Balsateres, Gonzalez-Santillan and Prendini, 2013

González-Santillán, Edmundo & Prendini, Lorenzo, 2018, Systematic Revision Of The North American Syntropine Vaejovid Scorpion Genera Balsateres, Kuarapu, And Thorellius, With Descriptions Of Three New Species, Bulletin of the American Museum of Natural History 2018 (420), pp. 1-1 : 8-16

Balsateres González- Santillán and Prendini, 2013

Figures 5 , 8 , 9 –15; table 8

Vaejovis cisnerosi Ponce-Saavedra and Sissom, 2004 [= Balsateres cisnerosi ( Ponce-Saavedra and Sissom, 2004) ], type species by monotypy.

Vaejovis incertae sedis Ponce-Saavedra and Sissom, 2004: 541.

Thorellius: Soleglad and Fet, 2008: 53 , 58, 67, 77, 94, 95, tables 4, 9.

Balsateres González-Santillán and Prendini, 2013: 3 , 6, 7, 18, 24, 53, 59, table 1, figs. 3; 2015a: 349, 351, 352, 355, 356, 360, 362, 363, 367, 372, 403, fig., table 5, 6; Quijano- Ravel and Ponce-Saavedra, 2014: 17, 18, table 2; 2016: 50; Santibáñez-López et al., 2015: 7; Dupré, 2017: 9.

DIAGNOSIS: Balsateres may be distinguished from other genera of Syntropinae by the smooth dorsal lateral and lateral median carinae of metasomal segments I–V. The carinae and intercarinal surfaces of the carapace, pedipalp chela and patella, tergites, metasoma, and telson are smooth (figs. 5A, 9A, 10A, 11A, 17, 18, 19) unlike other syntropine genera, in which they are granular. Counts of macrosetae on the carinae of metasomal segments I–IV are also greatly reduced in Balsateres , compared with other syntropine genera as follows: dl, 0/0:0/0:0/0:1/1:4/4; lm, 0/0:0/0:0/0:0/0:2/2; vl and vsm, 1/1:1/1:1/1:1/1:3/3. Two or more macrosetae are present on these carinae in other syntropine genera, including Thorellius , which also presents low setal counts, e.g., the following counts on seg- ments I–V in T. cristimanus : dl, 2/2:3/3:3/3:3/3:7/6; Thorellius cisnerosi: Soleglad and Fet, 2008: 26 , lm, 1/1:3/3:3/3:4/4:4/4; vl, 2/2:3/3:3/3:3/3:7/7; vsm, 31, 53, 58, 67, 73, 77, 94, 95, tables 4, 9; 3/3:3/3:3/3:3/3:5/5. Baldazo-Monsivaiz et al., 2012: 146, table 1; Balsateres is most closely related to Thorellius, 2013: 101 , table 1; Ponce-Saavedra and with which it shares broad pedipalpal and meta- Francke, 2013: 77, table 2. somal carinae, but all carinae are smooth in Balsateres , instead of moderately to densely granular, Balsateres cisnerosi: González-Santillán and as in Thorellius . The pedipalp chela fingers of Bal- Prendini, 2013: 3, 6, 7, 18, 21, 24, 28, 37, 53, sateres are sublinear, without a distinct proximal 55, 57, 59, table 1, figs. 3, 16A, 20A, 28A; gap, when closed, unlike Thorellius , in which a 2015a: 351–353, 357, 363, 376, 380, 382, distinct proximal gap is evident in both sexes. The 387, 399, figs. 3–5, 6; Quijano-Ravel and two genera also differ in base coloration and Ponce-Saavedra, 2014: 17, 18, 20, table 2; infuscation. Balsateres is yellowish and almost 2016: 50; Dupré, 2017: 9. immaculate, except for the ocular tubercle, which

TYPE MATERIAL: MEXICO: Michoacán: is infuscate, whereas most Thorellius species are

Municipio de Churumuco: Holotype ♂ ( UMSNH) darker and reddish in color, with infuscation on

[lost], Churumuco, 18°40′15″N 101°38′39″W, the carapace and tergites. Only T. intrepidus is

7.i.2000, J. Ponce. Paratypes: Churumuco, paler and mostly immaculate. Balsateres also

18°40′15″N 101°38′39″W, 1 ♀ ( UMSNH) [lost], shares with T. intrepidus a pair of prolateral den-

9.ix.2000, J. Ponce, 1 ♂, 1 subad. ♂, 1 juv. ♂ ticles at the seventh position on the pedipalp chela

( UMSNH) [lost], 11.xi.2000, R. Moreno, 1 ♂, 1 ♀ movable finger, whereas other species of Thorellius

( UMSNH) [lost]; Cerro de Turitzio, 18°31′41″N possess a single denticle at the seventh position.

