Diplocentrus keyserlingii Karsch, 1880

Santibäñez-López, Carlos E., Francke, Oscar F. & Prendini, Lorenzo, 2013, Systematics of the keyserlingii group of Diplocentrus Peters, 1861 (Scorpiones: Diplocentridae), with descriptions of three new species from Oaxaca, Mexico, American Museum Novitates 2013 (3777), pp. 1-48 : 20-28

Diplocentrus keyserlingii Karsch, 1880

Figure 5A ; tables 1, 2

Diplocentrus keyserlingii Karsch, 1880: 57 ; Stahnke, 1981: 34–44; Sissom, 1991: 156; Sissom and Walker, 1992: 130; Sissom, 1994: 257–261; Kovařík, 1998: 130; Sissom and Jackman, 1997: 151; Beutelspacher and Trujillo-Olvera, 1999: 9 (part); Beutelspacher, 2000: 29; Sissom and Fet, 2000: 338; Stockwell and Baldwin, 2001: 304 (part); Armas and Martín-Frías, 2003: 74.

Diplocentrus whitei: Kraepelin, 1894: 13–15 (part, misidentification).

Diplocentrus keyserlingi: Kraepelin, 1899: 102 (part); Banks, 1910: 185, 188; Herrera, 1917: 270; 1921: 159, figs. 30–33 (part); Ewing, 1928: 5; Gertsch, 1939: 17; Pelaez, 1962: 72 (part); Stahnke, 1970a: 25; Bücherl, 1971: 324; Guijosa, 1973: 145; Rowland and Reddell, 1976: 5; Moritz and Fischer, 1980: 316; Francke and Ponce-Saavedra, 2005: 49, 52.

Diplocentrus keyserlingi keyserlingi (not conspecific): Hoffmann, 1931: 313–317; 1938: 317.

Didymocentrus keyserlingii: Werner, 1934: 275 .

Diplocentrus moritzi: Stahnke, 1981: 44 , figs. 3, 4 (synonymized by Sissom, 1991: 156).

Diplocentrus reticulatus Francke, 1977: 150 , 191–194, 198, figs. 8, 13, 24, 55–58 (synonymized by Sissom, 1991: 156); Beutelspacher and Trujillo-Olvera, 1999: 8; Beutelspacher, 2000: 33.

TYPE MATERIAL: MEXICO: OAXACA: Lectotype ♂, paralectotype ♂ ( ZMB 3248 ) [designated by Stahnke, 1981: 34; not examined]. DISTRITO DE ETLA: Municipio de Santiago Tenango : Holotype ♂, paratype ♂ [ Diplocentrus reticulatus ] ( AMNH), near Tejocote [17°14′N 97°00′W], 7800 ft, summer 1963, C.M. Bogert, under logs . DISTRITO DE IXTLÁN DE JUÁREZ: Municipio de Santa Catarina Ixtepeji : Paratype ♂ [ Diplocentrus reticulatus ] ( AMNH), El Punto, road to Ixtlán de Juarez [17°13.30′N 96°35.03′W], 19.viii.1961, C.M. and M. R. Bogert .

ADDITIONAL MATERIAL: MEXICO: OAXACA: DISTRITO DE ETLA: Municipio de Santa María Peñoles: Santa Catarina Estetla , 17°01.6′N 97°05.766′W, 20.ix.2007, P. Lara, 1 ♂ ( CALA) . DISTRITO DE ZAACHILA: Municipio de Santa Inés del Monte: Santa Inés del Monte , 3 km E, 16°56.445′N 96°51.6312′W, 2665 m, 12.xii.2005, O. Francke, H. Montaño, C. Santibáñez, and A. Valdez, 1 ♂, 1 ♀ ( CNAN) , 2 ♀ ( AMNH) , 1 ♀ ( AMCC [ LP 6517 ]) ; Santa Inés del Monte , 16°56.442′N 96°51.629′W, 2270 m, 19.ix.2009, R. Paredes, C. Santibáñez, A. Valdez, and J. Cruz, 1 ♂, 5 juv. ( CNAN) , 1 juv. [leg] ( AMCC [ LP 11052 ]) .

