Paravaejovis Williams 1980
Creators: name: González-Santillán, Edmundo
name: Prendini, Lorenzo
Publisher: name: Zenodo
Year: 2013
Subjects: Biodiversity
Taxonomy
Animalia
Arthropoda
Arachnida
Scorpiones
Vaejovidae
Paravaejovis

description: Paravaejovis Williams, 1980
Figures 1C, 2C, E, 3, 7, 19C, D, 22C, 27C, 28C; table 1
Vejovis pumilis Williams, 1970 [= Paravaejovis pumilis (Williams, 1980)], type species, by monotypy.
Vejovis first section (part): Hoffmann, 1931: 134, 139.
Vejovis eusthenura group (part): Williams, 1970a: 395, 396.
Vejovis pumilis group: Williams, 1970b: 297, figs. 13, 14.
Vaejovis eusthenura group (part): Williams, 1980: 55; Sissom and Francke, 1985: 1; Sissom, 1991: 26; Stockwell, 1992: 408, 409; Sissom, 1993: 68; Lourenço and Sissom, 2000: 135; Sissom, 2000: 530, 532, 551; Armas and Martín-Frías, 2001: 8; McWest, 2009: 8, 48, 52, 56, 61, 64, 98, 101–103, 108, table 1; González-Santillán, 2004: 29; PonceSaavedra and Sissom, 2004: 541; Francke and Ponce-Saavedra, 2005: 67; Sissom and Hendrixson, 2005a: 131; 2005b: 33, 34; Fet et al., 2006a: 7; 2006b: tables 1, 9; Graham and Soleglad, 2007: 9, 11, 12; Soleglad et al., 2007: 134, 135; SantibáñezLópez and Sissom, 2010: 49.
Paravaejovis Williams, 1980: 29, 30, fig. 32A–D; Haradon, 1984b: 319; Francke, 1985: 11, 21; Sissom, 1990: 107, 110, 114, 106, fig. 3.19A; Nenilin and Fet, 1992: 9; Stockwell, 1992: 409, 416, figs. 62, 63; Fet et al., 1998: 617; Kovařík, 1998: 143; Beutelspacher, 2000: 56, 63, 151; Sissom, 2000: 504, 505; Soleglad and Fet, 2003: 88; 2005: 5, 6; Prendini and Wheeler, 2005: 482, table 10; Fet et al., 2006a: 4, 7, 9, 13, fig. 6, tables 1, 2,; 2006b: 1, 2, 4, 5, 7, 8, figs. 5, 6, tables 1, 2; 2006c: 1, 2, 4, 5, 8, figs. 5, 6, table 1; Soleglad and Fet, 2006: 5, 6, 10, 12, 13, 26, tables 2, 3; 2008: 1, 3, 5, 6, 12, 13, 24, 25, 28, 30, 34, 35, 37, 38, 40, 43, 51, 54, 55, 60, 62, 63, 70, 71, 73, 76–78, 82–84, 102, figs. 1, 7, 18, 19, 57, 97–99, 126, 163, 189, 191, 196, 197, 201, tables 1, 2, 7, 8, 9; Dupre´, 2007: 9, 16, 17.
Lissovaejovis Ponce-Saavedra and Beutelspacher, 2001, syn. nov.: 88 [nomen nudum].
Hoffmannius Soleglad and Fet, 2008, syn. nov.: 1, 4, 26, 57, 60, 89, 91, 96, 102; Buthus eusthenura Wood, 1863 [= Paravaejovis eusthenura (Wood, 1863), comb. nov.], type species, by original designation; Ayrey and Soleglad, 2011: 1.
DIAGNOSIS: Species of Paravaejovis may be separated from other genera of Syntropinae by the following combination of characters. The pedipalp chela manus drl carina and the patellar rlds and rlm carinae are usually complete and smooth, costate (fig. 19C, D), becoming obsolete in some species, e.g., P. pumilis. Pedipalp chela trichobothrium esb is situated between RD5 and RD6, and/or their associated macrosetae, and trichobothrium Et 5 at the base of the fixed finger, distal to the movable finger articulation and removed from Et 4 (fig. 19D), the exception being P. pumilis, in which esb is situated at the position of RD6 (only the associated macroseta is present) and Et 5 is situated at the base of the fixed finger, proximal to the movable finger articulation but slightly distal to Et 4. The median denticle row on the pedipalp chela fixed finger comprises six primary subrows of median denticles, separated by six retrolateral denticles (fig. 19C, D) in all Paravaejovis species except P. pumilis, P. puritanus, comb. nov., and P. schwenkmeyeri, comb. nov., in which it comprises five primary subrows of median denticles, separated by five retrolateral denticles.
