Kochius Soleglad and Fet, 2008

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 : 29-30

Kochius Soleglad and Fet, 2008

Figures 1B , 2B , 5 , 7 , 9–11 , 16C , 18A, B , 22A , 23A , 24A , 25A ; table 1

Buthus punctipalpi Wood, 1983 [= Kochius punctipalpi (Wood, 1983) ], type species, by original designation.

Vaejovis punctipalpi group (part): Williams, 1971a: 37; 1971b: 44; 1980: 80; Sissom and Francke, 1985: 1; Sissom, 1989: 180; 1991: 26; Stockwell, 1992: 408, 409; Sissom, 1993: 68; Lourenço and Sissom, 2000: 135; Sissom, 2000: 532, 548, 551; Armas and Martín-Frías, 2001: 1; Hendrixson, 2001: 1; Ponce-Saavedra and Sissom, 2004: 539, 541; Sissom and Hendrixson, 2005a: 131; 2005b: 35; Soleglad and Fet, 2005: 4, 6, 7; Graham and Fet, 2006: 7, 8, 10; Fet et al., 2006a: 9, table 1 ; 2006b: 8; 2006c: 7; Soleglad and Fet, 2006: 6, 26, tables 2 , 3 ; Graham and Soleglad, 2007: 11, 12; Francke and González-Santillán, 2007: 586, 587, 590; Soleglad et al., 2007: 134, 135; Soleglad and Fet, 2008: 1, 26, 57, 60, 94, 95, 102; McWest, 2009: figs. 322–326, tables 1 , 53, 89, 91, 93, 101– 105, 108, 123.

Kochius Soleglad and Fet, 2008: 1 , 3, 34, 38, 51, 53, 54, 57, 58, 60–69, 71, 73, 74, 76, 79, 91–96, 102, 104, figs. 192, 193–196, 204, tables 2 , 4, 6, 9 (part); Ayrey and Soleglad, 2011: 1 (part).

DIAGNOSIS: Kochius differs from all other genera of Syntropinae in possessing the longest macrosetae on the telson vesicle of the female and, in some species, of both sexes (fig. 25A); some macrosetae are as long as or longer than the aculeus, becoming narrower and translucent terminally. Species of Kochius are also characterized by a raised, whitish boss, restricted to the posteromedial third of mesosomal sternite VII (fig. 23A). Although other genera of Syntropinae possess whitish glandular surfaces on sternite VII (fig. 23B), the structure in Kochius , which is noticeably more elevated and usually shaped like a truncate triangle, is unique. Most Syntropinae also display a glandular surface posteromedially on sternite V, but it is more pronounced in Kochius (fig. 23A). Additional diagnostic characters of Kochius are as follows. Species of Kochius possess more pronounced, granular carinae on the pedipalp chela manus, especially the drs, drsa, and rlm carinae (fig. 18A, B), and the pedipalp patellar rlds and rlm carinae are more coarsely granular than in other genera of Syntropinae . The pedipalp carinae of Kochius are also distinctive in comprising moniliform granules (fig. 18A, B), rather than clustered granules, as in other genera (fig. 17A–D). The patellar dorsal intercarinal surface (between the dpl and drl carinae) is flat in Kochius , compared to other genera in which it is slightly concave. The densely and coarsely granular (shagreened) carapace (fig. 16C), markedly shagreened mesosomal posttergites I–VII and sternite VII (fig. 23A), and uniformly finely granular (matte) ventral intercarinal surfaces of the pedipalp chela manus, retrolateral intercarinal surfaces of the pedipalp patella and intercarinal surfaces of metasomal segments I–V (fig. 24A) are also characteristic for the genus. Pedipalp chela trichobothrium esb is situated at RD6 and/or its associated macroseta, and trichobothrium Db is situated dorsal to the drl carina, compared with Paravaejovis , in which esb is situated between RD6 and RD5 (fig. 19D), and Mesomexovis , gen. nov., in which Db is situated on the drl carina (fig. 19A). The macrosetae of legs I–IV also tend to be shorter and stouter in Kochius than in other genera (fig. 22A).