DIAGNOSIS: The following character combination, updated from Francke (1977), is diagnostic for D. keyserlingii . Total length (adult), ca. 45 mm. Base coloration (adult) reddish brown to brown. Carapacial anteromedian notch moderately deep, V-shaped; frontal lobes and interocular surface moderately granular, other surfaces finely granular. Pedipalp femur, dorsal surface moderately granular. Pedipalp patella, dorsomedian carina strongly developed, crenulate to smooth (♂); dorsoexternal carina moderately developed, smooth to slightly crenulate (♂); externomedian carina weakly developed to obsolete, smooth (♂); ventromedian carina obsolete to weakly developed, slightly granular (♂, ♀). Pedipalp chela manus, dorsal surface markedly reticulate (♂; fig. 5A) or smooth (♀); digital carina strongly developed, smooth (♂) or obsolete, smooth (♀); dorsal secondary carina moderately developed, smooth to slightly crenulate; fingers gently curved (fig. 5A). Legs I–IV telotarsi, counts of spiniform macrosetae in pro- and retroventral rows, 4/5:5/5:5/5:5/6 (variation in table 1). Pectinal tooth count, 7–9, mode = 7 (♂) or 6–8, mode = 7 (♀) (variation in table 2).

Diplocentrus keyserlingii resembles D. kraepelini , n. sp., D. rectimanus , and D. sagittipalpus , n. sp., in adult size and coloration but can be distinguished as follows. The counts of spiniform macrosetae on the telotarsi of legs III and IV are lower in D. keyserlingii (5/5:5/6) than in D. kraepelini , n. sp. (6/6:6/6), D. rectimanus (6/6:6/6), and D. sagittipalpus , n. sp. (6/6:6/6). The pectinal tooth count (♂) is lower in D. keyserlingii (7–9) than in D. sagittipalpus , n. sp. (9–12). The pedipalp chela manus (♂) is rounded, the fingers shorter than the manus, in D. keyserlingii , but slender, the fingers longer than the manus, in D. rectimanus , and the manus, dorsal surface (♀) is smooth in D. keyserlingii but granular in D. rectimanus .

DISTRIBUTION: Diplocentrus keyserlingii is known only from the central valleys and adjacent mountains (part of the Sierra Norte) of Oaxaca in the following municipalities: Santa Catarina Ixtepeji, Santa Inés del Monte, Santa María Peñoles, and Santiago Tenango (fig. 3B).

ECOLOGY: This species was collected under logs and stones, and excavated from burrows constructed at an angle of ca. 20° to the ground surface, ca. 20 cm deep and fairly straight with some turns due to the presence of stones in the soil matrix. The dominant vegetation in the area was oak forest. Centruroides nigrovariatus ( Pocock, 1898) was collected in sympatry. The habitat and habitus of D. keyserlingii are consistent with the pelophilous ecomorphotype ( Prendini, 2001).

Diplocentrus kraepelini , n. sp.

Figures 7B , 8B , 9B , 10B , 11B , 14 , 15 ; tables 1–3

TYPE MATERIAL: MEXICO: OAXACA: DISTRITO DE COIXTLAHUACA: Municipio de San Cristóbal Suchixtlahuaca: Holotype ♂ ( CNAN-T0671 ), 2 ♂, 3 ♀, 2 subad. ♀, 2 juv. paratypes ( AMNH), 1 ♂, 3 ♀, 1 subad. ♂, 2 juv. paratypes ( CNAN-T0672 ), 1 ♀ paratype ( AMCC [ LP 6426 ]), Km 2 road San Cristóbal Suchixtlahuaca–Santiago Tejupan , 17°42.240′N 97°23.667′W, 2290 m, 28.vi.2006, O. Francke, G. Villegas, H. Montaño, and A. Valdez; 1 subad. ♀ paratype ( CNAN), 1 subad. ♀ paratype [leg] ( AMCC [ LP 10973 ]), Suchixtlahuaca, 8 km NE, 17°42.124′N 97°23.776′W, 2030 m, 25.iii.2010, O. Francke, A. Valdez, C. Santibáñez, and J. Cruz, oak forest, under rock, daytime rock rolling.

ADDITIONAL MATERIAL: MEXICO: OAXACA: DISTRITO DE TEPOSCOLULA: Municipio San Bartolo Soyaltepec: Caballo Blanco [17°35.432′N 97°18.414′W], 12.vii.1963, G. Sludder, under logs, 1 ♂ ( AMNH) .

