Vaejovinae Thorell, 1876

Chávez-Samayoa, Fernanda, Díaz-Plascencia, José Eduardo & González-Santillán, Edmundo, 2022, Two new species of Vaejovis (Scorpiones: Vaejovidae) belonging to the mexicanus group from Aguascalientes, Mexico, with comments on the homology and function of the hemispermatophore, Zoologischer Anzeiger (Zool. Anz.) 298, pp. 148-169 : 151-164

Subfamily Vaejovinae Thorell, 1876 . Genus Vaejovis Thorell, 1876 . Vaejovis aguazarca Díaz-Plascencia and Gonz´alez-Santill´an sp. nov. LSID: urn:lsid:zoobank.org:act:0C495778–F61D-4049-973E-6351BC

0BFA52.

Fig. 1C, D , 2A , 3 , 4A , 5A , 6A , 7-9 , 10A, B , 11A , 12A , 13 A-C, 14A, 22,

23; Tables 1–3; Appendix A. Type Material: M´exico: Aguascalientes: Municipio San Jose ´de

Gracia: Holotype 6 (CNAN-T01492) Sierra Fría, Estaci´on Biologica´Agua

Zarca, 22.090555 ◦ N 102.556298 ◦ W, 2173 m, 31.vii.2015, J. E. DíazPlascencia and J. L. Aguilar. Paratypes 16, 4♀ ( CNAN-T01493 ), 22.09221 ◦ N 102.55774 ◦ W, 2190 m, 7.ix.2020. F. Ch´avez-Samayoa and D. F. Simijaca-Salcedo; 16, 1♀ ( CNAN-T01494 ), 22.09255 ◦ N 102.55728 ◦ W, 2190 m, 9.xi.2020, F. Ch´avez-Samayoa, D. F. Simijaca-Salcedo, and P. García-Macías; 16 ( CNAN-T01495 ), 22.09294 ◦ N 102.56112 ◦ W, 2181m, 19.ii.2021, F. Ch´avez-Samayoa and D. F. Simijaca-Salcedo .

Additional Material Examined: Mexico: Aguascalientes: Municipio Jesús María : 1♀ ( CZUUA SCO-382 ), Presa Abelardo Rodríguez , 21 ◦ 55’7.64"N 102 ◦ 24’28.62"W, 1927 m, 8.vi.2003. J. E. Valencia ; 1♀ ( CZUAA SCO-583 ), Los Arquitos , 21.921649 ◦ N 102.397052 ◦, 1935m, viii.2020, F. Ch´avez-Samayoa, S. S. Guti´errez-Ramírez, and D. F. Simijaca-Salcedo ; 1♀ ( CZUAA SCO-606 ), 21.921649 ◦ N 102.397052 ◦ W, 1935 m, 28.xi.2020, F. Ch´avez-Samayoa and D. F. Simijaca-Salcedo. Municipio Rincon ´de Romos : 1 ♀ ( CZUUA SCO-386 ), Pabell´on de Hidalgo, 22 ◦ 11’18.48"N 102 ◦ 21’22.56"W, 2046 m, 15.v.2013. H. Puga- Ar´evalo. Municipio San Francisco de los Romo : 1 ♀ ( CZUAA SCO-500 ), El Chicalote, 22.012016 ◦ N 102.253290 ◦ W, 1896 m, x.2019., M. J. Perales- Olivares. Municipio San Jos´ e de Gracia: 3 6 , 6 ♀ ( CZUAA SCO-552 ), Estaci´on Biologica ´Agua Zarca , 22.09221 ◦ N 102.55774 ◦ W, 2190 m, 7. ix.2020. F. Ch´avez-Samayoa and D. F. Simijaca-Salcedo; 56 ( CZUAA SCO-571 ), 22.09255 ◦ N 102.55728 ◦ W, 2190 m, 9.xi.2020, F. Ch´avez-Samayoa, D. F. Simijaca-Salcedo, and P. García-Macías; 1 6 ( CZUAA-608 ), 22.09294 ◦ N 102.56112 ◦ W, 2181 m, 19.ii.2021. F. Ch´avez-Samayoa and D. F. Simijaca-Salcedo; 1 6 , 2 ♀ ( CNAN-S4005 ), 22.092702 ◦ N 102.564780 ◦ W, 2201 m, 15.v.2015, J. E. Díaz-Plascencia, and C. F. Servín de la Mora .

Etymology: The specific epithet is used as a noun in apposition and refers to the type locality Estacion´Biol´ogica Agua Zarca where the type series was collected. The English translation of Zarca is “light blue”.

Diagnosis: Vaejovis aguazarca sp. nov. can be distinguished from V. aquascalentensis sp. nov. by the presence of one distomedial macroseta on the cheliceral manus instead of two ( Fig. 4A, B ). The pedipalp femur retrolateral dorsosubmedian carina ( Fig. 7A, B ) is serrated in both sexes, whereas in V. aquascalentensis sp. nov. it is weakly granular ( Fig. 16A, B ). The patella dorsal retrolateral carinae is granular ( Fig. 8A, B), instead of weakly granular ( Fig. 17A, B ); prolateral median carinae with a row of five to eight coarse granules ( Fig. 8C, D) instead of obsolete, and delimited by two large granules instead of a short row of granules as in V. aquascalentensis sp. nov ( Fig. 17C, D ). Chela manus dorsal and retrolateral intercarinal surfaces matte, finely and sparsely granular in V. aguazarca sp. nov ( Fig. 9A, B ). instead of smooth as in V. aquascalentensis sp. nov ( Fig. 18A, B ); prolateral dorsal, dorsal prosubmedian, and dorsal prolateral carinae strongly granular ( Fig. 9 ) instead of low granules ( Fig. 18 ) in V. aquascalentensis sp. nov. Movable finger with seven prolateral and six retrolateral denticles instead of seven prolateral and retrolateral denticles ( Figs. 10A, D ).