Four species of Chihuahuanus , gen. nov. ( C. cazieri , comb. nov., C. crassimanus , comb. nov., C. kovariki , comb. nov., and C. russelli , comb. nov.) resemble Kochius in possessing a raised, whitish boss in the posteromedial third of mesosomal sternite VII (fig. 23A), but may be separated from the latter by the small fusiform, whitish glandular area, anterior to the base of the aculeus, on the dorsal surface of the telson vesicle of adults; the shorter macrosetae on the telson vesicle of females; the less-developed (lower and smoother) pedipalp chelal rlm and rlvs carinae, and patellar rlds and rlm carinae; and the clustered instead of moniliform granules of the pedipalp carinae.

Species of Thorellius resemble Kochius with respect to the incrassate pedipalp chela manus and generally pronounced carination of the manus, especially the markedly granular rlm and rlvs carinae (figs. 17C, D, 18A, B), and metasoma, but can be separated from the latter by their considerably larger size and by the clustered instead of moniliform granules of the pedipalp carinae.

INCLUDED SPECIES: Kochius barbatus (Williams, 1971) , stat. nov.; Kochius bruneus (Williams, 1970) ; Kochius cerralvensis (Williams, 1971) , stat. nov.; Kochius insularis (Williams, 1971) ; Kochius hirsuticauda (Banks, 1910) ; Kochius magdalensis (Williams, 1971) ; Kochius punctipalpi (Wood, 1863) ; Kochius sonorae (Williams, 1971) ; Kochius villosus (Williams, 1971) , stat. nov.

DISTRIBUTION: Kochius is endemic to Mexico (recorded from Baja California, Baja California Sur, and Sonora) and the United States (recorded from Arizona, California, and Nevada; fig. 5). All except K. hirsuticauda and K. sonorae , occurring in the Mojave and Sonoran (including Colorado) deserts on the North American mainland, are endemic to the Baja California Peninsula and adjacent islands.

NATURAL HISTORY: Species of Kochius inhabit desert to semidesert habitats from sea level to 1350 m altitude. They have been collected under stones or other available debris on the ground, and on the surface at night, with UV light detection. Although Sissom (2000) suggested that these scorpions are burrowers, the absence of fossorial adaptations suggests otherwise. The cheliceral fingers are not enlarged, the pedipalp chela fingers are fairly slender, the tarsi are not laterally compressed and lack setal combs, and the metasoma lacks enlarged carinal processes and lobes. The habitat and habitus of Kochius are consistent with the lapidicolous ecomorphotype ( Prendini, 2001a).

REMARKS: As redefined here, this genus accommodates species previously assigned only to Williams’ (1971a) punctipalpi group of Vaejovis , for which Soleglad and Fet (2008) devised the name Kochius , without quantitatively testing its monophyly or composition. Kochius , as defined by Soleglad and Fet (2008), was consistently polyphyletic, and the group of species hereby assigned to it consistently monophyletic, in the phylogenetic analyses of González-Santillán and Prendini (in press) based on morphology and those based on morphology and DNA. The following species are therefore transferred to other genera in the present contribution: Mesomexovis atenango , comb. nov., previously assigned to the punctipalpi group ( Francke and González-Santillán, 2007) and then to Kochius ( Soleglad and Fet, 2008) ; C. cazieri , comb. nov., C. crassimanus , comb. nov., and C. russelli , comb. nov., previously assigned to the punctipalpi group ( Williams, 1971a, b; Sissom, 2000) and then to Kochius ( Soleglad and Fet, 2008) ; C. kovariki , comb. nov., previously assigned to Kochius ( Soleglad and Fet, 2008) .

The analyses of González-Santillán and Prendini (in press) identified sufficient, consistent diagnostic character differences to elevate to the rank of species, K. barbatus , stat. nov., and K. cerralvensis , stat. nov., previously considered subspecies of K. punctipalpi , and K. villosus , stat. nov., previously considered a subspecies of K. bruneus ( Williams, 1971a, 1980; Sissom, 2000). Vaejovis bruneus loretoensis Williams, 1971 , was not found to differ consistently from K. villosus , stat. nov., with which it is hereby synonymized: Vaejovis bruneus loretoensis Williams, 1971 = Kochius villosus (Williams, 1971) , syn. nov.