ETYMOLOGY: This species is dedicated to Karl Kraepelin for his contributions to scorpiology.

DIAGNOSIS: The following character combination is diagnostic for D. kraepelini , n. sp. Total length (adult), 50–55 mm. Base coloration (adult) brown to orange-brown. Carapace anteromedian notch moderately deep, U-shaped (fig. 7B). Pedipalp femur, dorsal surface finely and sparsely granular medially (fig. 9B). Pedipalp patella, dorsomedian carina moderately developed, crenulate (♂); dorsoexternal carina weakly developed to obsolete, smooth to slightly crenulate (♂); externomedian carina weakly developed, smooth (♂; fig. 11B); ventromedian carina moderately developed, smooth to slightly crenulate (♂, ♀). Pedipalp chela manus, dorsal surface markedly reticulate (♂; fig. 14A) or smooth (♀; fig. 14B); digital carina strongly developed, smooth to crenulate (♂) or weakly developed to obsolete, smooth (♀); dorsal secondary carina moderately to weakly developed, smooth (♂) or weakly developed to obsolete, smooth (♀); fingers markedly curved. Legs I–IV telotarsi, counts of spiniform macrosetae in pro- and retroventral rows, 4/5:5/5:6/6:6/6 (variation in table 2). Pectinal tooth count, 9–11, mode = 10 (♂) or 7–9, mode = 8 (♀) (variation in table 1).

Diplocentrus kraepelini , n. sp., D. keyserlingii and D. rectimanus possess similar counts of pectinal teeth and telotarsal spiniform macrosetae, but may be distinguished as follows. The pedipalp chela movable finger length is greater than the length of the carapace or metasomal segment V in D. kraepelini , n. sp., and D. rectimanus , but less than or subequal to the length of the carapace or metasomal segment V in D. keyserlingii . The counts of spiniform macrosetae on the telotarsi of legs I and II are lower (4/5) in D. kraepelini , n. sp., than in D. rectimanus (5/5), whereas the counts on the telotarsi of legs III and IV are higher (6/6:6/6) in D. kraepelini , n. sp., than in D. keyserlingii (5/5:5/6). The ventromedian carina is moderately developed, smooth to slightly crenulate in D. kraepelini , n. sp., but obsolete in D. rectimanus . The pedipalp chela fingers are markedly curved in D. kraepelini , n. sp., but straight to gently curved in D. rectimanus . The pedipalp chela manus, dorsal surface (♀) is smooth in D. kraepelini , n. sp., but granular in D. rectimanus .

Diplocentrus kraepelini , n. sp., resembles D. mitlae in size and coloration, but may be distinguished from the latter as follows. The pedipalp chela manus (♂), dorsal and external surfaces are reticulate and the digital carina strongly developed in D. kraepelini , n. sp., whereas the dorsal and external surfaces are smooth and the digital carina weakly developed in D. mitlae . The count of spiniform macrosetae on the telotarsi of legs I and II is higher in D. kraepelin i, n. sp. (6/6), than in D. mitlae (5/5).

DESCRIPTION: Based on holotype ♂ and paratype ♂ (fig. 15A, B) with differences in paratype ♀ (fig. 15C, D) noted. Measurements in table 3.

Coloration: Carapace brown to orange; moderately infuscate throughout, uniformly so around median ocelli, variegated elsewhere. Coxosternum pale orange. Pedipalps orange to reddish brown, carinae darker. Legs pale brown to pale orange, uniformly infuscate. Mesosoma brown; tergites brown (♂) or pale brown (♀), densely infuscate; sternites pale orange. Metasoma pale orange to brown; carinae weakly to moderately infuscate. Telson orange to reddish, uniformly infuscate.

Carapace: Anterior margin moderately setose; anteromedian notch moderately deep, U-shaped (fig. 7B). Frontal lobes and interocular surface moderately granular; other surfaces sparsely granular or shagreened. Three pairs of subequal lateral ocelli.