Vaejovis aguazarca sp. nov. is readily distinguished from V. tenamaztlei by the following features: anterior margin of carapace bilobed with a deep median notch instead of nearly straight with a shallow notch ( Fig. 5A, C ). Pedipalp manus carinae, dorsal, and retrolateral surfaces of femur and patella densely infuscate instead of infuscate only at the base of the macrosetae (Contreras-F´elix et al., 2015; Plate 1). Telotarsi with ventral spinules aligned in a single row instead of an unaligned cluster of spinules; one pair of ventrodistal spinules instead of two or three pairs. Metasoma segment IV lateral posterior end with one medial seta, absent in V. tenamaztlei . Greater total body length at 28.4–32.5 (6) or 29.3–41.1 (♀) instead of 19.4–22.1 (6) and 21.5–26.6 (♀).

Hemispermatophore laminar antero-distal process prominent in V. aguazarca sp. nov. instead of obsolete as in V. aquascalentensis sp. nov., or weak as in V. tenamaztlei ( Fig. 2 ). Clasper prominent and spiniform in V. aguazarca sp. nov. instead of inconspicuous as in V. aquascalentensis sp. nov. and V. tenamaztlei ( Fig. 2 ). Distal lamina basal lateral trough shallow in V. aguazarca sp. nov. and V. tenamaztlei instead of deep as in V. aquascalentensis sp. nov. Capsular basal carina with circa 13 hooklets in V. aguazarca sp. nov ( Fig. 1C ), absent in V. aquascalentensis sp. nov. and V. tenamaztlei ( Fig. 2 ).

Vaejovis aguazarca sp. nov. can be differentiated from the geographically close ( Fig. 22 ) species Vaejovis dugesi Pocock, 1902 from Guanajuato by a higher pectinal count, at 15–16 (6), 13–14 (♀) instead of 13-13 (6), 11–12 (♀) [Sissom, 1990]. All carinae of metasoma irregularly infuscate, even more so on the setae insertions instead of completely infuscate; dorsolateral carinae of segments I-IV with distal granules enlarged ( Fig. 13B ) instead of reduced; ventral median carinae of segment V complete granular instead of incomplete, present on anterior fourth fifths. Telson dorsal surface smooth without glandular area ( Fig. 14A ), instead of large glandular area (Contreras-F´elix and Francke 2019, Fig. 7a ).

Description: The following description is based on the type series including adults of both sexes and additional material examined.

Color and infuscation: Body base color yellowish (orange in alcohol). Cheliceral manus dorsal surface with faintly reticulate infuscation, base and distal margin of fixed fingers infuscate. Carapace with dusky markings ( Fig. 3A, C ). Pedipalps, all carinae of chela strongly infuscate except prolateral median and fingers faintly infuscate. Patella, femur, and legs densely but faintly infuscate. Coxosternal region, genital operculum, and pectines immaculate. Mesosomal tergites with two pairs of narrow, longitudinal infuscation along lateral margin and submedian surfaces; tergites I-VII dorsal median surface feebly infuscate more so on pretergites, sternites III-VII surfaces immaculate. Metasomal segments I–V with all dorsal carinae irregularly infuscate anteriorly and on setae insertions; intercarinal dorsal surfaces immaculate, lateral, and ventral densely infuscate, more so on segments IV and V lateral median, lateral inframedian, ventral lateral, and ventral submedian carinae irregularly to completely infuscate ( Fig. 3B, D ). Telson vesicle base color yellowish orange, ventral surface with three vestigial to obsolete longitudinal infuscation, setae insertions infuscate, median surface whitish except 30 to 40 percent of margins densely infuscate (6) or mostly infuscate (♀); aculeus yellowish basally and reddish brown distally ( Fig. 3 ).

Chelicerae: Manus dorsal surface smooth, with one macroseta located medially on anterior flat plate of manus, and two microsetae on prolateral surface ( Fig. 4A ). Movable finger ventral surface with serrula, comprising 17/18 tines in distal half.

Carapace: Length equal or slightly greater than posterior width (1.1), carapace length slightly greater than metasomal segment V on males and females (1.0), and longer than pedipalp femur (1.2). Surfaces shagreen, uniformly granular. Anterior margin bilobate, with deep median notch and three pairs of macrosetae ( Fig. 5A ). Lateral ocelli conform type 3A, PDMi, PLMa and MLMa subequal in size. Median ocular tubercle shallow, situated in anterior half of carapace, superciliary carinae costate-smooth. Median ocelli approximately three times the size of anterolateral ocelli. Anteriomedian and posteromedian sulcus deep and narrow, posterolateral sulcus shallow, and posterior transverse sulcus vestigial ( Fig. 5A ).

Coxosternal region: Sternum subequilateral pentagonal, anterior width slightly greater than length (1.4). Median sulcus deep, ventral surfaces smooth with four pairs of macrosetae ( Fig. 6A ). Coxae ventral surfaces entirely smooth. Coxa II, prolateral proximal margin smooth, subproximally with two oblique, minute, slit-like structures, adjacent to distinct, granular protuberance. Coxa IV length twice the length of coxa II (1.9/2.0).