100°55′27″W, 1 ♀ ( UMSNH) [lost]. Municipio de Balsateres resembles species of Mesomexovis

Arúa: Huetamo , 9.ix.2000, R. Moreno and R. in possessing obsolete ventral carinae on meta-

Cancino, 1 ♀ ( UMSNH) [lost]. Municipio de somal segments I–IV, but Mesomexovis are more

Carácuaro: El Carrizal, 19°09′00″N 101°06′19″W, setose and infuscate.

vii.2000, E. Miranda, 1 ex. ( UMSNH) [lost]. Neo-

INCLUDED SPECIES: Balsateres cisnerosi

type ♂ ( CNAN T 01249), Churumuco de Morelos, ( Ponce-Saavedra and Sissom, 2004).

18°40′15″N 101°38′39″W, 217 m, 1.vii.2000, J.

DISTRIBUTION: Balsateres cisnerosi is endemic

Ponce-Saavedra. The holotype and paratypes are to the Balsas Depression in the Mexican states of

apparently lost (J. Ponce-Saavedra and G. Mon- Michoacán and Estado de México (fig. 2).

tiel-Parra, personal commun.) and a neotype is

hereby designated.

Balsateres cisnerosi DIAGNOSIS: As for genus.

( Ponce-Saavedra and Sissom, 2004) DESCRIPTION: The following redescription,

which is based on topotypes and additional mate- Figures 1A , 2 , 5A , 7A, 8A, 9A 10A, 11A, 12A–

rial examined, supplements the original descrip-

C, 14A, 15A–C, 16–18; tables 1, 8

tion by Ponce-Saavedra and Sissom (2004). Vaejovis cisnerosi Ponce-Saavedra and Sissom, Color and infuscation: Chelicerae, carapace,

2004: 540, figs 1–4, table 1. pedipalps, legs, tergites, sternites, metasoma, and FIGURE 7. Balsateres González-Santillán and Prendini, 2013 , Kuarapu Francke and Ponce-Saavedra, 2010 , and Thorellius Soleglad and Fet, 2008 , sternum, genital operculum and pectines. A. Balsateres cisnerosi ( Ponce-Saavedra and Sissom, 2004) , ♂ (CNAN SC2441). B. Kuarapu purhepecha Francke and Ponce-Saavedra, 2010 , paratype ♂ (AMNH). C. Thorellius intrepidus (Thorell, 1876) , ♂ (AMNH). D. Thorellius tekuani , sp. nov., holotype ♂ (CNAN T01250). E. Thorellius wixarika , sp. nov., holotype ♂ (CNAN T01251). F. Thorellius yuyuawi , sp. nov., holotype ♂ (CNAN T01252). G. Thorellius cristimanus ( Pocock, 1898) , ♂ (AMNH). Scale bars = 1 mm.

telson base color uniformly yellow, mostly immaculate, except as follows. Carapace and tergites faintly infuscate. Coxosternal region, genital operculum, basal piece, and pectines pale whitish. Pedipalp chela fingers slightly orange to reddish proximally, darker than manus. Telson aculeus dark reddish posteriorly.

Chelicerae: Manus dorsal surface smooth, distal margin with two pairs of setae. Movable finger, ventral surface with serrula, comprising 14/14 tines, in distal half.

Carapace: Length 1.09/1.08× greater than posterior width (table 1). Anterior margin slightly emarginate, with shallow median notch, with three pairs of major macrosetae (fig. 5A). Three pairs of lateral ocelli, anterolateral and median lateral pairs equal in size, posterolateral pair approximately half the size. Median ocular tubercle slightly raised, situated in anterior half of carapace, superciliary carinae obsolete, lower than median ocelli. Median ocelli approximately twice the size of anterolateral ocelli. Anteromedian and posteromedian sulci moderate to deep, posterolateral and posterior transverse sulci shallow. Interocular surface glabrous, anterolateral surfaces with scattered granules, other surfaces mostly smooth (fig. 5A).