Pedipalps: Orthobothriotaxic, type C. Femur height greater than width (fig. 10B); dorsal intercarinal surface shallowly convex, finely and sparsely granular medially; external intercarinal surface smooth; ventral intercarinal surface flat, smooth; internal intercarinal surface coarsely and densely granular; dorsointernal carina strongly developed, granular; dorsoexternal carina weakly developed, granular proximally and smooth distally; ventroexternal carina obsolete to weakly developed; ventrointernal carina moderately developed, granular proximally, becoming obsolete distally. Patella, dorsal intercarinal surfaces slightly reticulate (♂) or smooth (♀); external and ventral intercarinal surfaces slightly reticulate (fig. 11B); internal intercarinal surface smooth; proximal tubercle moderately developed, bifurcate; dorsointernal carina weakly developed to obsolete; dorsomedian carina moderately developed, crenulate (♂) or weakly developed, smooth (♀); dorsoexternal carina weakly developed, smooth to slightly crenulate (♂) or obsolete, smooth (♀); externomedian weakly developed, smooth; ventroexternal carinae weakly to moderately developed, smooth; ventromedian carina moderately developed, smooth to slightly crenulate; ventrointernal carina strongly developed, comprising large granules. Chela manus, slender, height subequal to width (♂) or rounded, height greater than width (♀), densely (♂) or sparsely (♀) setose; dorsal intercarinal surface moderately reticulate (♂; fig. 14A) or smooth (♀; fig. 14B); external intercarinal surface smooth; dorsal marginal carina moderately developed, coarsely granular; digital carina strongly developed (♂) or obsolete to weakly developed (♀), smooth; dorsal secondary and external secondary carinae weakly to moderately developed (♂) or obsolete to weakly developed (♀), smooth; ventroexternal carina weakly developed, smooth distally, becoming obsolete proximally; ventromedian carina strongly developed, smooth proximally, becoming obsolete distally, directed toward midpoint of movable finger articulation; ventrointernal carina weakly to moderately developed, smooth; internodorsal carina weakly developed, granular; internomedian and internoventral carinae weakly developed, smooth; internal surface with shallow longitudinal depression where chela rests against patella. Chela fixed finger curved; length equal (♂) or subequal (♀) to femur length and patella length; dorsal surface smooth and densely setose proximally; external surface flat; internal surface shallowly concave.

Legs: Legs I–IV femora and tibiae, prolateral surfaces shagreened; telotarsi, counts of spiniform macrosetae in pro- and retroventral rows (dextral/sinistral), 4/5 4/5: 5/5 5/5: 6/6 6/6: 6/6 6/6.

Pectines: Tooth count: 10–10 (♂; fig. 8B); 8–9 (♀).

Mesosoma: Tergites I –VI, pretergites smooth, posttergites granular; VII granular. Sternites smooth; VII, dorsosubmedian and dorsolateral carinae weakly developed, crenulate.

Metasoma: Metasomal segments I–V, dorsal intercarinal surfaces weakly reticulate on segments I–IV, smooth on V; lateral intercarinal surfaces reticulate; ventral intercarinal surfaces smooth on I–III, sparsely granular on IV and V. Segments I–IV, dorsolateral carinae moderately developed (♂) to weakly developed (♀), granular; lateral supramedian carinae weakly to moderately developed on I, moderately developed, slightly granular to crenulate on II–IV; lateral inframedian carinae moderately developed, slightly granular to crenulate on I and II, weakly to moderately developed, slightly granular on III and IV; ventrolateral carinae moderately developed, crenulate to smooth on I–III, weakly developed, slightly granular on IV; ventrosubmedian carinae moderately developed, slightly crenulate to smooth on I–III, weakly developed, slightly crenulate to smooth on IV. Segment V length:pedipalp femur length ratio, 1.27 (♂), 1.39 (♀); dorsolateral and lateral inframedian carinae weakly to moderately developed, crenulate; ventrolateral carinae moderately developed, crenulate; ventromedian carina moderately developed, granular, with subspiniform granules posteriorly; ventral transverse carina moderately developed, comprising six subspiniform granules; anal arch semicircular; anal subterminal carina strongly developed, comprising 10 subspiniform granules; anal terminal carina moderately developed, granular.

Telson: Telson , width:length ratio, 0.56 (♂), 0.70 (♀). Vesicle, lateral surfaces smooth; ventral surface granular anteriorly. Subaculear tubercle stout, subconical. Aculeus, 1.5 length of subaculear tubercle, strongly curved.

