Typhlochactas Mitchell, 1971

Vignoli, Valerio & Prendini, Lorenzo, 2009, Systematic Revision Of The Troglomorphic North American Scorpion Family Typhlochactidae (Scorpiones: Chactoidea), Bulletin of the American Museum of Natural History 2009 (326), pp. 1-94 : 52-59

Typhlochactas Mitchell, 1971

Typhlochactas Mitchell, 1971: 238 ; type species by subsequent designation ( Mitchell, 1971: 238) Typhlochactas rhodesi Mitchell, 1968 .

Typhlochactas : Mitchell, 1968: 753–756, 771–774; 1971: 135, 138, 147 (part); Vachon, 1974: 914, 923; Soleglad, 1976: 253, 254; Mitchell and Peck, 1977: 159, 162, 164, 165, 167 (part; revised diagnosis); Francke, 1978: 44; Reddell, 1981: 115, 175; Francke, 1982a: 51–53, 57, 59–61; 1982b: 36; 1985: 14, 16, 20; 1986: 5, 8; Locket, 1986: 101, 111, 112; Sissom, 1988: 365, 370; 1990: 109, 114; Polis, 1990: 253; Nenilin and Fet, 1992: 10, 28; Stockwell, 1992: 410, 412–414, 419, figs. 3, 28, 30, 33, 35; Armas, 1994: 18, 19; Locket, 1995: 191; Kovařík, 1998: 142; Sissom and Cokendolpher, 1998: 285, 286; Beutelspacher Baigts, 2000: 6, 13–16, 18, 20, 48, 51, 53, 57, 151, fig. 11A, graph 1, tables 1, 2; Lourenço, 2000: 25; Lourenço and Sissom, 2000: 118; Sissom, 2000b: 498; Lourenço, 2001: 4; Soleglad and Sissom, 2001: 68, 69; Morrone et al., 2002: 93; Soleglad and Fet, 2003a: 7, 16, 17, 32, fig. 12; 2003b: 36, 67, 72, 74, 82, 106–109, 141,162, fig. 116, tables 3, 4, 9; Fet et al., 2004: 197; Rojas-Runjaic, 2004: 245; Prendini and Wheel- er, 2005: 454, 458, 460, 463, 468, 470, 477, 478, tables 3, 4, 5, 9, 10, figs. 18, 19; Fet and Soleglad, 2005: 12; Sissom and Hendrixson, 2005: 123, 130; Dupre´, 2006: 11; Francke and Savary, 2006: 29; Graham and Fet, 2006: 2; Volschenk and Prendini, 2008: 249; Botero-Trujillo and Francke, 2009: 2.

Typlochactas : Díaz Najera, 1975:3; Dupre´, 2007:11.

DIAGNOSIS: Typhlochactas is the sister taxon of Stygochactas (fig. 4). Species of Typhlochactas may be separated from Stygochactas and Sotanochactas on the basis of the following combination of characters. Size small to very small, total length (adult) less than 25 mm. Cheliceral fixed finger with three or four teeth, basal teeth usually not fused into a bicusp. Cheliceral movable finger with three, four, or (usually) five dorsal teeth (none, one, or two subdistal teeth). Carapace, anterior margin, median projection (epistome) usually present. Pedipalps not elongated, chela fingers similar in length to chela manus. Pedipalp femur, ventroexternal carina, patella, internomedian carina, and chela, dorsal secondary, digital, ventroexternal, ventromedian, and ventrointernal carinae, absent or obsolete. Chela fixed and movable fingers, median denticle row comprising 4–6 and 5–7 oblique primary subrows, respectively; basal primary subrows of fixed finger similar in length to other subrows. Patella trichobothrium v 1 situated level with or proximal to trichobothrium esb 1. Chela fixed finger, external trichobothria distributed across entire length of finger, with trichobothrium eb situated near base of finger; trichobothria it and ib situated near base of finger. Legs usually with prolateral pedal spurs. Basitarsi I and II, proventral surfaces usually with short row of closely aligned spinules subdistally. Telotarsi, ventral surface, usually with curved proximal row spinules and without ventromedian row of spinules. Tergite VII, dorsolateral carinae absent or vestigial. Metasomal segments I–IV, dorsosubmedian carinae obsolete; I–III, dorsolateral carinae absent or obsolete; I, median lateral carinae absent; I–V, ventrolateral carinae absent or obsolete; V, ventromedian carina absent.