Pedipalps: Femur length three times greater than width (3.0/3.2) [Appendix A]; intercarinal surfaces matte; dorsal prolateral, dorsal retrolateral and ventral prolateral carinae complete granular ( Fig. 7 ); ventral median carinae vestigial, strongly granular; retrolateral ventral carinae vestigial, weakly granular ( Fig. 7B ); ventral retrosubmedian carinae vestigial, reduced to few proximal granules; retrolateral dorsosubmedian carina partial, restricted to median part of segment, comprising six or seven serrated granules and two macrosetae; prolateral ventral carina vestigial, reduced to three conical granules, proximal and median granules with one macroseta, extending two-thirds the length of the segment ( Fig. 7D ); prolateral ventrosubmedian carina vestigial, reduced to three to four enlarged conical granules, proximal and two distal granules, each with one macroseta ( Fig. 7 ). Patella length three times greater than the width (3.0/2.9) and as wide as femur (1.1) [Appendix A]; intercarinal surfaces matte; dorsal prolateral, dorsal retrolateral, ventral prolateral, and ventral retrosubmedian carinae complete, granular ( Fig. 8); ventral median carina partial, granular, restricted to proximal third; prolateral ventral carina obsolete, delimited by two macrosetae medially; prolateral process well developed, prolateral median carina delimited by two large granules, each with a macroseta, and a row of five coarse granules curving toward dorsal prolateral carina ( Fig. 8A, B, D); retrolateral median carinae partial, irregular broken row of granules; retrolateral dorsosubmedian vestigial, with three granules in median third. Chela length (1.6/1.7) times greater than patella length, (1.7/1.8) times greater than femur length; width (1.1/1.2) times greater than patella width, (1.3/1.4) times greater than femur width (Appendix A). Manus incrassate ( Fig. 9 ); all intercarinal surfaces matte, finely and sparsely granular, comparatively smaller granules than femur and patella; dorsal retrolateral, dorsal median and retrolateral median carinae complete, costate, with clusters of weak granules ( Fig. 9A ); prolateral dorsal, dorsal prosubmedian, and dorsal prolateral carinae fused, strongly granular; dorsal retrolateral accessory carina vestigial, restricted to three or four proximal granules; retrolateral ventral carina vestigial, short row of proximal granules; retrolateral subventral accessory carina vestigial with short cluster of distal granules ( Fig. 9B ); ventral retrolateral carina granular, incomplete restricted to the level of trichobothrium V2 ( Fig. 9C ); ventral retrosubmedian carina partial, irregular cluster of granules restricted to median third, retrolateral subventral carina vestigial, distal short row of granules; ventral prolateral and prolateral ventral carinae complete, fused, with a cluster of granules; prolateral median and prolateral ventrosubmedian carinae partial, fused, with a cluster of enlarged granules restricted to median third; other carinae obsolete ( Fig. 9 ). Fixed and movable fingers dentate margin sublinear, notches and lobes absent; fixed finger median denticle row comprising six denticle subrows flanked by six prolateral and retrolateral denticles, retrolateral denticles aligned with subrows ( Fig. 10A ); movable finger denticle row comprising six denticle subrows, flanked by seven prolateral and six retrolateral denticles, retrolateral denticles aligned with medial rows except distal two slightly unaligned, terminal subrow comprising two denticles ( Fig. 10B ).

Trichobothrial pattern orthobothriotaxic Type C; chela trichobothrium Db situated on dorsal retrolateral carina, in proximal fifth of manus; Dt situated in proximal half of manus above dorsal retrolateral carina ( Fig. 9A ); ib and it situated on the base of fixed finger ( Fig. 9D ).

Legs: Basitarsi, prolateral ventral spinule rows on legs I-II complete, extending the whole length of the leg, incomplete on III, absent on IV; retrolateral ventral spinule row on legs I-II complete, incomplete on leg III irregular broken subrows of spinules restricted to distal half, absent on IV; retrolateral dorsal spinule row on legs I-III incomplete, vestigial sparse subrows of spinules restricted to distal half, absent or restricted to up to three spinules on distal third on IV; retrolateral median spinule row vestigial with few distal spinules on I-II, absent on III-IV ( Fig. 11A ). Telotarsi, macrosetal counts on I-IV, respectively: prolateral dorsal 4–6/ 4-6/4–7/4-7, prolateral median 1–3/2-3/1–4/2-4, retrolateral median 1–3/1-2/1–3/1-4, prolateral ventral 1–3/2-4/2–4/2-5, retrolateral ventral 3–5/4-6/4–6/3-5; proximal three macrosetae spiniform on II-IV; dorsal and retrolateral dorsal macrosetae arranged in two separate parallel to subparallel rows on I-IV. Telotarsi I-III, each with a single straight ventromedial row of spinules, curved proximally except on IV straight, and one to two pairs of ventrodistal spinules ( Fig. 11A ).

Genital operculum: Genital operculum (2.8/2.6) times wider than long with three pairs of macrosetae and variable accessory minor macrosetae ( Fig. 6A ); sclerites free longitudinally on anterior third, fused distally (6) or fused longitudinally (♀); genital papillae present, protruding posteriorly (6) or absent (♀).

Hemispermatophore: Laminar measurements (mm), lamina length slightly greater than stem (1.1); Blt_AL, 2.6; Clt_AL, 2.3; and Lcdc, 0.5. Laminar antero-distal process prominent; latero-distal crest prominent, located in the distal fifth of the lamina ( Fig. 2A ); laminar hooks moderately bifurcated. Capsular distal carina prominent and spiniform; basal lateral trough shallow; capsular basal carina with circa 13 hooklets; clasper prominent, spiniform ( Figs. 1C and 2A ). Stem axial carina prominent, parallel to posterior margin ( Fig. 2A ).

Pectines: Basal piece with two pairs (6) or three pairs of macrosetae (♀). Marginal lamella comprising three sclerites ( Fig. 6A ). Medial lamella proximal two or three sclerites fused, seven (♀) or eight (6) separate. Fulcra, 13/13 (♀) or 14/14 (6). Pectinal tooth count: 15–16 (6), 13–14 (♀). Pectines relatively long, third (distal) sclerite of marginal lamella aligned with midpoint of trochanter IV (6) or third sclerite of marginal lamella aligned with distal margin of coxa IV (♀).

Tergites: Tergites I-VII intercarinal surfaces pretergites and margin of anterior postergites matte, postergite shagreened, dorsal median and dorsal lateral carina obsolete on I and II, vestigial on III-IV, dorsal sublateral carinae vestigial, comprising few granules anteriorly, dorsal lateral and lateral median carinae converging anteromedially, with a spiniform terminal granule on VII.

Sternites: Sternites III-VI surfaces smooth, spiracles minute, slit-like, three times longer than wide; VII intercarinal median surfaces smooth, matte laterally, six pairs of setae including one pair on ventral margin, ventral submedian carina obsolete, ventral lateral carina costate-granular with two pairs of setae, lateral ventral carinae vestigial with few anterior granules ( Fig. 12A ).

