posterodorsal pattern. Ventral sensilla coeloconica in antennomeres 5 �� and 6 ��. Right mandible with a low, rounded terebral tooth on inner margin. Left mandible without teeth. Labium without special characters, the middle tooth is slightly rounded. Ligula with moderately prominent middle lobe and long paraglossae. Wide gula approximately twice as long as wide. Cephalic chaetotaxy: six pairs of labral setae (s-s-l-m-s-m / m-s-m-l-s-s), two pairs of clypeal setae (l-s / s-l), one pair of frontal setae, two pairs of supraocular setae (anterior and posterior), one pair of supraantennal setae, three pairs of occipital setae and two pairs of genal setae, as well as scattered pubescence. Labium with a pair of setae near base of the middle tooth, a pair of long setae near base of epilobes, a pair of very short setae near apex of epilobes and one or two pairs of very short setae near posterior suture. Prebasilar with a pair of lateral long setae near anterior margin, a pair of very short lateral setae in mid-region and two pairs of setae (lateral pair much longer) in posterior region, with some degree of individual variation over this basic pattern. Pronotum: subquadrate, longer (0.31���0.38 mm) than wide (0.30���0.35 mm). Anterior and posterior margins smoothly sinuate. Anterior margin slightly crenulate, with medial hiatus (approximately as wide as two adjacent intersetae spaces). Lateral margins with two or three denticles near posterior angles, blunt and irregular. Surface covered by subhexagonal microreticulation. Disc flattened, with medial line and pair of faint lateral sulci. Chaetotaxy: one pair of long setae in first third of lateral margins, one pair of long setae near posterior angles, a row of five to six pairs of setae [l-(l)-l-l-l-l / l-l-l-l-(l)-l] parallel to anterior margin, two, rarely three pairs of setae parallel to posterior margin [(s)-l-l / l-l-(s)], row of small, thin setae regularly placed in anterior (bulb shaped) and posterior (filiform) margins, row of short setae in lateral margins, and four or five pairs of longitudinal rows of short pubescence in disc. Proepisternal suture marked. Prosternal apophysis rounded. Anterior margin of prosternum with a row of long, thin setae and six to eight pairs of short setae parallel to it. Prosternum covered in scattered pubescence, absent in proepisternum. Elytra: approximately two times longer (0.60���0.72 mm) than wide (0.33���0.39 mm), subparallel. Lateral margins with 19���21 very strong denticles, ���hooked��� in the first third progressively less marked toward posterior, but reaching the apical region to the beginning of the apical denticles. Apical margin smoothly rounded, with four to six pairs of small denticles (affected by individual variation, five pairs in average). Disc flattened, with longitudinal lateral carinae associated to seventh stria but not reaching the apical margin. Surface with irregular subhexagonal microreticulation. Faint elytral pores, scattered along seventh stria, scutellar region and disc. Transverse scutellar organ with straight margin, and pair of ���buttonholes��� present near base of elytra. Chaetotaxy: umbilicate series composed by anterior group of four setae and posterior group of two setae (Fig. 2). One pair of scutellar setae. No discal setae. Discal pubescence arranged in five or six pairs of longitudinal rows. Apical row of short pubescent setae. Lateral margins with a short seta for every denticle, forming a row. Legs: similar in both sexes. Intermetacoxal space not widened. Metacoxae with smoothly rounded ���flap��� (a flat extension of the margin of metacoxae, overlapping over the base of metatrochanters, Fig. 2). Rounded metatrochanters, slightly angular metafemora and metatibiae with dilated anterodistal region, ending in a prominent dentiform projection. Inner side of meso- and metafemora covered by small scaly protuberances. Clearly pentamerous tarsi in all the legs. Abdomen: covered by irregular microreticulation, last segment with ���belt��� of scaly microsculpture, every scale with finely serrated edge in both sexes. Ventral foveae absent. Last ventrite without lateral teeth, pattern of five to six pairs of setae in posterior margin, sexually dimorphic: l-s-s-l-s / m-s-l-s-s-l (Fig. 2). Aedeagus: sickle-shaped median lobe (length: 0.18 mm) and subtriangular, smoothly rounded apex, bent to right (anatomically oriented) in dorsal view (Fig. 3 A, B). Endophallus with large and complex ���branched, stick-shaped��� sclerites (Fig. 3 B), with lateral projection smoothly curved upwards. Parameres subtriangular, with