Paravaejovis are most closely related to Chihuahuanus, gen. nov., and Mesomexovis, gen. nov. However, compared with Chihuahuanus, gen. nov., and Mesomexovis, gen. nov., which exhibit an incrassate chela manus and relatively short fingers, most species of Paravaejovis exhibit a slender chela manus and elongate fingers, the exceptions being P. gravicaudus, comb. nov., P. pumilis, and P. spinigerus, comb. nov., in which the manus is more incrassate. Most species of Paravaejovis present psammophilous adaptations that are absent in Mesomexovis, gen. nov., and most species of Chihuahuanus, gen. nov.: pale, immaculate coloration, glabrous integument, and numerous long macrosetae on the legs, with the dorsal and retrodorsal macrosetae on the basitarsi of legs I–III arranged in a single row (setal combs) (fig. 22C). Chihuahuanus globosus, comb. nov., exhibits such characters but may be separated from Paravaejovis by the presence of a small fusiform, whitish glandular area anterior to the base of the aculeus on the dorsal surface of the telson vesicle (as illustrated for C. coahuilae, comb. nov., in fig. 26A). In addition, the vl carinae of metasomal segments I–IV are usually granular in Chihuahuanus, gen. nov. (fig. 27A), but smooth in many species of Paravaejovis (fig. 27C), exceptions being P. confusus, comb. nov., P. waeringi, comb. nov., and P. pumilis (fig. 28C). Although there are notable exceptions (e.g., P. spinigerus, comb. nov.), most species of Paravaejovis differ from Mesomexovis, gen. nov., in the pale, immaculate integument of the carapace and mesosomal tergites, which are markedly infuscated in the latter.
INCLUDED SPECIES: Paravaejovis confusus (Stahnke, 1940), comb. nov.; Paravaejovis diazi (Williams, 1970), comb. nov.; Paravaejovis eusthenura (Wood, 1863), comb. nov.; Paravaejovis flavus Banks, 1900, comb. nov.; Paravaejovis galbus (Williams, 1970), comb. nov.; Paravaejovis gravicaudus (Williams, 1970), comb. nov.; Paravaejovis hoffmanni (Williams, 1970), comb. nov.; Paravaejovis pumilis (Williams, 1970); Paravaejovis puritanus (Gertsch, 1958), comb. nov.; Paravaejovis schwenkmeyeri (Williams, 1970), comb. nov.; Paravaejovis spinigerus (Wood, 1863), comb. nov.; Paravaejovis waeringi (Williams, 1970), comb. nov.
DISTRIBUTION: Paravaejovis is endemic to Mexico (recorded from Baja California, Baja California Sur, Chihuahua, Sinaloa, and Sonora) and the United States (recorded from Arizona, California, Nevada, New Mexico, Oregon, and Utah; fig. 3). All except three of the 12 currently recognized species (P. confusus, comb. nov.; P. spinigerus, comb. nov.; P. waeringi, comb. nov.), occurring in the Great Basin, Mojave, and Sonoran (including Colorado) deserts on the North American mainland, are endemic to the Baja California Peninsula, having diversified in isolation from their mainland relatives after the opening of the Gulf of California. This genus is absent from the Chihuahuan Desert.
NATURAL HISTORY: Most species of this genus are fossorial inhabitants of desert and semidesert regions, more commonly collected by UV light detection at night than under stones or other debris during the day. Species inhabiting the Baja California Peninsula have been collected from sea level to 1180 m altitude, those on the North American mainland mostly above 2000 m. Sandy substrata evidently played an important role in the diversification of Paravaejovis in Baja California. Most species on the peninsula are psammophilous or ultrapsammophilous, characterized by pale coloration, the absence of infuscation, a smooth, glabrous integument, and setal combs on the leg tarsi, for burrowing in sand (Fet et al., 1998; Prendini, 2001a). The mainland species, by contrast, appear to be less specialized and inhabit a greater range of substrata, from sand dunes to rock walls, and vegetation types, from desert to pine-oak forest which, although present on the peninsula, is uninhabited by Paravaejovis.