Metasoma: Length (1.6) times greater than mesosoma length. Segment I–V length (0.7/0.6), (0.8), (0.9/1), (1.4/1.5), (2.4/2.5) times greater than width respectively (Appendix A); intercarinal surfaces mostly smooth but with matte surfaces; dorsal lateral carinae complete, costate-granular terminating in enlarged spiniform granules posteriorly on I-IV, cluster of dense granulation on V; lateral median carinae complete, costate-granular, terminating in enlarged spiniform granules posteriorly on I-III, lobate posteriorly on IV, partial, with granulation in anterior half on V; lateral inframedian carinae complete, granular on I, partial, restricted to posterior third, granular on II, vestigial, restricted to few posterior granules, absent on IV-V; ventral lateral carinae complete, granular on I–V; ventral submedian carinae complete, costate, weakly crenular on I, complete granular on II-IV; ventral median carina complete, granular on V ( Fig. 13A–C ). Macrosetal counts on carinae of segments I–V, respectively: dorsal lateral 0-0:0-1:1-1:2-2:3-4; lateral median, 0-0:1-2:1-2:3-4:3-4; lateral inframedian, 1-2:0-1:0-1:1-1:0-0; ventral lateral, 2-2:2-3:2-4:3-4:5-7; ventral submedian, 3-3:3-3:3-3:5- 5:0-0; exclusive for segment V, ventral sublateral 2–3, ventral median 3–4.

Telson: Vesicle elongated; length (1.7/1.8) greater than width, (1.4/ 1.5) times greater than aculeus length (Appendix A). Dorsal surface smooth, without hyaline glandular area, flat ( Fig. 14A ). Ventral surface carinae weakly and finely granular, each with three pairs of macrosetae; subaculear tubercle obsolete. Aculeus laterobasal microserration absent.

Distribution: Vaejovis aguazarca sp. nov. is known from several localities within the Sierra Fría in Aguascalientes and its distribution probably extends to the southern part of Zacatecas ( Fig. 22 ).

Natural History: Specimens were collected under the bark of fallen logs or the bark of live trees, especially oaks ( Quercus eduardi Trel ), above ca. 0.4 m from the base; specimens have also been collected from the bark of live Yucca filifera Chaband. Although unfrequently, scorpions were also found under rocks and leaf litter. Specimens were collected during the whole year, but we noticed a peak of abundance during the rainy season, from July to September. Most scorpions were collected in humid microhabitats, such as the bottom of canyons or protected slopes. Three females kept in the lab at Estacion´Biologica´Agua Zarca during July 2015 gave birth to 27, 34, and 41 scorpionlings respectively, which took a random orientation on the back of the mother. The number of broods contrasts sharply with the numbers reported for V. tenamaztlei : 14, 18, and 22 (Contreras-F´elix et al., 2015), probably due to the larger body size of V. aguazarca sp. nov. Although it is possible that V. aguazarca sp. nov. and V. tenamaztlei might be found in sympatry, there appears to be a preference in habitat. Vaejovis aguazarca sp. nov. has been collected between 1927 and 2201 m of altitude, whereas V. tenamaztlei at 2390–2864 m. This difference of 100 m suggests that each species occupy different altitudinal levels and ecological niche, but further fieldwork is necessary to assess this observation.

Vaejovis aquascalentensis Ch´avez-Samayoa and Gonz´alez-Santill´an sp. nov.

LSID: urn:lsid:zoobank.org:act:016446C8-A814-413C-A6D9-F12C0BE187F7 .

Fig. 1A, B , 2B , 4B , 5B , 6B , 10C, D , 11B , 12B , 13 D-F, 14B, 15A-D, 16–18, 22, 23; Tables 1–3; Appendix A.

Pseudouroctonus sp. Escoto-Rocha and Delgado-Saldívar, 2008: 126–127, Table 3.12.2.

TYPE Material: M´exico: Aguascalientes: Municipio Calvillo: Holotype 1 6 ( CNAN-T 01496 ) Presa Los Alamitos , 21.730515 ◦ N 102.714031 ◦ W, 2367 m, 24.v.2021, F. Ch´avez-Samayoa, M. E. Samayoa-Sepúlveda. Paratype 2 6, 1 ♀ ( CNAN-T01497 ), 21.730515 ◦ N 102.714031 ◦ W, 2367 m, 17.v.2021, F. Ch´avez-Samayoa and D. Ortíz-Alvarez ´; 3 6 (CNAN- T01498 ) Presa el Adobe, inside a narrow canyon, 21.80559 ◦ N 102.68222 ◦ W, 2026 m, 3.viii.2017, J. E. Díaz-Plascencia and E. Gonz´alez-Santill´an .

Additional MATERIAL EXAMINED: Municipio Calvillo: 1 ♀ ( CZUAA SCO-397 ), Jaltiche de Arriba , 18.iv.2010, F. Rodríguez ; 1 ♀ ( CZUAA SCO-398 ), Los Alisos , 3.i.1983, C. Martínez-Saldana ˜; 3 ♀ ( CZUAA SCO-399 ), Cerro de la Manteca , Los Adobes, 1.ix.2015, J. E. Díaz-Plascencia; 8 6 ( CZUA SCO-610 ) Presa Los Alamitos, 21.730515 ◦ N 102.714031 ◦ W, 2367 m, 17.v.2021, F. Chavez-Samayoa ´and D. OrtízAlvarez; 1 6 ( CZUAA SCO-611 ), 21.730515 ◦ N 102.714031 ◦ W, 2367 m, 24.v.2021, F. Ch´avez-Samayoa .

Etymology: The specific epithet refers to the state of Aguascalientes and is used as a noun in apposition.

DIAGNOSIS: Vaejovis aquascalentensis sp. nov. is readily distinguished from V. aguazarca sp. nov. and V. tenamaztlei by the presence of two distal macrosetae on the chelicerae manus dorsal surface instead of one microseta and one macroseta ( Fig. 4B ). Pedipalp patella prolateral median carina with a short row of granules instead of a longer row of granules, or obsolete, delimited by two large granules ( Figs. 8C, D, 17C, D , 20C, D ). Pedipalp chela manus prolateral dorsal, dorsal prosubmedian, and dorsal prolateral carinae with low granules instead of finely granular or obsolete ( Figs. 9A , 18A and 21A ).