two medium-sized apical setae (Fig. 3 B). Female genitalia as described by Vigna-Taglianti (1972). Long tubular gonocoxites, with double apical setae. Lateral setae absent, but lateral pores in the gonocoxites. Short gonoduct with two regions: a thinner proximal (diameter 0.0017 mm) and thicker distal region (diameter 0.0045 mm). Irregularly ovoid spermatheca (length: 0.014 mm). Conical spermathecal gland (length: 0.016 mm), sclerotized in distal region (Fig. 3 C). Etymology. The name ��� acutangula ��� is referred to the characteristic diagnostic feature of the species, the prominent dentiform projection that gives the ���acute angled��� shape to the anterodistal region of metatibiae. Habitat. The type locality (Fig. 1) is found in open holm oak (Quercus ilex L.) environment, with abundant laudanum shrubs (Cistus ladanifer L.) and scarce asphodels (Asphodelus L.) and thistles. Reddish soil, with high clay content and high humidity at the moment of sampling, filled with various-sized boulders of breakable shales (Upper Carboniferous). The sample was taken under big boulders next to young holm-oak trees and laudanum shrubs. Variability. The range of variability observed in Typhlocharis acutangula n. sp. (34 specimens) is similar to that described for other species of baetica group (P��rez-Gonz��lez & Zaballos 2013 a). It mainly affects to the shape and number of posterolateral denticles of pronotum (two or three, irregularly marked); the degree of pronunciation of the lateral denticles of elytra (two specimens have blunt, worn lateral denticles, other two have denticles less marked than average, not hooked on the first third), the shape of transverse scutellar organ (from straight to subtriangular, rarely irregular), the shape of median lobe of ligula (very low in some specimens) and the apodemal ring of male genitalia. Variation in the apical denticles of elytra is very common, with individuals having four, five or six pairs. Asymmetries in the number of denticle pairs in each elytron are abundant and the observed combinations includes 4 /6, 5/4, 4/5, 6/ 5 and 5 / 7. Bifid or double denticles appear in two specimens. The most common combinations are 5 / 4 (17 % of the series) and 5 / 5 (32 % of the series) denticles. Chaetotaxy of labium, prebasilar and last ventrite shows subtle variations. There are asymmetric alterations of the pattern of sensilla coeloconica in 11 th antennomere in two specimens. Affinities. Typhlocharis acutangula n. sp. is included in the baetica group by the presence of four to six pairs of apical denticles in elytra (Zaballos & Banda 2001; P��rez-Gonz��lez & Zaballos 2013 a). Within the group, it shares an umbilicate series of 4 + 2 with T. pacensis Zaballos & Jeanne, 1987; T. furnayulensis Zaballos & Banda, 2001 and T. secunda Perez-Gonz��lez & Zaballos, 2013. The latter is the morphologically closest species, both sharing a similar shape of aedeagus, parameres and endophallic sclerites. The presence of protuberances in the inner margin of femora is shared with T. baetica Ehlers, 1883; T. tertia Perez-Gonz��lez & Zaballos, 2013 and T. quarta Perez-Gonz��lez & Zaballos, 2013. The new species can be immediately distinguished by the presence of the pointy dentiform projection in the anterodistal end of metatibiae, a feature only known in other species of the genus, T. crespoi Serrano & Aguiar, 2008, of quadridentata group (P��rez-Gonz��lez & Zaballos 2013 c). It is worth noting that other rare character, the bulb shaped setae in the anterior margin of pronotum is shared with T. baeturica Perez-Gonz��lez & Zaballos, 2013, included in quadridentata group., Published as part of P��rez-Gonz��lez, Sergio, Zaballos, Juan P. & Ghannem, Samir, 2013, Intraspecific variation in Typhlocharis Dieck, 1869 (Coleoptera, Carabidae, Anillini): the case of two new species of the baetica group in Zootaxa 3710 (1), DOI: 10.11646/zootaxa.3710.1.3, http://zenodo.org/record/223802