INCLUDED TAXA: Six species: Typhlochactas cavicola Francke, 1986 ; Typhlochactas mitchelli Sissom, 1988 ; Typhlochactas reddelli Mitchell, 1968 ; Typhlochactas rhodesi Mitchell, 1968 ; Typhlochactas sissomi Francke et al., 2009 ; Typhlochactas sylvestris Mitchell and Peck, 1977 .

DISTRIBUTION: Endemic to Mexico. Recorded from four states: Oaxaca, Queretaro, Tamaulipas, Veracruz.

Typhlochactas cavicola Francke, 1986 ( Figs. 1B , 4 , 5D , 8C , 9G, 9H , 12C , 23–25 ; table 8)

Typhlochactas cavicola Francke, 1986: 5–9 , figs. 1– 10, 17–19, table 1.

Typhlochactas cavicola : Sissom, 1988: 365; Armas, 1994: 21; Lourenço, 1994: 182, 183; Kovařík, 1998: 142. Sissom and Cokendolpher, 1998: 285, 287 (table 1); Beutelspacher Baigts, 2000: 18, 20, 52, 54, 147, 151, map 22; Sissom, 2000b: 499; Volschenk et al., 2001: 161; Soleglad and Fet, 2003a: 7; 2003b: 30, 32, 76; Vignoli and Kovařík, 2003: 131; Sissom and Hendrixson, 2005: 127, table 6.2; Volschenk and Prendini, 2008: 236 (table 1).

TYPE MATERIAL: Mexico: Tamaulipas: Município de Guemez: Holotype: 1 ♀ ( AMNH), Cueva del Vandalismo [23 ° 519540 N 99 ° 269450W], ca. 2600 m, 1 km SE Rancho Nuevo, D. Honea, 15.III.1982.

DIAGNOSIS: Typhlochactas cavicola is most closely related to T. rhodesi (fig. 4). It may be separated from T. rhodesi and all other Typhlochactas species by the following combination of characters. Cheliceral fixed finger with four teeth (subdistal present); median

TABLE 8

Meristic data for Typhlochactas cavicola Francke, 1986 , Typhlochactas reddelli Mitchell, 1968 , and Typhlochactas rhodesi Mitchell, 1968 Measurements (mm) follow Francke et al. (2009). Abbreviations as follows: AMNH 5 American Museum of Natural History , New York ; IBUNAM 5 Instituto de Biología, Universidad Nacional Autonóma de México, Mexico City ; MNHN 5 Muséum National d’Histoire Naturelle, Paris

TABLE 8 (Continued)

and basal teeth not fused into bicusp. Cheliceral movable finger with four dorsal teeth (one subdistal present). Carapace, anterior margin, median projection (epistome) absent. Pedipalp chela fixed and movable fingers, median denticle row comprising six and five oblique primary subrows, respectively; basal primary subrow of movable finger considerably longer than other subrows; terminal denticle of fixed finger slightly larger than preceding denticles, fingertips interlocking evenly when closed. Legs without prolateral pedal spurs. Telotarsi without ventromedian row of spinules. Sclerites of genital operculum (♀) mostly fused, but loosely connected by membrane along entire length of suture, posterior edges free.

DESCRIPTION: The following description supplements the original description by Francke (1986).

Color: Carapace, pedipalps, legs, tergites, and metasoma, Raw Sienna (136). Pedipalp chela fingers darker; cheliceral teeth, ungues paler than aculeus (fig. 23).