Paravaejovis pumilis from Baja California Sur evolved such specialized ultrapsammophilous adaptations that it was long thought related to the largely psammophilous subfamily Smeringurinae Soleglad and Fet, 2008, and to the borregoensis microgroup of Paruroctonus in particular, with which it was suggested to be subordinate (Haradon, 1984b; Sissom, 1990, 2000; Stockwell, 1989, 1992). The borregoensis microgroup comprises small psammophilous vaejovids characterized by the absence of a mid-retrosuperior (mrs) macroseta (part of the retroventral series in the terminology used in the present contribution) on the basitarsus of leg II, also observed in P. pumilis, which shares with the borregoensis microgroup several other characters, some sexually dimorphic, concerning the carapace, pectines, legs, and metasoma (Haradon, 1984b). The putative relationship of P. pumilis to Paruroctonus and other Smeringurinae, further elaborated by Soleglad and Fet (2003, 2008), was falsified by González-Santillán and Prendini (in press), who demonstrated that it forms a divergent sister species of P. eusthenura, comb. nov., P. diazi, comb. nov., and P. hoffmanni, comb. nov. (fig. 7), separated from the latter and other members of the genus by a series of apomorphic character states, mostly associated with psammophily. Psammophily in P. pumilis and other Paravaejovis species evolved independently from that in Smeringurinae.
REMARKS: As redefined here, Paravaejovis accommodates species mostly assigned to Hoffmann’s (1931) ‘‘first section’’ of Vaejovis, and then Williams’ (1970a) eusthenura group, for which Soleglad and Fet (2008) devised the name Hoffmannius, without quantitatively testing its monophyly or composition. Hoffmannius, as defined by Soleglad and Fet (2008), was consistently polyphyletic, and the group of species hereby assigned to Paravaejovis consistently monophyletic, in the phylogenetic analyses of González-Santillán and Prendini (in press) based on DNA and those based on morphology and DNA. The following species are therefore transferred to other genera in the present contribution: Chihuahuanus bilineatus, comb. nov., Chihuahuanus coahuilae, comb. nov., Chihuahuanus glabrimanus, comb. nov., Chihuahuanus globosus, comb. nov., Mesomexovis punctatus, comb. nov., Mesomexovis spadix, comb. nov. et stat., Mesomexovis variegatus, comb. nov., Maaykuyak vittatus, comb. nov., Maaykuyak waueri, comb. nov., and Vizcaino viscainensis, comb. nov., previously assigned to the eusthenura group (Williams, 1980; Sissom, 2000) and then to Hoffmannius (Soleglad and Fet, 2008); and Mesomexovis oaxaca, comb. nov., previously assigned to the eusthenura group (Santibáñez-López and Sissom, 2010).
In addition, P. pumilis was consistently retrieved, in the phylogenetic analyses of González-Santillán and Prendini (in press) based on DNA and those based on morphology and DNA, as the sister species of P. eusthenura, comb. nov., the type species of Hoffmannius, nested within a larger monophyletic group comprising the remaining species of Hoffmannius. Paravaejovis pumilis was never recovered close to Smeringurus grandis (Williams, 1970), the exemplar species for Smeringurinae, to which it was alleged to belong (Soleglad and Fet, 2008). Among several psammophilous character states, shared with other species of Paravaejovis including P. eusthenura, comb. nov., P. pumilis shares the diagnostic synapomorphy of Syntropinae, i.e., the spinose distal barb margin of the sclerotized hemimating plug on the hemispermatophore, noted but disregarded by Stockwell (1989, 1992) and Soleglad and Fet (2008). The other putatively diagnostic difference, on which basis P. pumilis was differentiated from other vaejovids, is the neotrichobothriotaxic pedipalp chela (Williams, 1980). Aside from the fact that neobothriotaxy, as expressed in P. pumilis, is autapomorphic and thus uninformative about its phylogenetic placement, ‘‘deviations’’ from the typical ‘‘ Type C’’ trichobothrial pattern are not uncommon among Vaejovidae (Haradon, 1984b), occurring, e.g., in three other, unrelated vaejovid taxa (Gertsch and Soleglad, 1972; Sissom and Francke, 1985; Sissom, 1991; Haradon, 1984b; Soleglad and Fet, 2003, 2008): Franckeus, Paruroctonus ammonastes Haradon, 1984, and Pseudouroctonus bogerti (Gertsch and Soleglad, 1972). Such changes may result from simple genetic processes like differential expression of chromosomes and homeotic genes (Scott and Weiner, 1984). Contrary to the prevailing dogma in scorpion systematics, trichobothria should not be given ‘‘more evolutionary or taxonomic significance’’ (Soleglad and Fet, 2008: 84) than other characters. Trichobothria are characters like any other.
