Syntropinae Kraepelin, 1905

González-Santillán, Edmundo & Prendini, Lorenzo, 2013, Redefinition And Generic Revision Of The North American Vaejovid Scorpion Subfamily Syntropinae Kraepelin, 1905, With Descriptions Of Six New Genera, Bulletin of the American Museum of Natural History 2013 (382), pp. 1-71 : 16-21

Subfamily Syntropinae Kraepelin, 1905

Syntropinae Kraepelin, 1905: 340 , type genus Syntropis Kraepelin, 1900 ; Birula, 1917a: 163, 1917b: 57; Werner, 1934: 281; Mello-Leitão, 1945: 118; Millot and Vachon, 1949: 428; Brues et al., 1954: 704; Gertsch, 1958: 14, 15; Stahnke, 1974: 112, 113; Stockwell, 1992: 408; Sissom, 2000: 107; Soleglad and Fet, 2003: 109; 2008: 1, 2, 4–6, 13, 26–28, 33, 35–40, 45, 46, 50, 51, 53, 54, 62, 69, 72–74, 76, 77, 82, 84–86, 88–90, 93–97, 103, figs. 34–36, 41–56, 122–125, 164–167, 183, 196, 202– 204, tables 1–3 , 8, 9.

Paravaejovini Soleglad and Fet, 2008 , syn. nov.: 1, 3, 13, 36, 46, 51, 75, 82, 84, 102, figs. 57, 126, 196, 197, tables 2 , 8, 9.

Syntropini: Soleglad and Fet, 2008: 1, 2, 4, 13, 30, 33, 34, 38, 40, 43, 45, 46, 49, 50, 51, 53, 54, 57, 69, 71–74, 76, 77, 84, 85, 89, 91, 94, 95, 103, figs. 75– 82, 103–110, 122–124, 170–181, 194–196, 203, 204; tables 2 , 3 , 8, 9.

Syntropina: Soleglad and Fet, 2008: 1, 32, 46, 51, 53, 74, 76, 77, 90, 91, figs. 196, 203, table 9.

Thorelliina Soleglad and Fet, 2008 , syn. nov.: 1, 45, 51, 53, 71, 74, 89, 91, 92, 95, 96, 103, figs. 196, 204, table 9.

DIAGNOSIS: The monophyletic group of vaejovid scorpion taxa redefined here as subfamily Syntropinae may be separated from all other vaejovid taxa by the following unique synapomorphy: margin of distal barb of sclerotized hemimating plug on hemispermatophore spinose (armed with spines, hooks or teeth; fig. 8C–E). The following other vaejovid taxa possess a sclerotized hemimating plug on the hemispermatophore, but the margin of the distal barb is smooth (i.e. not spinose; fig. 8F): the monophyletic group comprising Gertschius Graham and Soleglad, 2007 , Serradigitus , Stahnkeus Soleglad and Fet, 2006 , and Wernerius Soleglad and Fet, 2008 , formerly included in Syntropinae as tribe Stahnkeini ( Soleglad and Fet, 2008) ; Franckeus Soleglad and Fet, 2005 , and the nigrescens group of Vaejovis ; some species of the mexicanus group of Vaejovis , e.g., Vaejovis vorhiesi Stahnke, 1940 ; and some species of Pseudouroctonus , e.g., Pseudouroctonus apacheanus ( Gertsch and Soleglad, 1972) .

Syntropinae are most closely related to the monophyletic group comprising Gertschius , Serradigitus , Stahnkeus , and Wernerius , based on the fused sclerites of the female genital operculum, which are connected the entire length. Syntropinae may be further separated from these taxa as follows. The first to third pectinal teeth are unmodified in Syntropinae , but enlarged, ovoid, and devoid of sensilla in Gertschius , Serradigitus , Stahnkeus , and Wernerius . The pedipalp chela fingers of Syntropinae possess low, subserrated rows of denticles and small to moderately developed terminal retrolateral denticles (figs. 17–19), compared with the fingers of Serradigitus and Stahnkeus , which possess strongly serrated denticle rows and enlarged, hooklike denticles. Pedipalp chela trichobothrium est is situated between RD4 and RD5 (and/or associated macrosetae) in Syntropinae (figs. 10B, 17B, D, 18B, 19B, D) but between RD3 and RD4 (and/or associated macrosetae) in Stahnkeus . One to six pairs of ventrodistal spinules are present on the telotarsi of Syntropinae (figs. 21A–C, 22A, B), compared with only one pair on the telotarsi of Gertschius , Serradigitus , Stahnkeus , and Wernerius .