Chelicerae: Manus, dorsal and ventral surfaces smooth; dorsal surface with three microsetae situated near base of fixed finger; movable finger with two larger microsetae and one smaller microseta. Fixed finger, dorsal margin with four teeth (distal, subdistal, median, and basal; fig. 5D); median and basal teeth not fused into bicusp; distal tooth largest, subdistal and median teeth equal, smaller, basal tooth smallest. Movable finger, internal distal and external distal teeth opposable, internal distal tooth completely overlapping external distal tooth in dorsal view; dorsal margin with four teeth (internal distal, subdistal, median, and basal), internal distal tooth largest, subdistal and median teeth equal, smaller, basal tooth smallest; ventral surface with well-developed serrula, covered in part by dense brush of macrosetae.

Carapace: Length slightly greater than anterior width (table 8). Anterior margin sublinear, without median projection (epistome); asetose (fig. 23A). Posterior margin sublinear; asetose. Median and lateral ocelli absent. Median longitudinal, posterolateral and posterior transverse sulci shallow. Surface smooth, acarinate, with few microsetae.

Pedipalps: Femur, dorsoexternal, ventroexternal, and ventrointernal carinae obsolete, comprising a few small scattered granules, decreasing in number distally; dorsointernal carina absent. Dorsal and external surfaces smooth, with several short setae (fig. 25A); internal surface with few granules proximally. Patella, ventrointernal carina obsolete, granular; other carinae absent (fig. 25B–D). Intercarinal surfaces less granular than femur, with several short setae. Chela relatively large, longer than carapace (table 8). Manus acarinate (fig. 24); manus and fingers, surfaces mostly smooth, with several long setae; dorsal, external, and internal surfaces with small scattered granules; internal surface additionally with pair of prominent, isolated granules situated close together at base of fixed finger. Fixed finger, median denticle row comprising six oblique primary subrows (fig. 9G); internal denticles larger than external denticles; terminal denticle small, equal to internal denticle, fingertips interlocking evenly when closed. Movable finger, median denticle row comprising five oblique primary subrows of denticles, basal primary subrow considerably longer than other subrows (fig. 9H); terminal denticle small.

Trichobothria: Femur with three trichobothria (fig. 25A): one external (e), one dorsal (d), one internal (i). Patella with 19 trichobothria (fig. 25B–D), five petite (d 1, d 2, et 2, esb 2, eb 2), one accessory (em 3): two ventral (v 1, v 2); 14 external (et 1 –et 3, est, em 1 – em 3, esb 1, esb 2, eb 1 – eb 5); two dorsal (d 1, d 2); one internal (i). Chela with 26 trichobothria (fig. 24), seven petite (V 1, Et 4, Et 5, Esb, Db, esb , db): 16 on manus, four ventral (V 1 –V 4), 10 external (Et 1 – Et 5, Est, Esb, Eb 1 – Eb 3), two dorsal (Db, Dt); 10 on fixed finger, four external (et, est, esb , eb), four dorsal (dt, dst, dsb, db), two internal (it, ib).

Legs: All segments smooth, setose, especially telotarsi. Basitarsi I–IV without prolateral pedal spurs; I and II, proventral surface with short subdistal row of closely aligned spinules. Telotarsi, dorsomedian lobe with one microseta; ventral surface with curved row of spinules, proximally, and four or five submedian pairs of subspiniform macrosetae; ungues well developed, curved, equal in length; dactyl long, prominent.

Tergites: Surfaces I–VI, smooth, acarinate; VII, mostly smooth, posterolateral surfaces granular, dorsosubmedian carinae vestigial, reduced to few posterior granules, dorsolateral carinae absent (fig. 23A). Lateral and posterior margins setose.

Sternum: Posterior width slightly greater than length (fig. 8C); apex rounded; lateral margins converging anteriorly; lateral lobes flat; posterior depression moderate. Surfaces with few macrosetae.

Genital operculum: Sclerites mostly fused, but loosely connected by membrane along entire length of suture, posterior edges free (fig. 8C); suboval, each with pair of macrosetae and several microsetae on margin.