Coxosternal region: Sternum subequilateral pentagonal (fig. 7A); width 1.09/1.11× greater than length (table 1); median sulcus deep; surfaces smooth, with three pairs of macrosetae, anteriorly, medially and posteriorly on lobes. Coxae surfaces smooth; coxa II subproximal margin with three oblique slitlike structures, adjacent to smooth protuberance; coxal endite II proximal margin with moderate depression, medial margin smooth. Coxa IV 2× longer than coxa II (table 1).

Pedipalps: Femur, intercarinal surfaces glabrous and smooth (figs. 16, 17A); dorsal prolateral, dorsal retrolateral, and ventral prolateral carinae complete, granular (fig. 17A); retrolateral dorsosubmedian carina obsolete, mostly smooth, with few large granules proximally, and two macrosetae; prolateral ventrosubmedian carinae vestigial, reduced to proximal tubercle, six granules, and two macrosetae medially; prolateral ventral carina vestigial, reduced to low tubercle proximally, and three macrosetae in proximal half; retrolateral ventral, ventromedian, and ventral retrosubmedian carinae vestigial, each reduced to few granules proximally. Patella slightly wider than femur (table 1); intercarinal surfaces smooth and glabrous (fig. 17B–E); dorsal prolateral and ventral prolateral carinae complete, finely granular, becoming smooth distally; dorsal retrolateral and ventral retrosubmedian carinae complete, costate; ventral median carina vestigial, reduced to row of low granules proximally; retrolateral dorsosubmedian and retrolateral median carinae absent; prolateral process well developed; prolateral median carina partial, comprising proximal tubercle, two or three low granules, and two macrosetae, proximal larger than distal; prolateral ventral carina vestigial, reduced to low tubercle proximally, and two macrosetae, proximal larger than the distal. Chela 1.6× longer than patella and 1.7× longer than femur (table 1). Manus incrassate (fig. 18), 1.4× wider than patella, 1.6× wider than femur (table 1); intercarinal surfaces smooth and glabrous; prolateral dorsal, dorsal prosubmedian, and dorsal prolateral carinae fused into broad smooth costa; dorsal median, dorsal retrolateral, retrolateral median, ventral retrolateral, ventral median, ventral prolateral, and prolateral median carinae complete, broad, costate; dorsal retrosubmedian accessory carina vestigial, costate; other carinae absent. Fixed and movable fingers, dentate margins very shallowly emarginate, notches and lobes obsolete (♂), or sublinear, notches and lobes absent (♀), fitting together evenly such that no gap evident when closed (fig. 18B, D); fixed finger, median denticle row comprising five denticle subrows flanked by six prolateral and five retrolateral denticles; movable finger, median denticle row comprising six denticle subrows, flanked by eight prolateral and six retrolateral denticles, proximal two prolateral denticles adjacent, terminal subrow comprising single denticle. Trichobothrial pattern type C, orthobothriotaxic; chela trichobothrium Db situated on dorsal

TABLE 1

retrolateral carina, in proximal fifth of manus; Dt situated at midpoint of manus; db–dt and eb–et distributed along entire length of fixed finger with db situated proximal and eb subproximal; ib situated between sixth prolateral denticle and seta demarcating position of seventh denticle, which is absent; it aligned with sixth prolateral denticle (fig. 18D).

Legs: Basitarsi retrolateral dorsal spinule row complete on I and II, vestigial, reduced to few distal spinules on III and IV; retrolateral median spinule row absent on I–IV; prolateral ventral spinule row vestigial, reduced to few distal spinules on I–IV; retrolateral ventral spinule row complete on I–III, vestigial, reduced to few distal spinules on IV; macrosetal counts on legs I–IV, respectively: dorsal, 3:3:3:3; retrolateral dorsal, 4:4:4:4, prolateral ventral, 3:5:5:5, distal spinules small and delicate; retrolateral ventral, 4:8:9:9; dorsal and retrolateral dorsal macrosetae arranged into two separate rows on I–IV. Telotarsi I–IV, each with single ventromedian row of spinules, curved proximally, and two or three pairs of ventrodistal spinules (fig. 11A).

Genital operculum: Width greater than length (fig. 7A), 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 1.2× longer than trunk (table 8). Median lobe, ental terminus prominent, rounded (fig. 12A–C). Dorsal trough margin long, narrow, curving proximally, well separated from ventral trough (fig. 12C). Dorsal and ventral trough margins, terminal spinelike processes fused into prominent, bifurcate hook, situated distally on lamina dorsal margin (fig. 12A, B). Basal plate of inner lobe well separated from ventral trough (fig. 14A); spine of inner lobe long and markedly sclerotized. Hemimating plug developed from inner lobe; distal barb 0.4× longer than basal plate (fig. 15A–C); distal barb margin with 25 elongated spinules; secondary spine on ventromedian side of distal barb moderately developed.