Furthermore, V. aquascalentensis sp. nov. differs from V. aguazarca sp. nov. by the pedipalp femur retrolateral dorsosubmedian carinae costate in males and weakly granular in females instead of serrated granules ( Figs. 7B, C , 16B, C ). Pedipalp patella dorsal retrolateral carinae costate and weakly granular instead of complete and granular. Pedipalp chela manus intercarinal surfaces smooth instead of matte, finely and sparsely granular ( Figs. 9 and 18 ). Movable finger with seven prolateral and retrolateral denticles instead of seven prolateral and six retrolateral denticles ( Fig. 10A, C ).

Vaejovis aquascalentensis sp. nov. can also be distinguished from V. tenamaztlei by having six pairs of setae on the sternite VII instead of five ( Fig. 12B, C ). Metasomal ventral lateral carina granular on segments II-V instead of almost smooth with low granules ( Fig. 13F, I ).

Hemispermatophore laminar antero-distal process obsolete in V. aquascalentensis sp. nov ( Fig. 2B, C ). instead of prominent as in V. aguazarca sp. nov. or vestigial as in V. tenamaztlei ; basal lateral trough deep instead of shallow ( Figs. 1B and 2B ). Laminar antero-basal constriction conspicuous, instead of absent as in V. aguazarca sp. nov. or vestigial as in V. tenamaztlei . Capsular basal carina smooth and straight without hooklets in V. aquascalentensis sp. nov., instead of having ca. 13 hooklets as in V. aguazarca sp. nov., or V-shaped as in V. tenamaztlei ( Fig. 2 ); clasper obsolete, instead of spiniform in as in V. aguazarca sp. nov ( Fig. 1 ); capsular distal carina straight at the level of the laminar anterior margin, instead of strongly protruding and spiniform as in V. aguazarca sp. nov.

Vaejovis aquascalentensis sp. nov. resembles Vaejovis coalcoman Contreras-F´elix and Francke 2014 from Michoac´an state in the robust chela manus and similar pectinal count 14–15 (6), 12–13 (♀), but it can be differentiated by the following: Pedipalp patella retrolateral median carinae obsolete, retrolateral dorsosubmedian carina costate in females and weaker in males instead of granular in both sexes. Pedipalp chela fixed finger with six retrolateral denticles instead of five; movable finger with seven prolateral and retrolateral denticles instead of five retrolateral and six prolateral denticles. Mesosomal pretergites surface smooth and postergites matte in V. aquascalentensis sp. nov. but shagreened in V. coalcoman ; sternite VII lateral carinae costate smooth, instead of sparsely granulose. Metasomal ventral submedian carinae costate on segments I and II, and weakly granular on III instead of granular on segments I-III; dorsolateral carinae costate granular on I-IV, instead of finely granular.

Description: The following description is based on the type series, including adults of both sexes and additional material examined.

Color and infuscation: Body base color beige (in alcohol may turn brownish). Carapace, tergites infuscate with dusky markings ( Fig. 15 ). Cheliceral margin of the dorsal surface of manus feebly infuscate; manus immaculate, dorsal surface of movable finger infuscate. All pedipalp chela carinae infuscate as well as all metasomal carinae ( Fig. 15 ). Lateral and dorsal surfaces of metasomal segments III-V diffusely infuscate. Coxosternal region and sternites III-VII immaculate ( Fig. 15B, D ). Legs finely infuscate on dorsal surface. Telson yellowish (may turn brownish in alcohol), with feeble infuscation at the base of subaculear tubercle; aculeus brownish at the base, becomes reddish towards the apex ( Fig. 15 ).

Chelicerae: Manus dorsal surface smooth, with two macrosetae located medially on an anterior flat plate of the manus ( Fig. 4B ). Movable finger ventral surface with serrula, comprising 14 tines in distal half.

Carapace: Length equal or slightly greater than posterior width (1.1/ 1), carapace length slightly shorter than metasomal segment V on males and females (0.8/0.9), and longer than the pedipalp femur (1.2/1.2). Surfaces uniformly granular except for a heart-shaped, matte area surrounding the ocular tubercle ( Fig. 5B ). Anterior margin slightly curved, with subtle median notch and three pairs of macrosetae. Lateral ocelli type 3A, PDMi half the size of PLMa, and PLMa half the size of MLMa. Median ocular tubercle shallow, situated in anterior half of carapace, superciliary carinae costate-smooth. Median ocelli approximately three times the size of anterolateral ocelli. Anteromedian and posteromedian sulci deep and narrow, posterolateral sulcus shallow and posterior transverse sulcus vestigial ( Fig. 5B ).

Coxosternal region: Sternum subequilateral pentagonal, anterior width (1.5/1.38) times greater than length (Appendix A). Median sulcus wide, surfaces smooth, with four pairs of macrosetae ( Fig. 6B ). Coxae ventral surfaces entirely smooth. Coxa II, prolateral proximal margin smooth, subproximally with two oblique, minute, slit-like structures adjacent to a distinct smooth protuberance. Coxa IV length (1.74/1.9) times greater than coxa II length (Appendix A).

Pedipalps: Femur length (2.8) times greater than width (Appendix A); intercarinal surfaces matte ( Fig. 16 ); dorsal prolateral, dorsal retrolateral and ventral prolateral carinae complete and granular; ventral median carina granular; retrolateral ventral carina vestigial, with few proximal granules; ventral retrosubmedian carina vestigial, reduced to few proximal granules; retrolateral dorsosubmedian carina costate in males and costate and weakly granular in females, restricted to median part of segment, with two macrosetae ( Fig. 16A, B ); prolateral ventral carina vestigial, reduced to three conical granules with one macroseta each, extending two-thirds the length of the segment; prolateral ventrosubmedian carina vestigial, reduced to three to four enlarged conical granules, proximal and two distal granules each with one macroseta ( Fig. 16C ).

Patella width (1.1) times femur width (Appendix A); intercarinal surfaces matte ( Fig. 17 ); dorsal prolateral and ventral retrosubmedian carinae complete, coarsely granular; ventral prolateral carina complete and granular; dorsal retrolateral carina costate, weakly granular, almost entirely smooth; ventral median carina incomplete, restricted to proximal third, granular; prolateral ventral carina obsolete, delimited by two median macrosetae ( Fig. 17C, D ); prolateral process well developed ( Fig. 17A, C ); prolateral median carina obsolete, with one macroseta; retrolateral median carina obsolete; retrolateral dorsosubmedian carina partial, costate in females and weaker in males.