Additional diagnostic characters of Syntropinae are as follows: carapace with superciliary carinae usually higher than median ocelli (fig. 16); cheliceral serrula comprising up to 30 tines; pedipalp chela trichobothrium Db situated on or dorsal to the drl carina (figs. 10A, 17A, C, 18A, D, 19A, C); chela dps, plm, pld, and plvs carinae obsolete to costate, except in Kochius and a few species of Chihuahuanus , gen. nov., and Thorellius (figs. 10A, 11A, B, 17A, C, 18A); patella rlm and rlds carinae obsolete to costate, except in Kochius , Syntropis , and a few species of Thorellius (fig. 9F); telotarsi I–IV each with two proventral setae, two or three retro- ventral macrosetae, and 1–6 pairs of ventrodistal spinules (figs. 13B, 21A–C, 22A, B); aculeus with lateral microserration vestigial to well developed.

INCLUDED GENERA: Balsateres , gen. nov.; Chihuahuanus , gen. nov.; Kochius Soleglad and Fet, 2008 ; Konetontli , gen. nov.; Kuarapu Francke and Ponce-Saavedra, 2010 ; Maaykuyak , gen. nov.; Mesomexovis , gen. nov.; Paravaejovis Williams, 1980 ; Syntropis Kraepelin, 1900 ; Thorellius Soleglad and Fet, 2008 ; Vizcaino , gen. nov.

DISTRIBUTION: Subfamily Syntropinae is endemic to Mexico and the United States (figs. 3–6). Species of the subfamily have been recorded from 27 states in Mexico (Aguascalientes, Baja California, Baja California Sur, Coahuila, Colima, Chiapas, Chihuahua, Durango, Estado de México, Guanajuato, Guerrero, Hidalgo, Jalisco, Michoacán, Morelos, Nayarit, Nuevo León, Oaxaca, Puebla, Queretaro, San Luis Potosí, Sinaloa, Sonora, Tamaulipas, Tlaxcala, Veracruz, Zacatecas) and seven states in the United States (Arizona, California, Nevada, New Mexico, Oregon, Texas, Utah).

REMARKS: Based on phylogenetic analyses presented elsewhere (fig. 7; González-Santillán and Prendini, in press), Syntropinae is hereby restricted to the 11 genera listed above. Four genera, formerly accommodated in tribe Stahnkeini Soleglad and Fet, 2008 , are hereby removed from Syntropinae : Gertschius Graham and Soleglad, 2007 ; Serradigitus Stahnke, 1974 ; Stahnkeus Soleglad and Fet, 2006 ; Wernerius Soleglad and Fet, 2008 .

KEY TO IDENTIFICATION OF THE GENERA OF SYNTROPINAE

1. Telson, dorsal surface (adult ♂, ♀) with whitish glandular area (fig. 26); metasomal segment V, ventral surface glabrous (smooth) to matte; pedipalp chela fixed finger, median denticle row with five primary subrows of median denticles and five prolateral denticles............. 2

– Telson, dorsal surface (adult ♂, ♀) without whitish glandular area; metasomal segment V, ventral surface matte to shagreened; pedipalp chela fixed finger, median denticle row with five or six primary subrows of median denticles and five or six prolateral denticles........................ 3

2. Telson, dorsal surface (adult ♂, ♀) with small fusiform, whitish glandular area anterior to base of aculeus (fig. 26A; may be minute in Chihuahuanus bilineatus (Pocock, 1898) , comb. nov.); pedipalp chela, fixed finger trichobothrium et situated midway between RD3 and RD4 or closer to RD3 than to RD4 (fig. 17B)....... Chihuahuanus , gen. nov.

– Telson, dorsal surface (adult ♂, ♀) with medium-sized oval, whitish glandular area medially (fig. 26B; may be reduced in adult ♀); pedipalp chela, fixed finger trichobothrium et aligned with or closer to RD4 than to RD3......... Maaykuyak , gen. nov.

3. Sternite VII, ventral surface (adult ♂, ♀) with whitish glandular area medially (fig. 23)... 4

– Sternite VII, ventral surface (adult ♂, ♀) without whitish glandular area........ 6

4. Sternite VII, ventral surface (adult ♂, ♀) with whitish glandular area between ventrosubmedian carinae, extending almost entire length of segment (fig. 23B); metasomal segments I– IV, ventrosubmedian carinae absent, ventromedian carinae present, smooth to finely granular (fig. 24C)............ Syntropis

– Sternite VII, ventral surface (adult ♂, ♀) with raised, whitish triangular boss, restricted to posteromedial third of segment; metasomal segments I–IV, ventrosubmedian carinae present, granular, ventromedian carinae absent (fig. 24A, B).................. 5