Pectines: Basal piece with three pairs of major macrosetae. Marginal lamella comprising three sclerites. Medial lamella proximal sclerites fused, 15 distal sclerites separate (fig. 7A). Fulcra, 21–22 (♂), 19–20 (♀). Pectinal teeth, 20–23 (♂), 20–21 (♀). 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 smooth. Posttergites I–VI, intercarinal surfaces smooth medially, finely granular posterolaterally, dorsal median and dorsal lateral carinae obsolete, reduced to few, low granules; VII, intercarinal surfaces finely granular, dorsal median carina vestigial, restricted to anterior third, costategranular; dorsal submedian carinae vestigial, each reduced to anterior granule, dorsal lateral and lateral median carinae costate to costategranular, converging anteromedially, posterior granules slightly larger, spiniform.

Sternites: Sternites III–VII, surfaces acarinate, smooth; III–VI, spiracles elongate, slitlike, 4× longer than wide; V, posteromedian hyaline glandular area moderately developed (♂) or obsolete (♀).

Metasoma: Metasoma 1.5× longer than mesosoma (table 1). Segments I–V, respectively 1.3/1.4, 1.1, 1.0, 0.7, 0.5× wider than long; V, 1.3× wider than telson vesicle. Segments I–V, intercarinal surfaces smooth (figs. 8A, 9A); dorsal lateral carinae complete, costate, terminating in enlarged spiniform granules posteriorly on I–IV, obsolete on V; lateral median carinae vestigial, costate, terminating in enlarged spiniform granules posteriorly on I–III, lobate posteriorly on IV, vestigial, costate on V; lateral inframedian, ventral lateral, and ventral submedian carinae obsolete on I–V, except ventral lateral carinae granular on posterior third of V. Macrosetal counts on carinae of segments I–V, respectively: dorsal lateral carinae, 0:0:1:1:4; lateral median carinae, 0:1:1:1:2; lateral inframedian carinae, 0:0:0:0:0; ventral lateral carinae, 1:1:1:1:3; ventral submedian carinae, 1:1:1:1:3; ventral sublateral carinae, 0:0:0:0:2.

Telson: Vesicle globose, length 2.5× greater than width (table 1); ventral and dorsal surfaces smooth; ventral carinae each with two pairs of macrosetae. Subaculear tubercle vestigial, comprising low broad granule (fig. 10A). Aculeus, laterobasal microserration comprising 4–6 spinules.

DISTRIBUTION: As for genus.

ECOLOGY: Balsateres cisnerosi inhabits tropical deciduous forest at altitudes ranging from 195 to 672 m. Specimens have been observed at night, doorkeeping at the entrances of their burrows in open, sandy areas, and excavated from the burrows, approximately 15–20 cm deep with a circular entrance ( Ponce-Saavedra and Sissom, 2004). The habitat and habitus are consistent with the psammophilous ecomorphotype ( Prendini, 2001) although the leg setation, in which the dorsal and retrolateral dorsal macrosetae are arranged into two separate rows, not forming a setal comb, is atypical. Balsateres cisnerosi was collected in sympatry with the following species: the buthids Centruroides balsasensis Ponce-Saavedra and Francke, 2004 (observed preying on B. cisnerosi ) and Centruroides limpidus (Karsch, 1879) ; the diplocentrids Diplocentrus churumuco Francke and Ponce Saavedra, 2005 , and Kolotl poncei (Francke and Quijano-Ravell, 2009) ; and the vaejovids Konetontli aff. kuarapu ; Kuarapu purhepecha , Mesomexovis sp. , and Thorellius tekuani , sp. nov. (J. Ponce-Saavedra and A. Quijano-Ravel, personal commun.). Although sympatric, B. cisnerosi was collected in open, sandy areas, whereas K. purhepecha was collected on road cuts and under rock piles.