Chela length (1.8) times greater than patella length, (2.0/1.9) times greater than femur length; width (1.5) times greater than patella width, (1.6/1.7) times greater than femur width (Appendix A). Manus incrassate ( Fig. 18 ); all intercarinal surfaces smooth; dorsal retrolateral, dorsal median, and retrolateral median carinae complete, costate, almost completely smooth ( Fig. 18A, B ); prolateral dorsal, dorsal prosubmedian, and dorsal prolateral carinae fused, with a cluster of low granules of varying sizes; dorsal retrolateral accessory carina costate; retrolateral ventral and retrolateral subventral accessory carinae obsolete; ventral retrolateral carina granular, incomplete, restricted to the level of trichobothrium V2 ( Fig. 18C ); ventral retrosubmedian carina partial, with irregular cluster of granules, restricted to median third in females and obsolete in males; retrolateral subventral carina obsolete; ventral prolateral and prolateral ventral carinae complete, fused, with a cluster of granules in females, restricted distally in males; prolateral median and prolateral ventrosubmedian carinae partial, fused, with a cluster of enlarged granules restricted to median third; other carinae obsolete ( Fig. 18 ). Fixed and movable fingers dentate margin sublinear, notches and lobes absent; fixed finger median denticle row comprising six denticle subrows flanked by six prolateral and retrolateral denticles, retrolateral denticles aligned with subrows ( Fig. 10D ); movable finger median denticle row comprising seven denticle subrows, flanked by seven prolateral and retrolateral denticles, retrolateral denticles aligned with median rows except distal two slightly removed, terminal subrow comprising one or two denticles ( Fig. 10C ).

Trichobothrial pattern orthobothriotaxic, Type C; chela trichobothrium Db situated on dorsal retrolateral carina, in proximal fifth of manus; Dt situated in proximal half of manus above dorsal retrolateral carina ( Fig. 18A ); ib and it situated on the base of the fixed finger ( Fig. 18D ).

Legs: Basitarsi, prolateral ventral spinule rows on legs I-II complete, incomplete on III, absent on IV; retrolateral ventral spinule rows on legs I-II complete, incomplete on leg III irregular broken subrows of spinules restricted to distal half, absent on IV; retrolateral dorsal spinule rows on legs I-III incomplete, vestigial sparse subrows of spinules restricted to distal half, absent or restricted to up to three spinules on distal third on IV; retrolateral median spinule row absent on I-IV. Telotarsi, macrosetal counts on I-IV, respectively: prolateral dorsal 4–7/4-6/4–6/4-8, prolateral median 1–3/2-4/2–5/3-6, retrolateral median 2–3/2-3/2–5/2-6, prolateral ventral 3–4/4-6/4–7/4-7, retrolateral ventral 2–4/3-5/3–8/ 4-7 ( Table 2); proximal three macrosetae spiniform on II-IV; dorsal and retrolateral dorsal macrosetae arranged in two separate parallel to subparallel rows on I-IV. Telotarsi I-III, each with a single straight ventromedial row of spinules, curved proximally, except straight on IV, and one or two pairs of ventrodistal spinules ( Fig. 11B ).

Genital operculum: Genital operculum wider than it is long (0.4/0.3,

Fig. 6B ), with three pairs of macrosetae and variable accessory minor macrosetae; sclerites free longitudinally on anterior third, fused distally (6) or fused longitudinally (♀); genital papillae present, protruding posteriorly (6) or absent (♀).

Hemispermatophore: Laminar measurements (mm), lamina length slightly greater than stem (1.1); Blt_AL, 2.9; Clt_AL, 2.5; and Lcdc, 0.7. Latero-distal crest prominent, located in the distal fifth of lamina; laminar antero-distal process absent; laminar hooks strongly bifurcated ( Fig. 2B ). Capsular distal carina prominent but blunt; basal lateral trough deep ( Figs. 1B and 2B ); capsular basal carina smooth, straight. Stem with axial carina parallel to posterior margin ( Fig. 2B ).

Pectines: Basal piece with three (6) or four (♀) pairs of macrosetae. Marginal lamella comprising three sclerites. Medial lamella proximal with two or three sclerites fused, seven (♀) or eight (6) separate. Fulcra, 12/12 (♀) or 14/14 (6). Pectinal teeth, 14–15 (6), 12–13 (♀). Pectines relatively long ( Fig. 6B ), third (distal) sclerite of marginal lamella aligned with midpoint of trochanter IV (6) or third sclerite of marginal lamella aligned with distal margin of coxa IV (♀).

Tergites: Tergites I-VII intercarinal surfaces pretergites and margin of anterior postergite smooth (♀) or matte (6), posterior postergite matte; dorsal median and dorsal lateral carina obsolete on I-VI; dorsal sublateral carina vestigial, comprising few granules anteriorly (6) or obsolete (♀), dorsal lateral and lateral median carinae separated, not converging anteromedially and granular on VII ( Fig. 15A, C ).

Sternites: Sternites III-VI surfaces smooth, spiracles minute, slit-like ( Fig. 15B, D ), two times longer than wide; sternite VII intercarinal median surfaces smooth, matte laterally, with hyaline glandular area medially, six pairs of setae, including two pairs on posterior margin, ventral submedian carina obsolete, ventral lateral carina costate with two pairs of setae, lateral ventral carina obsolete ( Fig. 12B ).

