Systematic Revision Of The North American Syntropine Vaejovid Scorpion Genera Balsateres, Kuarapu, And Thorellius, With Descriptions Of Three New Species Author González-Santillán, Edmundo City University of New York; Scorpion Systematics Research Group, Division of Invertebrate Zoology, American Museum of Natural History; Current address: Instituto de Biotecnología, Universidad Nacional Autónoma de México, Cuernavaca, Morelos, Mexico Facultad de Ciencias, Departamento de Biología Comparada, Universidad Nacional Autonóma de México, Mexico City Author Prendini, Lorenzo Scorpion Systematics Research Group, Division of Invertebrate Zoology, American Museum of Natural History text Bulletin of the American Museum of Natural History 2018 2018-06-20 2018 420 1 1 http://www.bioone.org/doi/10.1206/0003-0090-420.1.1 journal article 10.1206/0003-0090-420.1.1 0003-0090 6499473 A4D48D7A-5088-49F2-B88B-361D392B3F96 Kuarapu purhepecha Francke and Ponce-Saavedra, 2010 Figures 1B , 4 , 5B , 7B, 8B, 9B, 10B, 11B, 12D–F, 14B, 15D, E, 19–21; tables 2, 8 Kuarapu purhepecha Francke and Ponce-Saavedra, 2010: 52 ; Ayrey, 2011: 1 ; González-Santillán and Prendini, 2013: 3 , 6, 8, 11, 18, 21, 24, 28, 33, 43, 45, 51, 52, 56, 59, figs. 4, 18C, D, 24B, table 1; 2015a: 344, 345, 349, 351– 354, 356, 357, 360, 362–364, 367, 373, 377, 380, 384, 388 400, 403, fig. 4–6, tables 5, 6; Ponce-Saavedra and Francke, 2013: 77 , table 2; Quijano-Ravel and Ponce-Saavedra, 2014: 17 , 18, 20, figs. 1–3, table 2; 2016: 50; Dupré, 2017: 11 . TYPE MATERIAL : MEXICO : Michoacán : Municipio de la Huacana : Holotype ♂ ( CNAN T-0411 ), “El Vado” [seasonal drainage underpass], km 17 [road marker], road from Zicuarán to Churumuco , 18.814°N 101.916°W , 248 m , 30.vii.2008 , O. Francke , H. Montaño , J. Ponce , and A. Quijano. Paratypes : 2 ♂ , 2 ♀ ( CNAN T-0412 ), 1 ♂ , 1 ♀ ( AMNH ), same data as holotype ; 1 ♀ ( CNAN T-0413 ), same locality as holotype, 20.v.2007 , O. Francke , J. Ballesteros , J. Ponce , A. Quijano , and M. Villaseñor . DIAGNOSIS: As for genus. DESCRIPTION: The following redescription, which is based on the type material and additional material examined, supplements the original description by Francke and Ponce- Saavedra (2010). Color and infuscation : Chelicerae, carapace, pedipalps, legs, tergites, sternites, metasoma, and telson base color pale yellow to orange. Cheliceral manus, dorsal surface with reticulate infuscation. Carapace and tergites with variegated infuscation. Coxosternal region, genital operculum, basal piece, and pectines pale whitish. Pedipalp femur, patella, and chela dorsal, prolateral, and retrolateral intercarinal surfaces faintly infuscate; carinae densely infuscate; chela fingers infuscate proximally. Legs prolateral surfaces partially infuscate. Sternites mostly immaculate, lateral margins faintly infuscate. Metasoma with variegated infuscation, more densely so on segments III–V. Telson vesicle dorsal surface entirely infuscate; ventral surface with three broad, longitudinal bands of infuscation, one median and two lateral; aculeus reddish to brownish. Chelicerae : Manus dorsal surface smooth, with three macrosetae distally, decreasing in size TABLE 2 Measurements (mm) of adult Kuarapu purhepecha Francke and Ponce-Saavedra, 2010 Specimens deposited in American Museum of Natural History (AMNH) and Colección Nacional de Arácnidos (CNAN), Instituto de Biología, Universidad Nacional Autónoma de México.