MATERIAL EXAMINED: Kochius barbatus (Williams, 1971) , stat. nov.: MEXICO: Baja California Sur: Municipio de La Paz : Las Cruces , 25 ft, 29.vii.1968, S.C. Williams and M.A. Cazier, holotype ♀ of Vaejovis punctipalpi barbatus Williams, 1971 ( CAS Type No. 10469), paratype ♀ ( AMNH); Las Cruces, 4 km NW, 24 ° 13.5108 ′ N 110 ° 7.41 ′ W, 31 m, 24.vi.2008, H. Montaño and E. González, 18 ( AMNH [ ARA 2884 ]), 18 ( IBUNAM [ ARA 2274 ]). Kochius bruneus (Williams, 1970) : MEXICO: Baja California Sur: Municipio de Comondú : San José de Comondú , 5 mi. SW, 1000 ft, 2.vii.1968, S.C. Williams and M.A. Cazier, holotype ♂ ( CAS Type No. 10410); San José de Comondú, 30 km N, between Rosarito and San Isidro , 26 ° 21.2520 ′ N 111 ° 50.10 ′ W, 258 m, 27.vi.2008, H. Montaño and E. González, 28, 2♀ ( AMNH [ ARA 2854 ]). Kochius cerralvensis (Williams, 1971) , stat. nov.: MEXICO: Baja California Sur: Municipio de La Paz : Isla Cerralvo , Bahía Limona , 31.v.1969, S.C. Williams, holotype ♂ ( CAS Type No. 11470); Isla Cerralvo, Piedras Gordas , 17.v.1970, S.C. Williams and V. F. Lee, 18, 1♀ paratypes ( AMNH); Isla Cerralvo , SW side, 24 ° 10.9332 ′ N 109 ° 53.2332 ′ W, 50 m, 27.v.2008, E. González and I.G. Nieto, 18, 1♀ ( AMNH [ ARA 2880 ]). Kochius hirsuticauda (Banks, 1910) : U.S.A.: Arizona: Yuma Co. : Palm Canyon, Kofa Mountains , 12.x.1968, M.A. Cazier et al., 18, 1♀ ( AMNH). California: San Diego Co.: Anza-Borrego Desert State Park: Pinyon Mountain Road , ca. 4.8 km towards Pinyon Mountain Valley from junction with Route S2, 33 ° 03.043 ′ N 116 ° 23.270 ′ W, 870 m, 30.viii.2005, R. Mercurio and L. Prendini , 18, 1♀ ( AMNH). Kochius insularis (Williams, 1970) : MEXICO: Baja California Sur: Municipio de La Paz : Isla Partida , Central Valley , 10.vii.1968, S.C. Williams, M. Bentzien, and B. Fox, holotype ♂ ( CAS Type No. 10422), allotype ♀ ( CAS). Municipio de La Paz : Isla Espíritu Santo, Playa Bonanza , 24 ° 27.3636 ′ N 110 ° 18.47 ′ W, 50 m, 31.v.2008, E. González and I.G. Nieto, 18, 1♀ ( AMNH [ ARA 2875 ]), 18, 1♀ ( IBUNAM [ ARA 2260 ]). Kochius magdalensis (Williams, 1971) : MEXICO: Baja California Sur: Municipio de La Paz : La Paz , 75 mi. NW, 200 ft, 4.vii.1968, S.C. Williams and M.A. Cazier, holotype ♂ ( CAS Type No. 10468); El Pilar, ca. 20 km NE Las Pocitas , 24 ° 28.7616 ′ N 111 ° 1.12 ′ W, 92 m, 25.vi.2008, H. Montaño and E. González, 