Metasoma: Length (1.77/1.51) times greater than mesosoma length (Appendix A). Segments I–V length (0.71/0.63), (0.88/0.78), (1.18/ 0.86), (1.39/1.19), (2.91/2.49) times greater than width, respectively (Appendix A); intercarinal surfaces matte; dorsal lateral carina complete, costate-granular, terminating in enlarged spiniform granules posteriorly on I-IV, cluster of dense granulation on V; lateral supramedian carina complete, low granules, terminating in enlarged spiniform granules posteriorly on I-III, lobate posteriorly on IV, partial, with granulation in anterior half on V; lateral inframedian carina complete, granular on I, partial, restricted to posterior third, granular on II, vestigial, restricted to few posterior granules on III, absent on IV-V; ventral lateral carina complete, low granules on I, granular on II-V; ventral submedian carina complete, costate to finely granular on I-II, finely granular on III, granular on IV; ventral median carina complete, granular on V ( Fig. 13D–F ). Macrosetal counts on carinae of segments I–V: dorsal lateral 0-0:0-1:1-1:1-2:3-4; lateral median 0-0:1-2:1-2:2-3:2- 3; lateral inframedian 1-3:0-2:0-2:1-1:0-0; ventral lateral 1-2:2-2:2-3:2- 4:4-5; ventral submedian 2-3:3-3:3-3:3-4:0-0; exclusive for segment V, ventral sublateral 2–3, ventral median 1–3.

Telson: Vesicle elongate; length (1.75/1.42) greater than width, (2.09/1.89) times greater than aculeus length (Appendix A). Dorsal surface smooth, flat, with an annular hyaline thickening near the base of the aculeus. Ventral surface carina vestigial, each with three pairs of macrosetae; subaculear tubercle obsolete ( Fig. 14B ). Aculeus laterobasal microserration absent.

DISTRIBUTION: Vaejovis aquascalentensis sp. nov. is known from several localities within the Sierra del Laurel in Aguascalientes, and its distribution probably extends to the southern part of Zacatecas and northern part of Jalisco ( Fig. 22 ).

NATURAL HISTORY: Specimens were collected on the side of a hiking road in an oak forest in the Los Alamitos dam, and in a protected forested canyon with subtropical vegetation in the Los Adobes dam. The altitude at which specimens were collected varied from 1601 to 2367 m. Unlike V. aguazarca sp. nov. (1927–2201 m) and V. tenamaztlei (2390–2864 m), Vaejovis aquascalentensis sp. nov. exhibits a wider altitudinal gradient. However, all records of it have been registered within the Sierra del Laurel. It has been hypothesized that the Sierra Fría and the Sierra del Laurel have very different orographic origins, despite being separated by less than 20 km. Additionally, it has also been hypothesized that, Francke (2019). Additionally, the main diversity of Pseudouroctonus is restricted to the isolated mountain ranges of Northern Mexico, in Chihuahua, Coahuila, Nuevo Leon´, Sonora, and most likely Tamaulipas, as well as the southern states of the USA ( Bryson et al. 2013). Aguascalientes is far south of the distribution of Pseudouroctonus , so it is unlikely that this genus would reach the state.

historically, the deep basin created by the Calvillo, Labor, and Malpaso Rivers in the southwestern extreme of the state may have served as a barrier to the dispersion of fauna between the two sierras ( Bryson et al. 2008). This idea has been suggested to propose the possibility that V. tenamaztlei may be restricted to the Sierra del Laurel. However, we report here that V. tenamaztlei is distributed beyond the Sierra del Laurel, having been registered in the Sierra Fría and the Sierra de Juan el Grande as well. On the other hand, our geographical records of V. aquascalentensis sp. nov. suggest that this species is in fact restricted to the Sierra del Laurel.

REMARKS: This species was previously misidentified and reported as Pseudouroctonus sp. ( Escoto Rocha and Delgado Saldivar 2008). The genus Pseudouroctonus comprises small but robust scorpions with incrassate and granular pedipalp carinae, short and granular carinated metasomal segments, and a telson with a constriction at the base of the aculeus ( Gertsch and Soleglad 1972). All these characters can be observed in V. aquascalentensis sp. nov.; however, the absence of a sclerotized mating plug, the laminar hooks contiguous to the capsule, and the presence of infuscation patterns on dorsal surfaces of the body in V. aquascalentensis sp. nov. permitted us to separate these two genera. Furthermore, we confirm that this species complies with the diagnostic characters of the mexicanus group as defined by Contreras-F´elix and

Figures

Fig. 1. Sinistral hemispermatophore of Vaejovis aquascalentensis Ch´avez-Samayoa and Gonz´alez- Santill´an sp. nov. (Upper plate) and Vaejovis aguazarca Díaz-Plascencia and Gonz´alez-Santill´an sp. nov. (Lower plate), cleared of soft tissue by hand. A., C. Contralateral aspect; B., D. Lateral aspect. Abbreviations on all gray squares refer to structures present in the whole Order Scorpiones: Blt, Basal lateral trough. Cbc, Capsular basal carina. Cbch, Capsular basal carina hooklets. Cl, Clasper. Cdc, Capsular distal carina. Clt, Contralateral trough. Lam, Laminar anterior margin. Lh, Laminar hook. Lpm, Laminar posterior margin. Sd, Sperm duct. Sub, Subex. Tec, Tectum. Tf, Truncal flexure.

Fig. 2. Sinistral hemispermatophore, cleared of soft tissue by hand. Contralateral aspect (left) and lateral aspect (right). A. Vaejovis aguazarca Díaz-Plascencia and Gonz´alez-Santill´an sp. nov. B. Vaejovis aquascalentensis Ch´avez-Samayoa and Gonz´alez-Santill´an sp. nov. C. Vaejovis tenamaztlei Contreras-F´elix, Francke and Bryson, 2015. The tridimensional boxes are enclosing the total extension of the capsular distal carina, including the basal and distal sections delimited by the presence of the laminar hook. Abbreviations: Ac, Axial carina, Cdc, Capsular distal carina. Lac, Laminar antero-basal constriction, Lap, Laminar antero-distal process. Llc, laminar latero-distal crest.

Fig. 3. Vaejovis aguazarca Díaz-Plascencia and Gonz´alez-Santill´an sp. nov., habitus. A, B 6, C, D. ♀ (CNAN-S4005). A, C. Dorsal aspect. B, D. Ventral aspect. Scale bar =5 mm.