Collection	CNAN	AMNH	CNAN	AMNH
Number	T0411	T0412	1234	T0412	T0413
Type	Holo.	Para.	Para.
Sex	♂	♂	♂	♂	♀	♀	♀	♀	♀
Carapace	length	3.2	3.2	3.0	3.0	4.1	4.3	4.0	3.9	3.8
ant. width	1.5	1.5	1.6	1.6	2.2	2.2	2.0	1.9	2.1
post. width	2.7	2.7	2.5	2.5	3.8	4.0	2.5	3.5	3.3
Femur	length	2.6	2.7	2.6	2.5	3.5	3.6	3.3	3.2	3.2
width	0.9	0.8	0.8	0.8	1.2	1.2	1.1	1.1	1.1
Patella	length	3.1	3.1	3.0	2.9	4.0	4.1	3.9	3.8	3.9
width	1.0	0.9	0.9	1.0	1.3	1.4	1.3	1.2	1.2
Chela	length	4.6	4.7	4.1	4.4	5.7	5.8	5.2	5.2	5.5
Manus	width	1.1	1.1	1.0	1.1	1.3	1.3	1.2	1.2	1.1
Manus	height	1.1	1.0	1.0	1.0	1.1	1.2	1.2	1.1	0.9
Manus	length	2.4	2.5	2.2	2.3	2.8	3.0	2.7	2.6	2.8
Fixed finger	length	2.2	2.2	2.0	2.1	2.9	2.8	2.5	2.6	2.8
Mov. finger	length	2.7	2.8	2.5	2.7	3.5	3.6	3.3	3.2	3.5
Coxa II	length	1.3	1.2	1.3	1.2	1.6	1.8	1.6	1.6	1.5
Coxa IV	length	2.5	2.5	2.3	2.1	3.3	3.6	3.1	3.2	3.0
Sternum	length	0.6	0.7	0.6	0.7	0.8	0.9	0.7	0.8	0.9
width	0.7	0.7	0.8	0.7	0.9	0.9	0.9	0.9	1.0
Mesosoma	length	7.1	6.6	6.8	5.7	6.3	10.4	9.5	9.1	6.8
Metasoma	length	10.8	9.4	9.9	9.9	13.1	13.1	11.9	11.9	12.0
Segment I	length	1.4	1.4	1.3	1.3	1.8	1.8	1.7	1.6	1.6
width	1.8	1.7	1.7	1.6	2.4	2.4	2.2	2.2	2.2
height	1.5	1.5	1.3	1.2	2.1	2.0	1.8	1.9	1.5
Segment II	length	1.7	1.6	1.5	1.5	2.1	2.0	1.7	1.9	1.9
width	1.8	1.7	1.6	1.6	2.3	2.3	2.2	2.1	2.1
height	1.4	1.4	1.3	1.2	2.0	1.9	1.7	1.8	1.5
Segment III	length	1.8	1.7	1.7	1.7	2.2	2.2	2.0	1.9	2.0
width	1.7	1.6	1.6	1.0	2.2	2.3	2.2	2.0	2.1
height	1.4	1.4	1.3	1.5	2.0	1.9	1.8	1.8	1.5
Segment IV	length	2.5	2.4	2.3	2.3	3.0	3.0	2.7	2.7	2.7
width	1.7	1.6	1.6	1.0	2.2	2.2	2.1	2.0	2.0
height	1.4	1.4	1.3	1.5	2.0	1.9	1.8	1.8	1.4
Segment V	length	3.6	2.2	3.1	3.3	4.1	4.3	3.9	3.9	3.9
width	1.6	1.6	1.6	1.2	2.1	2.1	2.0	1.9	1.9
height	1.4	1.3	1.3	1.5	1.9	1.8	1.7	1.7	1.4
Telson	length	3.1	3.0	2.7	2.9	3.7	4.0	3.7	3.8	3.6
Vesicle	length	2.1	2.0	1.8	1.9	2.4	2.6	2.3	2.3	2.2
width	1.2	1.2	1.1	0.9	1.5	1.6	1.5	1.5	1.5
height	1.0	1.0	0.9	1.1	1.3	1.4	1.2	1.2	1.1
Aculeus	length	1.1	1.0	0.9	1.1	1.4	1.4	1.4	1.5	1.5
Total	length	24.3	22.2	22.5	21.4	27.2	31.7	29.1	28.6	26.2

from dorsomedian to prolateral surface. Movable finger, ventral surface with serrula, comprising 14/14 tines, in distal half. Carapace : Length 1.18/1.16× greater than posterior width (table 2). Anterior margin sublinear, with vestigial median notch, and three pairs of major macrosetae (fig. 5B). Three pairs of lateral ocelli, anterolateral and median pairs equal in size, posterolateral pair half the size. Median ocular tubercle shallow, situated in anterior half of carapace, superciliary carinae smooth, lower than median ocelli. Median ocelli more than twice the size of anterolateral ocelli. Anteromedian, posteromedian, and posterolateral sulci moderate, posterior transverse sulci obsolete. Surfaces matte ( ♂ ) or shagreened, covered with minute rounded granules ( ♀ ). Coxosternal region : Sternum subequilateral pentagonal (fig. 7B); length 0.89/0.88× longer than wide (table 2); median sulcus deep; surfaces matte, with four pairs of macrosetae. Coxae surfaces matte, smooth in places; coxa II subproximal margin with two oblique slitlike structures, adjacent to small, smooth protuberance; coxal endite II proximal margin with shallow depression, medial margin smooth. Coxa IV 1.92/2.04× longer than coxa II (table 2). Pedipalps : Femur intercarinal surfaces matte (fig. 20A); dorsal