Typhlocharis acutangula n. sp. Figs 2, 3 Type series. Holotype, ♂, SPAIN, Huelva, San Silvestre de Guzmán (0.7 km N), 21 - 11-2012, 175 m (37 º 23 ’ N, 07º 21 ’ W), J.P. Zaballos, S. Pérez & S. Ghannem leg. (Coll. J.P. Zaballos, UCM). Paratypes: 18 ♂♂, 13 ♀♀ same data as holotype (Coll. J.P. Zaballos and Coll. S. Pérez-González, UCM). 1 ♂, 1 ♀ same data as holotype (MNCN, Madrid, Catalogue number: 9999). Diagnosis. Small endogean beetle, anophthalmous, with narrow and subrectangular body covered by microreticulate integument and scattered pubescence. Vertex with pars stridens. Elytra well serrated in lateral margins and apical margin with four to six pairs of small denticles. Umbilicate series with six setae (4 + 2). Metatibiae with a distinct dentiform projection at the anterodistal end. Abdomen without ventral foveae. Aedeagus as in Fig. 3, ‘stick-shaped’ endophallic sclerite. Female genitalia as in Fig. 3, with ovoid spermatheca. Description. Length 1.14–1.35 mm (males), 1.17–1.35 mm (females) (Fig. 2). Head: wider (0.25–0.30 mm) than long (0.24–0.27 mm), with subhexagonal microsculpture and stridulatory organ (pars stridens) in vertex region, in both sexes. Posterolateral semilunar notch in both sides of cephalic capsule. Subrectangular labrum, with rounded edges and triangular area with middle button of thicker cuticle. Clypeus with straight anterior margin. Moniliform antennae with 11 antennomeres progressively more quadrate in lateral view (morph 1 sensu Pérez- González & Zaballos 2013 b), last one pyriform. Pedicel of third antennomere not elongate. Sensilla coeloconica of last antennomere distributed in a three anterodorsal and one posterodorsal pattern. Ventral sensilla coeloconica in antennomeres 5 º and 6 º. Right mandible with a low, rounded terebral tooth on inner margin. Left mandible without teeth. Labium without special characters, the middle tooth is slightly rounded. Ligula with moderately prominent middle lobe and long paraglossae. Wide gula approximately twice as long as wide. Cephalic chaetotaxy: six pairs of labral setae (s-s-l-m-s-m / m-s-m-l-s-s), two pairs of clypeal setae (l-s / s-l), one pair of frontal setae, two pairs of supraocular setae (anterior and posterior), one pair of supraantennal setae, three pairs of occipital setae and two pairs of genal setae, as well as scattered pubescence. Labium with a pair of setae near base of the middle tooth, a pair of long setae near base of epilobes, a pair of very short setae near apex of epilobes and one or two pairs of very short setae near posterior suture. Prebasilar with a pair of lateral long setae near anterior margin, a pair of very short lateral setae in mid-region and two pairs of setae (lateral pair much longer) in posterior region, with some degree of individual variation over this basic pattern. Pronotum: subquadrate, longer (0.31–0.38 mm) than wide (0.30–0.35 mm). Anterior and posterior margins smoothly sinuate. Anterior margin slightly crenulate, with medial hiatus (approximately as wide as two adjacent intersetae spaces). Lateral margins with two or three denticles near posterior angles, blunt and irregular. Surface covered by subhexagonal microreticulation. Disc flattened, with medial line and pair of faint lateral sulci. Chaetotaxy: one pair of long setae in first third of lateral margins, one pair of long setae near posterior angles, a row of five to six pairs of setae [l-(l)-l-l-l-l / l-l-l-l-(l)-l] parallel to anterior margin, two, rarely three pairs of setae parallel to posterior margin [(s)-l-l / l-l-(s)], row of small, thin setae regularly placed in anterior (bulb shaped) and posterior (filiform) margins, row of short setae in lateral margins, and four or five pairs of longitudinal rows of short pubescence in disc. Proepisternal suture marked. Prosternal apophysis rounded. Anterior margin of prosternum with a row of long, thin setae and six to eight pairs of short setae parallel to it. Prosternum covered in scattered pubescence, absent in proepisternum. Elytra: approximately two times longer (0.60–0.72 mm) than wide (0.33–0.39 mm), subparallel. Lateral margins with 19–21 very strong denticles, “hooked” in the first third progressively less marked toward posterior, but reaching the apical region to the beginning of the apical denticles. Apical margin smoothly rounded, with four to six pairs of small denticles (affected by individual variation, five pairs in average). Disc flattened, with longitudinal lateral carinae associated to seventh stria but not reaching the apical margin. Surface with irregular subhexagonal microreticulation. Faint elytral pores, scattered along seventh stria, scutellar region and disc. Transverse scutellar organ with straight margin, and pair of ‘buttonholes’ present near base of elytra. Chaetotaxy: umbilicate series composed by anterior group of four setae and posterior group of two setae (Fig. 2). One pair of scutellar setae. No discal setae. Discal pubescence arranged in five or six pairs of longitudinal rows. Apical row of short pubescent setae. Lateral margins with a short seta for every denticle, forming a row. Legs: similar in both sexes. Intermetacoxal space not widened. Metacoxae with smoothly rounded “flap” (a flat extension of the margin