Fig. 4. Chelicerae dorsal and ventral aspects under UV light (6). A. Vaejovis aguazarca Díaz-Plascencia and´Gonzalez-Santillan´sp. nov. B. Vaejovis aquascalentensis Ch´avez-Samayoa and Gonz´alez-Santill´an sp. nov. C. Vaejovis tenamaztlei Contreras-F´elix, Francke and Bryson, 2015.

Fig. 5. Carapace dorsal surface under UV light (6). A. Vaejovis aguazarca Díaz-Plascencia and Gonz´alez-Santill´an sp. nov. B. Vaejovis aquascalentensis Ch´avez- Samayoa and Gonz´alez-Santill´an sp. nov. C. Vaejovis tenamaztlei Contreras-F´elix, Francke and Bryson, 2015.

Fig. 6. Coxal region, genital operculum, and pectines under UV light. Left side (6) and right side (♀). A. Vaejovis aguazarca Díaz-Plascencia and Gonz´alez-Santill´an sp. nov. B. Vaejovis aquascalentensis Ch´avez-Samayoa and Gonz´alez-Santill´an sp. nov. C. Vaejovis tenamaztlei Contreras-F´elix, Francke and Bryson, 2015.

Fig. 7. Pedipalp femur of Vaejovis aguazarca Díaz-Plascencia and Gonz´alez- Santill´an sp. nov. under UV light (6). A. Dorsal, B. Retrolateral, C. Ventral, and D. Prolateral aspects.

Fig. 10. Pedipalp chela, fixed (left) and movable (right) fingers under UV light (6). A. Vaejovis aguazarca Díaz-Plascencia and Gonz´alez-Santill´an sp. nov. B. Vaejovis aquascalentensis Ch´avez-Samayoa and Gonz´alez-Santill´an sp. nov. C. Vaejovis tenamaztlei Contreras-F´elix, Francke and Bryson, 2015.

Fig. 11. Leg, basitarsal and telotarsal armature under UV light (6). Dorsal (left) and Ventral (right) aspects. A. Vaejovis aguazarca Díaz-Plascencia and Gonz´alez- Santill´an sp. nov. B. Vaejovis aquascalentensis Ch´avez-Samayoa and Gonz´alez-Santill´an sp. nov. C. Vaejovis tenamaztlei Contreras-F´elix, Francke and Bryson, 2015.

Fig. 12. Sternite VII ventral surface ornamentation under UV light (6). A. Vaejovis aguazarca Díaz-Plascencia and Gonz´alez-Santill´an sp. nov. B. Vaejovis aquascalentensis Ch´avez-Samayoa and Gonz´alez-Santill´an sp. nov. C. Vaejovis tenamaztlei Contreras-F´elix, Francke and Bryson, 2015.

Fig. 13. Metasomal segments I-IV under UV light (6). A., D., G. Dorsal, B., E., H. Lateral, and C., F., I. Ventral aspects. A-C. Vaejovis aguazarca Díaz-Plascencia and Gonz´alez-Santill´an sp. nov. D-F. Vaejovis aquascalentensis Ch´avez-Samayoa and Gonz´alez-Santill´an sp. nov. G-I. Vaejovis tenamaztlei Contreras-F´elix, Francke and Bryson, 2015.

Fig. 16. Pedipalp femur of Vaejovis aquascalentensis Ch´avez-Samayoa and Gonz´alez-Santill´an sp. nov. under UV light (6). A. Dorsal, B. Retrolateral, C. Ventral, and D. Prolateral aspects.

Fig. 17. Pedipalp patella of Vaejovis aquascalentensis Ch´avez-Samayoa and Gonz´alez-Santill´an sp. nov. under UV light (6). A. Dorsal, B. Retrolateral, C. Ventral, and D. Prolateral aspects.

Fig. 9. Pedipalp chela of Vaejovis aguazarca Díaz-Plascencia and Gonz´alez- Santill´an sp. nov. under UV light (6). A. Dorsal, B. Retrolateral, C. Ventral, and D. Prolateral aspects.

Fig. 18. Pedipalp chela of Vaejovis aquascalentensis Ch´avez-Samayoa and Gonz´alez-Santill´an sp. nov. under UV light (6). A. Dorsal, B. Retrolateral, C. Ventral, and D. Prolateral aspects.

Fig. 22. Map of Central Mexico and Aguascalientes state showing the distribution of Vaejovis aguazarca Díaz-Plascencia and Gonz´alez-Santill´an sp. nov. (gray circles), Vaejovis aquascalentensis Ch´avez-Samayoa and Gonz´alez-Santill´an sp. nov. (red diamonds), Vaejovis tenamaztlei Contreras-F´elix, Francke and Bryson, 2015 (green stars), Vaejovis coalcoman Contreras-F´elix and Francke 2014 (orange square), and Vaejovis dugesi Pocock, 1902 (blue triangle). Mountain ranges are indicated as: Sierra Fria (1), Sierra del Laurel (2) and Juan el Grande foothills (3). (For interpretation of the references to colour in this figure legend, the reader is referred to the Web version of this article.)

Fig. 14. Telson dorsal (left) and lateral (right) aspects under UV light (6). A. Vaejovis aguazarca Díaz-Plascencia and Gonz´alez-Santill´an sp. nov. B. Vaejovis aquascalentensis Ch´avez-Samayoa and Gonz´alez-Santill´an sp. nov. C. Vaejovis tenamaztlei Contreras-F´elix, Francke and Bryson, 2015.

Fig. 20. Pedipalp patella of Vaejovis tenamaztlei Contreras-F´elix, Francke and Bryson, 2015 under UV light (6). A. Dorsal, B. Retrolateral, C. Ventral, and D. Prolateral aspects.

Fig. 21. Pedipalp chela of Vaejovis tenamaztlei Contreras-Felix´, Francke and Bryson, 2015 under UV light (6). A. Dorsal, B. Retrolateral, C. Ventral, and D. Prolateral aspects.

Fig. 15. Vaejovis aquascalentensis Ch´avez-Samayoa and Gonz´alez-Santill´an sp. nov., habitus. A, B 6, C, D. ♀ (CNAN-S4005). A, C. Dorsal aspect. B, D. Ventral aspect. Scale bar =5 mm.