prolateral, dorsal retrolateral, and ventral prolateral carinae complete, granular to costate-granular; retrolateral dorsosubmedian carina complete, serrate, with proximal tubercle and two major macrosetae medially; prolateral ventrosubmedian carina partial, reduced to row of conical granules and two minor macrosetae medially; prolateral ventral carina vestigial, reduced to proximal and distal tubercles and three conical granules, one bicuspid, medially, each with macroseta; ventral median and retrolateral ventral carinae vestigial, reduced to scattered serrate granules; ventral retrosubmedian carina obsolete. Patella 1.16/1.14× wider than femur (table 2); intercarinal surfaces matte (fig. 20B–E); dorsal prolateral, dorsal retrolateral, and ventral prolateral carinae complete, finely granular; ventral retrosubmedian carina vestigial, reduced to proximal spiniform granules; ventral median carina vestigial, reduced to few low granules; retrolateral dorsosubmedian and retrolateral median carinae obsolete; prolateral process moderately developed; prolateral median carina complete, comprising tubercle with macroseta proximally, row of serrate granules medially, and tubercle with minor macroseta distally; prolateral ventral carina vestigial, comprising proximal and distal granules, each with major macroseta, and minor macrosetae medially. Chela 1.46/1.40× longer than patella, 1.69/1.62× longer than femur (table 2). Manus elongate and slender (fig. 21), 1.12/0.95× wider than patella, 1.30/1.08× wider than femur; intercarinal surfaces matte; acarinate. Fixed and movable fingers, dentate margins emarginate, fixed finger with shallow proximal notch and median lobe (fig. 21B, D), movable finger with shallow proximal lobe, fitting together unevenly such that moderate proximal gap evident when closed ( ♂ ) or dentate margins sublinear, notches and lobes absent, fitting together evenly such that no gap evident when closed ( ♀ ); fixed finger, median denticle row comprising five denticle subrows flanked by six prolateral and five retrolateral serrate denticles, prolateral and retrolateral denticles equal in size; movable finger, median denticle row comprising five denticle subrows, flanked by seven prolateral denticles and five retrolateral denticles, subpaired, terminal subrow comprising single denticle; terminal retrolateral denticle of fixed and movable fingers considerably larger and sharper than preceding denticles. Trichobothrial pattern type C, orthobothriotaxic; chela trichobothrium Db situated on dorsal retrolateral carina; Dt situated dorsal to dorsal retrolateral carina, near midpoint of manus, but slightly toward distal half; db–dt and eb–et distributed along entire length of fixed finger with db situated proximal and eb subproximal; ib aligned with sixth prolateral denticle (fig. 21D); it situated between fifth and sixth prolateral denticles. Legs : Basitarsi retrolateral dorsal, prolateral ventral, and retrolateral ventral spinule rows complete on I and II, retrolateral dorsal spinule row partial, restricted to distal half of III and IV, prolateral ventral spinule row vestigial, reduced to few distal spinules on III, absent on IV; retrolateral ventral spinule row complete on III, absent on IV; retrolateral median spinule row vestigial, reduced to few distal spinules on I–IV; macrosetal counts on legs