of metacoxae, overlapping over the base of metatrochanters, Fig. 2). Rounded metatrochanters, slightly angular metafemora and metatibiae with dilated anterodistal region, ending in a prominent dentiform projection. Inner side of meso- and metafemora covered by small scaly protuberances. Clearly pentamerous tarsi in all the legs. Abdomen: covered by irregular microreticulation, last segment with ‘belt’ of scaly microsculpture, every scale with finely serrated edge in both sexes. Ventral foveae absent. Last ventrite without lateral teeth, pattern of five to six pairs of setae in posterior margin, sexually dimorphic: l-s-s-l-s / m-s-l-s-s-l (Fig. 2). Aedeagus: sickle-shaped median lobe (length: 0.18 mm) and subtriangular, smoothly rounded apex, bent to right (anatomically oriented) in dorsal view (Fig. 3 A, B). Endophallus with large and complex “branched, stick-shaped” sclerites (Fig. 3 B), with lateral projection smoothly curved upwards. Parameres subtriangular, with two medium-sized apical setae (Fig. 3 B). Female genitalia as described by Vigna-Taglianti (1972). Long tubular gonocoxites, with double apical setae. Lateral setae absent, but lateral pores in the gonocoxites. Short gonoduct with two regions: a thinner proximal (diameter 0.0017 mm) and thicker distal region (diameter 0.0045 mm). Irregularly ovoid spermatheca (length: 0.014 mm). Conical spermathecal gland (length: 0.016 mm), sclerotized in distal region (Fig. 3 C). Etymology. The name “ acutangula ” is referred to the characteristic diagnostic feature of the species, the prominent dentiform projection that gives the “acute angled” shape to the anterodistal region of metatibiae. Habitat. The type locality (Fig. 1) is found in open holm oak (Quercus ilex L.) environment, with abundant laudanum shrubs (Cistus ladanifer L.) and scarce asphodels (Asphodelus L.) and thistles. Reddish soil, with high clay content and high humidity at the moment of sampling, filled with various-sized boulders of breakable shales (Upper Carboniferous). The sample was taken under big boulders next to young holm-oak trees and laudanum shrubs. Variability. The range of variability observed in Typhlocharis acutangula n. sp. (34 specimens) is similar to that described for other species of baetica group (Pérez-González & Zaballos 2013 a). It mainly affects to the shape and number of posterolateral denticles of pronotum (two or three, irregularly marked); the degree of pronunciation of the lateral denticles of elytra (two specimens have blunt, worn lateral denticles, other two have denticles less marked than average, not hooked on the first third), the shape of transverse scutellar organ (from straight to subtriangular, rarely irregular), the shape of median lobe of ligula (very low in some specimens) and the apodemal ring of male genitalia. Variation in the apical denticles of elytra is very common, with individuals having four, five or six pairs. Asymmetries in the number of denticle pairs in each elytron are abundant and the observed combinations includes 4 /6, 5/4, 4/5, 6/ 5 and 5 / 7. Bifid or double denticles appear in two specimens. The most common combinations are 5 / 4 (17 % of the series) and 5 / 5 (32 % of the series) denticles. Chaetotaxy of labium, prebasilar and last ventrite shows subtle variations. There are asymmetric alterations of the pattern of sensilla coeloconica in 11 th antennomere in two specimens. Affinities. Typhlocharis acutangula n. sp. is included in the baetica group by the presence of four to six pairs of apical denticles in elytra (Zaballos & Banda 2001; Pérez-González & Zaballos 2013 a). Within the group, it shares an umbilicate series of 4 + 2 with T. pacensis Zaballos & Jeanne, 1987; T. furnayulensis Zaballos & Banda, 2001 and T. secunda Perez-González & Zaballos, 2013. The latter is the morphologically closest species, both sharing a similar shape of aedeagus, parameres and endophallic sclerites. The presence of protuberances in the inner margin of femora is shared with T. baetica Ehlers, 1883; T. tertia Perez-González & Zaballos, 2013 and T. quarta Perez-González & Zaballos, 2013. The new species can be immediately distinguished by the presence of the pointy dentiform projection in the anterodistal end of metatibiae, a feature only known in other species of the genus, T. crespoi Serrano & Aguiar, 2008, of quadridentata group (Pérez-González & Zaballos 2013 c). It is worth noting that other rare character, the bulb shaped setae in the anterior margin of pronotum is shared with T. baeturica Perez-González & Zaballos, 2013, included in quadridentata group.