In view of the unambiguous phylogenetic position of the type species of Paravaejovis, Hoffmannius Soleglad and Fet, 2008, is hereby abolished: Hoffmannius Soleglad and Fet, 2008 = Paravaejovis Williams, 1980, syn. nov. Ten species listed above, formerly placed in Hoffmannius, are transferred to Paravaejovis.
González-Santillán and Prendini (in press) also identified sufficient, consistent diagnostic character differences to remove P. schwenkmeyeri, comb. nov., from synonymy with P. puritanus, comb. nov. However, Vaejovis coloradensis Williams, 1970, was not found to differ consistently from its senior synonym, P. waeringi, comb. nov., with which it is retained in synonymy, following Williams (1980). Two subspecies were not found to differ consistently from the nominotypical forms, with which they are hereby synonymized: Vaejovis diazi transmontanus Williams, 1970 = Paravaejovis diazi (Williams, 1970), syn. nov.; Vaejovis hoffmanni fuscus Williams, 1970 = Paravaejovis hoffmanni (Williams, 1970), syn. nov.
The identity of one species assigned to this genus, P. flavus, comb. nov., has been problematic since its description. Vaejovis flavus Banks, 1900, was briefly described in a key couplet. Soleglad (1973) redescribed the species based on a specimen from the Museum of Comparative Zoology (MCZ), Harvard University, presumed to be the holotype, but which turned out not to be (Sissom, 2000). The holotype was apparently deposited in the U.S. National Museum of Natural History, Smithsonian Institution, Washington, DC (USNM), the MCZ specimens belong to an undescribed species in the former punctipalpi group, and the USNM specimen to the former eusthenura group (Sissom, 2000). The type locality of this species is Albuquerque, New Mexico. Several attempts to collect new material matching the description, from the vicinity of the type locality and elsewhere in New Mexico have been unsuccessful (Sissom, 2000), as have attempts to locate the holotype in the USNM. Due to the uncertainty regarding the identity of P. flavus, comb. nov., it is considered a nomen dubium pending further investigation.
MATERIAL EXAMINED: Paravaejovis confusus (Stahnke, 1940), comb. nov.: U.S.A.: Arizona: Maricopa Co.: Wickenburg, 20.iv. 1938, A.L. Corbin, 2♀ syntypes (CAS Type No. 105170). Mohave Co.: Warm Springs Wash, 4.7 km N Topock, 34 ° 44.8662 ′ N 114 ° 28.8996 ′ W, 147 m, 9.ix.2007, L. Prendini and J. Huff, 58, 2♀ (AMNH [ARA 3117]). Paravaejovis diazi (Williams, 1970), comb. nov.: MEXICO: Baja California Sur: Municipio de Comondú: Ciudad Constitución, 34.4 km W (= El Crucero, 21.4 mi. W), Magdalena Plain, 75 ft, 26.vii.1968, S.C. Williams and M.A. Cazier, holotype ♂ (CAS Type No. 10413); Punta San Telmo on Gulf coast, 25 ft, 26.v.1969, S.C. Williams, holotype ♂ of V. diazi transmontanus (CAS Type No. 10414). Municipio de La Paz: Las Cruces, 8 km SW, 30.vii.1968, S.C. Williams and M.A. Cazier, 18, 1♀ paratypes (AMNH). Paravaejovis eusthenura (Wood, 1863), comb. nov.: MEXICO: Baja California Sur: Municipio de Los Cabos: 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 3177]). Municipio de La Paz: La Paz, 18 km SE, 24 ° 2.7498 ′ N 110 ° 8.85 ′ W, 626 m, 8.vii.2004, O.F. Francke, E. González, and A. Valdez, 18, 1♀ (AMNH [ARA 1192]). Paravaejovis galbus (Williams, 1970), comb. nov.: MEXICO: Baja California Sur: Municipio de