I–IV, respectively: dorsal, 3:3:3:3; retrolateral dorsal, 2:3:3:3; prolateral ventral, 3:3:5:5; retrolateral ventral, 3:5:6:6; dorsal and retrolateral dorsal macrosetae arranged in two separate rows on I–IV. Telotarsi I–IV, each with single ventromedian row of spinules, curved proximally, with two pairs of ventrodistal spinules (fig. 11B). Genital operculum : Width greater than length (fig. 7B), with three pairs of macrosetae. Sclerites separated, but unable to open more than 45° ( ♂ ) or fused longitudinally ( ♀ ). Genital papillae present, protruding posteriorly ( ♂ ) or absent ( ♀ ). Hemispermatophore : Lamina 1.76× longer than trunk (table 8). Median lobe, ental terminus moderately developed, rounded (fig. 12D, F). Dorsal trough margin long, narrow, curving proximally, well separated from ventral trough (fig. 12F). Dorsal and ventral trough margins, terminal spinelike processes fused into prominent, bifurcate hook, situated distally on lamina dorsal margin (fig. 12E, F). Basal plate of inner lobe situated proximal to ventral trough (fig. 14B); spine of inner lobe moderately developed. Hemimating plug developed from inner lobe; distal barb 0.6× longer than basal plate; distal barb margin with 9 elongated spinules; secondary spine on ventromedian side of distal barb moderately developed (fig. 15D, E). Pectines : Basal piece with two pairs of minor macrosetae. Marginal lamella comprising three sclerites. Medial lamella proximal sclerites fused, 11–13 distal sclerites separate (fig 5D). Fulcra, 16–17 ( ♂ ), 15–16 ( ♀ ). Pectinal teeth, 17–18 ( ♂ ), 16–17 ( ♀ ). Pectines relatively long, midpoint ( ♂ ) or distal margin ( ♀ ) of medial sclerite of marginal lamella aligned with distal margin of coxa VI. Tergites : Pretergites I–VII, surfaces matte. Posttergites I–VI, intercarinal surfaces matte, except posterolateral surfaces more coarsely granular, dorsal median and dorsal lateral carinae obsolete, granular, restricted to posterior half; VII, intercarinal surfaces matte, except posterolateral surfaces more coarsely granular, dorsal median carina partial, restricted to anterior half, costate, dorsal submedian carinae vestigial, each reduced to anterior tubercle; dorsal lateral and lateral median carinae granular, converging anteromedially, posterior granules larger, spiniform. Sternites : Sternites III–VI, surfaces acarinate, matte; spiracles elongate, slitlike, ca. 2× longer than wide; V , posteromedian hyaline glandular area protruding slightly from adjacent surfaces ( ♂ ) or obsolete ( ♀ ) ; VII, posteromedian hyaline glandular area rounded, protruding moderately, ventral submedian carinae obsolete, each with three macrosetae, ventral lateral carinae weakly costate-granular. Metasoma : Metasoma 1.59/1.48× longer than mesosoma (table 2). Segments I – V , respectively 1.25/1.35, 1.06/1.16, 0.87/1.04, 0.61/0.74, 0.48/0.49× wider than long ; V , 1.03/1.07× wider than telson vesicle. Segments I– V , all intercarinal surfaces matte (figs. 9B, 10B) ; dorsal lateral carinae serrate, terminating in conical spiniform granules posteriorly on I–IV, serrate in anterior half, becoming finely granular posteriorly on V; lateral median carinae serrate, terminating in conical spiniform granules posteriorly on I–III, lobate on IV, partial, restricted to posterior two-thirds, serrate anteriorly, becoming