REMARKS: Due to the unique morphology of V. cisnerosi, Ponce-Saavedra and Sissom (2004) refrained from assigning it to one of the five species groups of Vaejovis recognized at the time. Soleglad and Fet (2008) transferred it to Thorellius , the name devised for Hoffmann’s (1931) “second section” of Vaejovis , later termed the intrepidus group by Sissom (1989), without quantitatively testing its placement. Santibáñez- López and Sissom (2010) also referred this species to the intrepidus group. González-Santillán and Prendini (2013) justified the creation of Balsateres based on its unique diagnostic character combination.

ADDITIONAL MATERIAL EXAMINED: MEX- ICO: Estado de México: Municipio de Luvianus: Pungarancho, 3 km E, 19°01.775′N 100°28.406′W, 16.iv.2002, E. González, O. Francke and S. Reynaud, tropical dry forest, 1 ♀ ( CNAN SC2662 ). Michoacán: Municipio de Carácuaro : La Berea, 19°08.173′N 102°13.195′W, 445 m, 11.vii.2005, O. Francke, J. Ponce, M. Córdova, A. Jaimes, G. Francke and V. Capovila, 1 juv. ♀ ( AMCC [ LP 5320 ]) ; Las Cocinas, Hwy Nocupetaro – Carácuaro, 19°01.880′N 101°14.067′W, 672 m, 14. vii.2005, O. Francke, J. Ponce, M. Córdova, A. Jaimes, G. Francke, and V. Capovilla, 1 subad. ♂ ( AMCC [ LP 5269 ]), [18°39.393′N 101°39.124′W], 8.ix.2001, J. Ponce, 1 ♂ ( CNAN SC2665 ), 1 ♀ ( AMCC [ LP 2019 ]), R. Cancino, 1 ♂ ( AMCC [ LP 2018 ]). Municipio de Huetamo: Arua [18°35′N 100°53′W], 9.ix.2000, R. Cancino, 2 ♀ ( CNAN SC2663 ), 23.viii.2001, 1 ♂ ( CNAN SC2661 ). Municipio de La Huacana : El Chauz [18°52′50.14″N 102°00′27.55″W], 20.ii.2004 R. Lanoy Cardenas, 1 juv. ( CNAN SC3030 ) ; El Vado, 18°48.908′N 101°54.976′W, 198 m, 20.v.2007, O. Francke, J. Ponce, M. Villaseñor, and A. Quijano, 1 ♂, 2 ♀ ( AMNH), 1 ♂, 2 ♀ ( CNAN), 18°48′51.34″N 101°55′10.31″W, 248 m, 30. viii.2008, O. Francke, H. Montaño, J. Ponce, and A. Quijano, 2 ♂, 2 ♀, 2 subad. ♂ ( CNAN SC2441 ). Municipio de Parácuaro: La Berea, 19°08.173′N 102°13.195′W, 445 m, 11.vii.2005, O. Francke, J. Ponce, M. Córdova, A. Jaimes, G. Francke, and V. Capovilla, 1 juv. ( AMCC [ LP 5320 ]). Municipio de Zingangio : Las Juntas Ranch, 24 km W of Capeo, 18°32.270′N 101°11.603′W, 195 m, 8.iv.2012, O. Francke et al., 5 ♂, 1 subad. ♂ ( CNAN SC2442 ) .

Figures

FIGURE 1. Habitus in life. A. Balsateres cisnerosi (Ponce-Saavedra and Sissom, 2004), ♀; B. Kuarapu purhepecha Francke and Ponce-Saavedra, 2010, ♀; C. Thorellius cristimanus (Pocock, 1898), ♂; and D. Thorellius intrepidus (Thorell, 1876), ♀.

FIGURE 2. Map of central Mexico, plotting known locality records for the vaejovid scorpion species, Balsateres cisnerosi (Ponce-Saavedra and Sissom, 2004) (triangles) and Thorellius cristimanus (Pocock, 1898) (squares), based on data collected in the present study.

FIGURE 5. Balsateres González-Santillán and Prendini, 2013, Kuarapu Francke and Ponce-Saavedra, 2010, and Thorellius Soleglad and Fet, 2008, carapaces. 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). Scale bars = 1 mm

FIGURE 8. Balsateres González-Santillán and Prendini, 2013, Kuarapu Francke and Ponce-Saavedra, 2010, and ThorelliusSoleglad and Fet, 2008, metasomal segments I–V, lateral 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.

FIGURE 9. Balsateres González-Santillán and Prendini, 2013, Kuarapu Francke and Ponce-Saavedra, 2010, and Thorellius Soleglad and Fet, 2008, metasomal segments I–V, 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.