Pectines: Pectinal plate, surface with pair of macrosetae. Lamella comprising three segments (fig. 8C); surfaces with several macrosetae and microsetae. Tooth count, 5/ 5; teeth equal to subequal, except for distal tooth, larger, pointed.

Sternites: Surfaces smooth, acarinate (fig. 23B); surfaces and margins with several microsetae. Respiratory spiracles (stigmata) small, round, situated posterolaterally.

Metasoma: Segments elongated, progressively increasing in length, decreasing in width (fig. 12C), I and II width greater than length, III–V length greater than width (table 8). Dorsosubmedian carinae, segments I–IV, obsolete, granular, distal granules not noticeably larger than preceding granules (fig. 12C). Dorsolateral carinae, segment V, obsolete, granular. Dorsolateral, median lateral, and ventrosubmedian carinae, segments I–IV, ventrolateral carinae, segments I–V, and ventromedian carina, segment V, absent. Dorsal intercarinal surfaces, segments I–V, slightly concave, uniformly finely granular; other surfaces smooth. All segments setose.

Telson : Vesicle broad, elongated, flattened dorsally, rounded ventrally; anterodorsal lateral lobes absent (fig. 12C); surface smooth, setae distributed uniformly across ventral surface. Aculeus short, slightly curved, arising gradually from vesicle.

Hemispermatophore: Male unknown.

Ontogenetic variation: It is unclear whether the holotype female is adult.

Sexual dimorphism: Male unknown.

Geographical variation: Single specimen.

REMARKS: Francke (1986: 5, 6) described the holotype as a male and stated that the genital opercula were ‘‘without membranous connection,’’ but noted that genital papillae ‘‘could not be detected.’’ We confirmed the absence of genital papillae but observed that the genital opercula are mostly fused, being loosely connected by a membrane along the entire length of the suture, although the posterior edges are free. These observations, together with the shape of the pectinal teeth, lead us to conclude that the holotype is female. Males of all other typhlochactid species display genital papillae and divided genital opercula.

DISTRIBUTION: Known only from the type locality, Cueva del Vandalismo, in the Sistema Purificación karstic region, northwest of Ciudad Victoria, Tamaulipas (central-eastern Mexico) (fig. 1B).

ECOLOGY: No additional data are available for the cave from which this troglobitic species was collected.

Figures

Fig. 1. Map of Mexico, showing the known localities of Typhlochactidae Mitchell, 1971. A. Known localities of Alacraninae, new subfamily: Type locality of Alacran tartarus Francke, 1982, on the Huautla Plateau, Oaxaca (1); Te Cimutaá Cave, Oaxaca, a new record for the genus possibly representing a new species of Alacran Francke, 1982 (2). B. Type localities of Typhlochactinae Mitchell, 1971: Sotanochactas elliotti (Mitchell, 1971), El Sótano de Yerbaniz, San Luis Potosí (1); Stygochactas granulosus (Sissom and Cokendolpher, 1998), Sótano de Poncho, Veracruz (2); Typhlochactas cavicola Francke, 1986, Cueva del Vandalismo, Tamaulipas (3); Typhlochactas mitchelli Sissom, 1988, Cerro Ocote, Oaxaca (4); Typhlochactas reddelli Mitchell, 1968, La Cueva del Ojo de Agua de Tlilapan, Veracruz (5); Typhlochactas rhodesi Mitchell, 1968, La Cueva de la Mina, Tamaulipas (6); Typhlochactas sissomi Francke et al., 2009, Cañada de La Joya, Queretaro (7); Typhlochactas sylvestris Mitchell and Peck, 1977, 25 km S Valle Nacional, Oaxaca (8).

Fig. 4. Single most parsimonious tree (312 steps, CI: 57, RI: 56) obtained by cladistic analysis of 142 morphological characters (appendix 1) scored for nine taxa in the family Typhlochactidae Mitchell, 1971, and three outgroup taxa under weighting regimes that maximized fit and minimized length (Prendini et al., 2009). Unambiguous synapomorphies are indicated with bars. Solid bars indicate uniquely derived apomorphic states, whereas empty bars indicate parallel derivations of apomorphic states. The number above each bar gives the character number, the number below gives the character state.