costate posteriorly on V; lateral inframedian carinae complete, serrate on I, partial, obsolete anteriorly, granular in posterior half on II, partial, serrate, restricted to posterior third on III, absent on IV; ventral lateral carinae finely serrate on I–V; ventral submedian carinae costate on I and II, costate in anterior two-thirds, finely serrate in posterior third, on III, finely serrate on IV; ventral median carina finely serrate on V . Macrosetal counts on carinae of segments I– V , respectively: dorsal lateral carinae, 0:0:1:1:4 ( ♂ ), 1:2:3:3:9 ( ♀ ) ; lateral median carinae, 1:1:1:2:2 ( ♂ ), 1:2:3:4:5 ( ♀ ); lateral inframedian carinae, 1:1:1:0:0 ( ♂ ), 3:2:1:0:0 ( ♀ ); ventral lateral carinae, 2:2:2:3:5 ( ♂ ), 3:3:3:4:12 ( ♀ ); ventral submedian carinae, 3:3:3:3 ( ♂ ), 3:3:4:3:0 ( ♀ ); ventral sublateral carinae, 0:0:0:0:3 ( ♂ ), 0:0:0:0:3 ( ♀ ); ventral median carinae, 0:0:0:0:4 ( ♂ ), 0:0:0:0:6 ( ♀ ); segments II– IV each with three accessory macrosetae on ventral median surface ( ♀ ). FIGURE 11. Balsateres González-Santillán and Prendini, 2013 , Kuarapu Francke and Ponce-Saavedra, 2010 , and Thorellius Soleglad and Fet, 2008 , dextral leg III, basitarsus and telotarsus, ventral aspect. A. Balsateres cisnerosi ( Ponce-Saavedra and Sissom, 2004 ) , ♂ (CNAN SC2441). B. Kuarapu purhepecha Francke and Ponce-Saavedra, 2010 , paratype ♂ (AMNH). C. Thorellius cristimanus ( Pocock, 1898 ) , ♂ (AMNH). D. Thorellius intrepidus (Thorell, 1876) , ♂ (AMNH). E. Thorellius tekuani , sp. nov. , holotype ♂ (CNAN T01250). F. Thorellius wixarika , sp. nov. , holotype ♂ (CNAN T01251). G. Thorellius yuyuawi , sp. nov. , holotype ♂ (CNAN T01252). Scale bars = 1 mm. Telson : Vesicle relatively elongate (fig. 10B); length 2.73/2.52× greater than width (table 2). Ventral and dorsal surfaces smooth, relatively hirsute; ventral carinae each with three ( ♂ ) or five to six ( ♀ ) macrosetae posteriorly. Subaculear tubercle vestigial, comprising low, broad tubercle, protruding weakly from adjacent surface. Aculeus, laterobasal microserration comprising seven sharp spinules. DISTRIBUTION: As for genus. ECOLOGY: Kuarapu purhepecha inhabits tropical deciduous forest at an altitude of approximately 250 m . It has been collected with UV light detection at night on the surfaces of road cuts ( Francke and Ponce-Saavedra, 2010 ) and during the day from rock piles at the base of talus slopes covered with vegetation and leaf litter. The habitat and habitus of Kuarapu are consistent with the lithophilous ecomorphotype ( Prendini, 2001 ). Kuarapu purhepecha has been collected in sympatry with Balsateres cisnerosi , Centruroides balsasensis Ponce-Saavedra and Francke, 2004 , Kolotl poncei (Francke and Quijano- Ravell, 2009), Konetontli sp. , Mesomexovis sp. , and Thorellius tekuani at the type locality (J. Ponce-Saavedra and A. Quijano-Ravel, personal commun.). REMARKS: Two adult specimens from El Valle labeled as paratypes were not listed as such in the original description. ADDITIONAL MATERIAL EXAMINED: MEX- ICO: Michoacán : Municipio de La Huacana : El Vado, 18°48′51″N 101°54′57″W , 248 m , 30. vii.2008 , O. Francke, H. Montaño , J. Ponce, and A. Quijano, 2 juv. (AMCC [LP 9517]). Municipio de Parácuaro : El Valle [ca. 5 km S of Parácuaro, 19°06′12.41″N 102°13′09.73″W ], 21.x.2000 , J. Ponce et al., 1 ♂ (AMNH), 1 ♀ (CNAN).