Loreto: Loreto, 8 km S, 16.v.1969, S.C. Williams, holotype ♂ (CAS Type No. 10415), 1♀ allotype, 18, 1♀ paratypes (CAS). Paravaejovis gravicaudus (Williams, 1970), comb. nov.: MEXICO: Baja California Sur: Municipio de La Paz: Los Aripes, 21.4 mi. W, 900 ft, 25.vii.1968, S.C. Williams, J. Bigelow, and M. Bentzien, holotype ♂ (CAS Type No. 10418). Municipio de Comondú: San Miguel de Comondú, 2 km SW, 3.vii.1968, M.A. Cazier et al., 18, 1♀ paratypes (AMNH). Paravaejovis hoffmanni (Williams, 1970), comb. nov.: MEXICO: Baja California Sur: Municipio de Comondú: San Jose de Comondú, 24 mi. NE, 900 ft, 15.v.1969, S.C. Williams, holotype ♂ of V. hoffmanni fuscus (CAS Type No. 10421). Baja California: Municipio de Ensenada: Laguna Manuela, 4.8 km N, 152 m, 22.vi.1968, S.C. Williams and M.A. Cazier, 18, 1♀ paratypes (AMNH); Manuela, 3 mi. N, 500 ft, 22.vi.1968, S.C. Williams and M.A. Cazier, holotype ♂ (CAS Type No. 10420). Paravaejovis pumilis (Williams, 1970): MEXICO: Baja California Sur: Ciudad Constitución (= El Crucero), 26.8 mi. W, Magdalena Plain, 50 m, 26.vii.1968, S.C. Williams and M.A. Cazier, holotype ♂ (CAS Type No. 10425). Municipio de Comondú: El Cayuco Fish camp, sand dunes, 24 ° 34.6656 ′ N 111 ° 40.6326 ′ W, 6 m, 11.vii.2005, W.E. Savary, E. González, L. Prendini, and R. Mercurio, 28, 2♀ (AMNH [ARA 3173]). Paravaejovis puritanus (Gertsch, 1958), comb. nov.: MEXICO: Baja California: Municipio de Ensenada: Santo Tomas, 8.vii.1953, W.J. and J.W. Gertsch, holotype ♂ (AMNH). Baja California Sur: Rancho Canipole´, 1 mi. SW, 800 ft, 15.v.1969, S.C. Williams, holotype ♂ of Vaejovis terradomus Williams, 1970 (CAS Type No. 10428). Paravaejovis schwenkmeyeri (Williams, 1970), comb. nov.: MEXICO: Baja California: Municipio de Ensenada: Bahia de Los Angeles, 25 ft, 19.vi.1968, S.C. Williams and M.A. Cazier, holotype ♂ (CAS Type No. 10426). Baja California Sur: Municipio de Ensenada: San Raymundo, 12.9 km NW, 152 m, 30.vi.1968, M.A. Cazier et al., 18, 1♀ (AMNH). Paravaejovis spinigerus (Wood, 1863), comb. nov.: U.S.A.: Arizona: Maricopa Co.: Sunflower, turnoff to Bushnell Tanks, 33 ° 52 ′ 08.8 " N 111 ° 27 ′ 54.4 " W, 1057 m, 6.iv. 2007, J. Huff and L. Prendini, 18, 1♀ (AMNH). Paravaejovis waeringi (Williams, 1970), comb. nov.: MEXICO: Baja California: Municipio de Ensenada: Oakies Landing, 27 mi. S of Puertocitos, 12.vi.1968, S.C. Williams and M.A. Cazier, holotype ♂ (CAS Type No. 10431), allotype ♀ (CAS); Puertocitos, 11.vi.1968, S.C. Williams and M.A. Cazier, 28, 1♀ (CAS); Puertocitos, 44 km S, Oakies Landing, 12.vi.1968, S.C. Williams and M.A. Cazier, paratype ♂ (AMNH). U.S.A.: California: Imperial Co.: Algodones Dunes, along Grays Wells Road, S of Interstate Hwy 8, W of Imperial Sand Dunes Recreation Area, 32 ° 44.1756 ′ N 114 ° 53.43 ′ W, 50 m, 31.viii.2005, R. Mercurio and L. Prendini, 1♀ (AMNH [ARA 1499]); Andrade, 2 mi. W, 6.vii.1969, S.C. Williams and V.F. Lee, holotype ♂ (CAS Type No. 10411); San Diego Co.: Anza-Borrego Desert State Park: Culp Valley Camp, 33 ° 13.4118 ′ N 116 ° 27.26 ′ W, 1033 m, 30.viii. 2005, R. Mercurio and L. Prendini, 18, 1♀ (AMNH [ARA 1497]).
description: Published as part of 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, pp. 1-71 in Bulletin of the American Museum of Natural History 2013 (382) on pages 46-49, DOI: 10.1206/830.1, http://zenodo.org/record/4611867