18, 1♀ ( AMNH [ ARA 2874 ]), 18, 1♀ ( IBUNAM [ ARA 2259 ]); Junction Route 1 and road to Juan de la Costa , 5 km N, 24 ° 08.500 ′ N 110 ° 28.566 ′ W, 30 m, 9.vi.2005, E. González, W.E. Savary, R. Mercurio, and L. Prendini , 18 ( AMNH). Kochius punctipalpi (Wood, 1863) : MEXICO: Baja California Sur: Municipio de Los Cabos: Santiago , 23 ° 26.4 ′ N 109 ° 43.5666 ′ W, 225 m, 9.vii.2004, O.F. Francke, E. González, and A. Valdez, 18, 1♀ ( AMNH [ ARA 1471 ]); San José del Cabo, ca. 10 km S off Route 1, 23 ° 1.7622 ′ N 109 ° 43.49 ′ W, 50 m, 10.vii.2005, W.E. Savary, E. González, and R. Mercurio , 18, 1♀ ( AMNH [ ARA 1473 ]). Municipio de La Paz: La Paz , 15 km SE, 24 ° 3.7332 ′ N 110 ° 10.53 ′ W, 476 m, 8.vii.2004, O.F. Francke, E. González, and A. Valdez, 18 ( AMNH [ ARA 1472 ]). Kochius sonorae (Williams, 1971) : MEXICO: Sonora: Municipio de Alamos : Alamos , 11.2 mi. W, 23.i.1966, R. Winokur, C. Mays , and M.A. Nikerson, holotype ♂ ( CAS Type No. 10472). Municipio de Navojoa: Microwave antenna Cerro Prieto , 15 km E Navojoa , 27 ° 4.9830 ′ N 109 ° 17.4498 ′ W, 375 m, 23.viii.2009, E. González and J.L. Castelo, 18, 1♀ ( AMNH [ ARA 3157 ]); Navojoa, 32 km S, 12.ix.1966, J. and W. Ivie, 18, 1♀ ( AMNH). Municipio de Soyopa: Sierra el Encinal, 9 km from Hwy MEX 16 on dirt road to El Encinal , 28 ° 35.4120 ′ N 109 ° 27.1476 ′ W, 380 m, 24.vi. 2006, E. González and P. Berea, 28 ( AMNH [ ARA 2539 ]); dirt road between El Encinal and Tonichi , 28 ° 33.2388 ′ N 109 ° 21.64 ′ W, 645 m, 1.vii.2006, E. González and P. Berea, 28 ( AMNH [ ARA 3015 ]). Kochius villosus (Williams, 1971) , stat. nov.: MEXICO: Baja California Sur: Municipio de Comondú : San Miguel Comondú , 5–10 mi. SW, 800–1000 ft, 3.vii.1968, S.C. Williams and M.A. Cazier, holotype ♂ of Vaejovis bruneus villosus Williams, 1971 ( CAS Type No. 10467), 18, 1♀ paratypes ( AMNH). Municipio de La Paz : San Evaristo , 10–15 km S, dirt road, 24 ° 47.7570 ′ N 110 ° 40.5018 ′ W, 50 m, 12.vii.2008, H. Montaño and E. González, 18 ( IBUNAM [ ARA 2275 ]), 18, 1♀ ( AMNH [ ARA 2888 ]). Municipio de Loreto: Juncalito , 25 ° 49.8918 ′ N 111 ° 19.6908 ′ W, 5 m, 20.v.2004, R. Mercurio and M. Nishiguchi, 18, 1♀ ( AMNH [ ARA 3132 ]). Municipio de Mulegé: San Ignacio , 152 m, 24.vi.1968, S.C. Williams and M.A. Cazier, holotype ♂ of V. b. loretoensis ( CAS Type No. 10466), allotype ♀, 18, 1♀ paratypes ( AMNH).