Fig. 5. Typhlochactidae Mitchell, 1971, chelicerae, dorsal aspect. A. Alacran tartarus Francke, 1982, paratype ³ (AMNH). B. Sotanochactas elliotti (Mitchell, 1971), paratype ³ (WDS). C. Stygochactas granulosus (Sissom and Cokendolpher, 1998), juv. ³ holotype (AMNH). D. Typhlochactas cavicola Francke, 1986, holotype ♀ (AMNH). E. Typhlochactas mitchelli Sissom, 1988, holotype ³ (AMNH). F. Typhlochactas reddelli Mitchell, 1968, holotype ♀ (AMNH). G. Typhlochactas rhodesi Mitchell, 1968, holotype ♀ (AMNH). H. Typhlochactas sissomi Francke et al., 2009, subad. ³ holotype (IBUNAM), dextral chelicera. I. Typhlochactas sylvestris Mitchell and Peck, 1977, holotype ♀ (AMNH), sinistral chelicera. Scales: A 5 1 mm; B–H 5 0.1 mm.

Fig. 8. Typhlochactidae Mitchell, 1971, sternum, genital operculum and pectines, ventral aspect. A. Stygochactas granulosus (Sissom and Cokendolpher, 1998), juv. ³ holotype (AMNH). B. Typhlochactas mitchelli Sissom, 1988, holotype ³ (AMNH). C. Typhlochactas cavicola Francke, 1986, holotype ♀ (AMNH). D. Typhlochactas sylvestris Mitchell and Peck, 1977, holotype ♀ (AMNH). Scales 5 0.5 mm.

Fig. 9. Typhlochactidae Mitchell, 1971, dextral pedipalp chela fixed (left) and movable (right) fingers, dentition patterns. A, B. Alacran tartarus Francke, 1982. C, D. Sotanochactas elliotti (Mitchell, 1971). E, F. Stygochactas granulosus (Sissom and Cokendolpher, 1998). G, H. Typhlochactas cavicola Francke, 1986. I, J. Typhlochactas mitchelli Sissom, 1988. K, L. Typhlochactas reddelli Mitchell, 1968. M, N. Typhlochactas rhodesi Mitchell, 1968. O, P. Typhlochactas sissomi Francke et al., 2009. Q, R. Typhlochactas sylvestris Mitchell and Peck, 1977.

Fig. 12. Typhlochactinae Mitchell, 1971, metasoma and telson, lateral aspect. A. Sotanochactas elliotti (Mitchell, 1971), paratype ³ (WDS). B. Stygochactas granulosus (Sissom and Cokendolpher, 1998), juv. ³ holotype (AMNH). C. Typhlochactas cavicola Francke, 1986, holotype ♀ (AMNH). D. Typhlochactas mitchelli Sissom, 1988, holotype ³ (AMNH). E. Typhlochactas reddelli Mitchell, 1968, holotype ♀ (AMNH), segments IV, V and telson only. F. Typhlochactas rhodesi Mitchell, 1971, holotype ♀ (AMNH). G. Typhlochactas sylvestris Mitchell and Peck, 1977, holotype ♀ (AMNH). Scales 5 1 mm.

Fig. 23. Typhlochactas cavicola Francke, 1986, holotype ♀ (AMNH), habitus. A. Dorsal aspect. B. Ventral aspect. Scale 5 1 mm.

Fig. 24. Typhlochactas cavicola Francke, 1986, holotype ♀ (AMNH), sinistral pedipalp chela showing carinae and distribution of trichobothria. A. Dorsal aspect. B. External aspect. C. Ventral aspect. Scale 5 0.5 mm.

Fig. 25. Typhlochactas cavicola Francke, 1986, holotype ♀ (AMNH), sinistral pedipalp femur (A) and patella (B–D) showing carinae and distribution of trichobothria. A, B. Dorsal aspect. C. External aspect. D. Ventral aspect. Scales 5 0.5 mm.