Hemispermatophore: Lamelliform, weakly sclerotized (fig. 9B); total length, 6.5 mm; distal lamella, length, 3.5 mm; capsular region, width, 1.6 mm; median lobe narrow, margin entire.

DISTRIBUTION: Diplocentrus kraepelini , n. sp., is known only from the type locality in the San Cristóbal Suchixtlahuaca municipality of Oaxaca (fig. 3A) .

ECOLOGY: This species was observed doorkeeping at burrow entrances at night with UV detection, and was excavated during the daytime. The burrows, constructed at an angle of ca. 30° to the ground surface, were ca. 30 cm long, and mostly straight with some turns around stones in the soil matrix. The dominant vegetation in the area was oak forest. An undetermined species of Centruroides Marx, 1890 , related to C. nigrovariatus , Vaejovis dzahui Santibáñez- López and Francke, 2010, and an undetermined species of Vaejovis were collected in sympatry. The habitat and habitus of D. kraepelini , n. sp., are consistent with the pelophilous ecomorphotype ( Prendini, 2001).

Figures

FIGURE 5. Diplocentrus keyserlingii group, dextral pedipalp chela, dorsoexternal aspect. A. Diplocentrus keyserlingii Karsch, 1880, holotype ♂ (AMNH) of Diplocentrus reticulatus Francke, 1977. B. Diplocentrus mitlae Francke, 1977, holotype ♂ (AMNH). Scale bar = 1 mm.

FIGURE 7. Diplocentrus keyserlingii group, carapace, dorsal aspect. A. Diplocentrus formosus Armas and Martín-Frías, 2003, ♂ (CNAN). B. Diplocentrus kraepelini, n. sp., paratype ♂ (AMNH). C. Diplocentrus sagittipalpus, n. sp., paratype ♂ (AMNH). D. Diplocentrus sissomi, n. sp., paratype ♂ (AMNH). Scale bars = 1 mm.

FIGURE 8. Diplocentrus keyserlingii group, sternum, genital operculum, and pectines, ventral aspect. A. Diplocentrus formosus Armas and Martín-Frías, 2003, ♂ (CNAN). B. Diplocentrus kraepelini, n. sp., paratype ♂ (AMNH). C. Diplocentrus sagittipalpus, n. sp., paratype ♂ (AMNH). D. Diplocentrus sissomi, n. sp., paratype ♂ (AMNH). Scale bars = 1 mm.

FIGURE 9. Diplocentrus keyserlingii group, sinistral hemispermatophore, dorsal aspect. A. Diplocentrus formosus Armas and Martín-Frías, 2003, ♂ (CNAN). B. Diplocentrus kraepelini, n. sp., paratype ♂ (AMNH). C. Diplocentrus sagittipalpus, n. sp., paratype ♂ (AMNH).D. Diplocentrus sissomi, n. sp., paratype ♂ (AMNH). Scale bars = 1 mm.

FIGURE 10. Diplocentrus keyserlingii group, dextral pedipalp femur, dorsal aspect. A. Diplocentrus formosus Armas and Martín-Frías, 2003, ♂ (CNAN). B. Diplocentrus kraepelini, n. sp., paratype ♂ (AMNH). C. Diplocentrus sagittipalpus, n. sp., paratype ♂ (AMNH). D. Diplocentrus sissomi, n. sp., paratype ♂ (AMNH). Scale bars = 1 mm.

FIGURE 11. Diplocentrus keyserlingii group, dextral pedipalp patella, external aspect. A. Diplocentrus formosus Armas and Martín-Frías, 2003, ♂ (CNAN). B. Diplocentrus kraepelini, n. sp., paratype ♂ (AMNH). C. Diplocentrus sagittipalpus, n. sp., paratype ♂ (AMNH). D. Diplocentrus sissomi, n. sp., paratype ♂ (AMNH). Scale bars = 1 mm.

FIGURE 14. Diplocentrus kraepelini, n. sp., dextral pedipalp chela, dorsoexternal aspect. A. Paratype ♂ (AMNH). B. Paratype ♀ (AMNH). Scale bar = 1 mm.

FIGURE 15. Diplocentrus kraepelini, n. sp., habitus, dorsal (A, C) and ventral (B, D) aspect. A, B. Paratype ♂ (AMNH). C, D. Paratype ♀ (AMNH). Scale bars = 10 mm.