Figures

Fig. 1. Representative species in the North American vaejovid scorpion subfamily Syntropinae Kraepelin, 1905. A. Chihuahuanus bilineatus (Pocock, 1898), comb. nov., ♀; B. Kochius bruneus (Williams, 1970), 8; C. Mesomexovis punctatus (Karsch, 1879), comb. nov., 8; D. Syntropis macrura Kraepelin, 1900, ♀; E. Thorellius sp., 8; F. Vizcaino viscainensis (Williams, 1970), comb. nov., 8. Photos in B, D and E courtesy of R. Mercurio.

Fig. 2. Representative habitats of species in the North American vaejovid scorpion subfamily Syntropinae Kraepelin, 1905. A. Cuatro Cienegas de Carranza, Coahuila, Mexico, habitat of Chihuahuanus cazieri (Williams, 1968), comb. nov., Chihuahuanus coahuilae (Williams, 1968), comb. nov., and Chihuahuanus globosus (Borelli, 1915), comb. nov. B. Death Valley, California, habitat of Kochius hirsuticauda (Banks, 1910). C. Isla Espiritu Santo, La Paz, Baja California Sur, Mexico, habitat of Kochius insularis (Williams, 1971), Paravaejovis diazi (Williams, 1970), comb. nov., and Paravaejovis gravicaudus (Williams, 1970), comb. nov. D. Cuicatlán, Oaxaca, Mexico, habitat of Mesomexovis subcristatus (Pocock, 1898), comb. nov. E. Pinacate desert, Sonora, Mexico, habitat of Paravaejovis confusus (Stahnke, 1940), comb. nov. F. West of La Paz, Baja California Sur, Mexico, habitat of Syntropis williamsi Soleglad et al., 2007. Photos in B and E courtesy of R. Mercurio and in C courtesy of I.G. Nieto.

Fig. 5. Map of southwestern North America, plotting known locality records for two genera in the vaejovid scorpion subfamily Syntropinae Kraepelin, 1905, based on data collected in the present study: Kochius Soleglad and Fet, 2008 (circles); Thorellius Soleglad and Fet, 2008 (squares).

Fig. 7. Most parsimonious tree (length: 10894.593, CI: 0.190, RI: 0.628, fit: 244.08), obtained by simultaneous phylogenetic analysis of 250 qualitative and quantitative morphological characters and 4221 aligned DNA nucleotides from three mitochondrial and two nuclear gene markers, for 18 genera and 61 species in the North American vaejovid scorpion subfamily Syntropinae Kraepelin, 1905. Analyses were conducted with an implied weighting regime (k 518) that maximized average support (González-Santillán and Prendini, in press). Synapomorphies are optimized with accelerated transformation. Black bars indicate uniquely derived apomorphic states, white bars indicate parallel derivations of apomorphic states, numbers above indicate characters, and numbers below indicate states (appendix 1).

Fig. 9. Kochius punctipalpi (Wood, 1863), 8 (AMNH), dextral pedipalp femur (A–D) and patella (E– H), dorsal (A, E), retrolateral (B, F), ventral (C, G) and prolateral (D, H) aspects, illustrating carinae (boldface) and trichobothria (italics). Abbreviations: dpl, dorsal prolateral; drl, dorsal retrolateral; plm, prolateral median; pls, prolateral subdorsal (absent in Syntropinae Kraepelin, 1905, but present in other vaejovids); pv, prolateral ventral; plvs, prolateral ventrosubmedian; rlds, retrolateral dorsosubmedian; rv, retrolateral ventral; vm, ventral median; vpl, ventral prolateral; vrl, ventral retrolateral; vrs, ventral retrosubmedian. d, dorsal; e, external; eb, external basal; esb, external suprabasal; em, external medial; est, external subterminal; et, external terminal; i, internal; v, ventral. Scale bars 5 1 mm.

Fig. 10. Kochius punctipalpi (Wood, 1863), 8 (AMNH), dextral pedipalp chela, dorsal (A) and retrolateral (B) aspects, illustrating carinae (boldface), finger lobes and notches (uppercase), and trichobothria (italics). Abbreviations: dm, dorsal median; dpl, dorsal prolateral; dps, dorsal prosubmedian; drl, dorsal retrolateral; drs, dorsal retrosubmedian; drsa, retrosubmedian accessory; pld, prolateral dorsal; rd, retrolateral dorsal; rlm, retrolateral median; rls, retrolateral subventral; rlsa, retrolateral subventral accessory; rlvs, retrolateral ventrosubmedian; vm, ventral median; vps, ventral prosubmedian; vrl, ventral retrolateral. ML, medial lobe; MN, medial notch; PN, proximal [5 basal] notch. db, Db dorsal basal; dsb, dorsal suprabasal; dst, dorsal subterminal; dt, Dt, dorsal terminal; eb, Eb, external basal; esb, Esb, external suprabasal; est, Est, external subterminal; et, Et, external terminal. Scale bar 5 1 mm.