5. Pedipalp chela manus (adult ♂, ♀) incrassate, with carinae well developed, granular and immaculate (fig. 18A, B); sternite VII, ventrolateral surfaces shagreened; telson (♀), lateral and especially ventral surfaces, macrosetae as long as or longer than aculeus (fig. 25A).... Kochius

– Pedipalp chela manus (adult ♂, ♀) slender, with carinae obsolete and infuscate (fig. 18C, D); sternite VII, ventrolateral surfaces glabrous; telson (♀), lateral and ventral surfaces, macrosetae shorter than aculeus.... Kuarapu

6. Pedipalp chela movable finger, median denticle row with five retrolateral denticles and four primary subrows of median denticles, terminal row absent (fig. 20C); legs I–III, basitarsi, dorsal and retrodorsal macrosetae elongated, arranged into sublinear row, forming setal comb (fig. 21C); telotarsi, ventral spinules elongate and slender (fig. 21C)............... Vizcaino , gen. nov.

– Pedipalp chela movable finger, median denticle row with five or more retrolateral denticles and more than four primary subrows of median denticles, terminal row present (figs. 12B, 17C, 19A, C, 20A, B); legs I–III, basitarsi, dorsal and retrodorsal macrosetae usually short and arranged into separate rows, not forming setal comb (fig. 21B), except in Paravaejovis pumilis ( Williams, 1980) (fig. 22C); telotarsi, ventral spinules short and stout (fig. 21B)...... 7

7. Total body length (adult ♂, ♀) less than 25 mm; telotarsi each with one pair of ventrodistal spinules; telson with conspicuous subaculear tubercle (figs. 25B, C, 28B).................... Konetontli , gen. nov.

– Total body length (adult ♂, ♀) more than 26 mm; telotarsi each with two or more pairs of ventrodistal spinules (fig. 21B; except in P. pumilis , fig. 22C); telson without conspicuous subaculear tubercle (figs. 27B, C, 28A, C)... 8

8. Carapace and tergites densely infuscate, except in Mesomexovis atenango ( Francke and González-Santillán, 2007) , comb. nov.; pedipalp chela manus, dorsal carinae (dm, dpl, drl, drs, drsa) smooth (fig. 19A, B); metasomal segments I–IV, ventrosubmedian and ventrolateral carinae infuscate, ventrosubmedian carinae absent on I and II, obsolete to weakly granular on III and IV (fig. 27B)....... Mesomexovis , gen. nov.

– Carapace and tergites immaculate to weakly infuscate; pedipalp chela manus, dorsal carinae (dm, dpl, drl, drs, drsa) smooth or granular (figs. 17C, D, 19C, D); metasomal segments I–IV, ventrosubmedian and ventrolateral carinae immaculate to weakly infuscate, ventrosubmedian carinae absent to obsolete on I and II, weakly costate to strongly granular on III and IV (fig. 27C)........ 9

9. Habitus gracile, total body length (adult ♂, ♀) 37–68 mm; base color pale, yellowish (fig. 1C); pedipalp chela manus, carinae obsolete to weakly developed, barely protruding above intercarinal surfaces, and rarely granular; trichobothrium Et 5 situated at base of fixed finger far removed from trichobothrium Et 4 (fig. 19D), less obvious in P. pumilis ............... Paravaejovis

– Habitus robust, total body length (adult ♂, ♀) 45–94 mm; base color dark brown to reddish, except in Balsateres cisnerosi ( Ponce-Saavedra and Sissom, 2004) , comb. nov.; pedipalp chela manus, carinae well developed, protruding markedly above intercarinal surfaces, smooth or granular (fig. 17C, D); trichobothrium Et 5 situated on manus close to trichobothrium Et 4 (fig. 17C, D).................... 10

10. Carapace, pedipalp chela and patella, tergites, metasoma, and telson, carinae and intercarinal surfaces smooth (figs. 16A, 28A); base color pale, yellowish; tergites immaculate; metasomal macrosetae counts low: 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; pedipalp chela fingers, proximal gap weak to obsolete........................... Balsateres , gen. nov.

– Carapace, pedipalp chela and patella, tergites, metasoma, and telson, carinae and intercarinal surfaces granular or crenulate (figs. 17C, D); base color dark brown to reddish; tergites infuscate; metasomal macrosetae counts moderate: dl, 2/2:3/3:3/3:3/3:7/6; lm, 1/1:3/3:3/3:4/ 4:4/4; vl, 2/2:3/3:3/3:3/3:7/7; vsm, 3/3:3/3:3/3:3/ 3:5/5 or greater; pedipalp chela fingers, proximal gap moderate to strong.... Thorellius