Fig. 11. Kochius punctipalpi (Wood, 1863), 8 (AMNH), dextral pedipalp chela, ventral (A) and prolateral (B) aspects, illustrating carinae (boldface), finger lobes and notches (uppercase), and trichobothria (italics). Abbreviations: dpl, dorsal prolateral; dps, dorsal prosubmedian; pld, prolateral dorsal; plm, prolateral median; pv, prolateral ventral; plvs, prolateral ventrosubmedian; rls, retrolateral subventral; rlsa, retrolateral subventral accessory; vm, ventral median; vpl, ventral prolateral; vps, ventral prosubmedian; vrl, ventral retrolateral; vrs, ventral retrosubmedian. ML, medial lobe; MN, medial notch; PN, proximal [5 basal] notch. Eb, external basal; Est, external subterminal; Et, external terminal; ib, internal basal; it, internal terminal; V, ventral. Scale bar 5 1 mm.

Fig. 16. Syntropinae Kraepelin, 1905, carapace, dorsal aspect, illustrating surface ornamentation. A. Balsateres cisnerosi (Ponce-Saavedra and Sissom, 2004), comb. nov., 8 (IBUNAM). B. Chihuahuanus crassimanus (Pocock, 1898), comb. nov., 8 (AMNH). C. Kochius bruneus (Williams, 1970), 8 (AMNH). D. Konetontli acapulco (Armas and Martín-Frías, 2001), comb. nov., holotype 8 (IBUNAM). Scale bars 5 1 mm (A–C), 0.5 mm (D).

Fig. 18. Syntropinae Kraepelin, 1905, dextral pedipalp chela, dorsal (A, C), retrolateral (B) and prolateral (D) aspects, illustrating carinae and distribution of trichobothria. A, B. Kochius punctipalpi (Wood, 1863), 8 (AMNH). C, D. Kuarapu purhepecha Francke and Ponce-Saavedra, 2010, paratype 8 (AMNH). Scale bars 5 1 mm (A, B), 0.5 mm (C, D).

Fig. 22. Syntropinae Kraepelin, 1905, dextral leg III, basitarsus and telotarsus, ventral aspect, illustrating macrosetae and spinules. A. Kochius punctipalpi (Wood, 1863), 8 (AMNH). B. Maaykuyak waueri (Gertsch and Soleglad, 1972), comb. nov., 8 (AMNH). C. Paravaejovis pumilis (Williams, 1980), 8 (AMNH). Scale bars 5 0.5 mm.

Fig. 23. Syntropinae Kraepelin, 1905, mesosomal sternites, ventral aspect, illustrating glandular areas on ventral surfaces. A. Kochius bruneus (Williams, 1970), 8 (AMNH), sternites V–VII. B. Syntropis macrura Kraepelin, 1900, 8 (AMNH), sternite VII. Scale bars 5 1 mm.

Fig. 24. Syntropinae Kraepelin, 1905, metasomal segments I–V, ventral aspect, illustrating carinae. A. Kochius bruneus (Williams, 1970), 8 (AMNH). B. Kuarapu purhepecha Francke and Ponce-Saavedra, 2010, paratype 8 (AMNH). C. Syntropis macrura Kraepelin, 1900, 8 (AMNH). Scale bars 5 1 mm (A, B), 2.5 mm (C).

Fig. 25. Syntropinae Kraepelin, 1905, telson, lateral aspect, illustrating macrosetae and subaculear tubercles. A. Kochius hirsuticauda (Banks, 1910), 8 (AMNH). B. Konetontli acapulco (Armas and Martín- Frías, 2004), comb. nov., holotype 8 (IBUNAM). C. Konetontli nayarit (Armas and Martín-Frías, 2004), comb. nov., holotype 8 (IBUNAM). Scale bars 5 0.5 mm.