Paruroctonus Werner, 1934

Soleglad, Michael E. & Fet, Victor, 2008, Contributions to scorpion systematics. III. Subfamilies Smeringurinae and Syntropinae (Scorpiones: Vaejovidae), Euscorpius 71 (71), pp. 1-115 : 75-81

Genus Paruroctonus Werner, 1934

Type Species. Uroctonoides gracilior Hoffmann ,

1931 [= Paruroctonus gracilior (Hoffmann, 1931) ].

Synonyms:

Uroctonoides Hoffmann, 1931: 405 ; a junior homonym of Uroctonoides Chamberlin, 1920 (= Teuthraustes Simon, 1878 : Scorpiones , Chactidae ).

Hoffmanniellius Mello-Leitão, 1934: 80 ; a replacement name for Uroctonoides Hoffmann, 1931 (see Sissom, 2000: 505).

References (selected):

Paruroctonus : Werner, 1934: 283, fig. 363 (a replacement name for Uroctonoides Hoffmann, 1931 ); Williams, 1972: 1–3 (in part); Stahnke, 1974: 119, 136 (in part); Williams, 1974: 15 (in part); Williams, 1980: 31–34, figs. 35–37 (in part); Sissom, 1990a: 110, 114 (in part); Stockwell, 1992: 408, 409, 416, 419, figs. 12, 37, 39, 58; Kovařík, 1998: 143; Sissom et al., 1998: 17–19; ICZN, 1999b: 209–210; Beutelspacher, 2000: 56, 65, 152 (in part); Sissom, 2000: 505– 514; Soleglad & Fet, 2003b: 88.

Vejovis (Paruroctonus) : Gertsch & Allred, 1965: 4 (in part); Williams, 1970c: 277 (in part); Gertsch & Soleglad, 1972: 4 (in part); Hjelle, 1972: 26 (in part).

Paruroctonus (Paruroctonus) : Haradon, 1983: 256; Haradon, 1984a: 205–209; Haradon, 1984 b: 317– 318; Haradon, 1985: 19–21.

Composition. This genus includes the following 33

species and subspecies:

P. ammonastes Haradon, 1984

P. arenicola arenicola Haradon, 1984 P. arenicola nudipes Haradon, 1984 P. arnaudi Williams, 1972

P. baergi (Williams et Hadley, 1967) P. bajae Williams, 1972

P. bantai bantai (Gertsch et Soleglad, 1966) P. bantai saratoga Haradon, 1985

P. becki (Gertsch et Allred, 1965) P. boquillas Sissom et Henson, 1998 P. boreus (Girard, 1854)

P. borregoensis borregoensis Williams, 1972 P. borregoensis actites Haradon, 1984

Paravaejovini

Paravaejovis

Smeringurinae

Smeringurini Paruroctonus

Smeringurus

Vejovoidus Syntropina Hoffmannius Syntropini Syntropis

Thorelliina Thorellius

Kochius Syntropinae

Stahnkeini Gertschius

Wernerius

Serradigitus

Stahnkeus

Pseudouroctonus

Uroctonites Vaejovinae *

Vaejovis

"mexicanus " group

Franckeus

Vaejovis

"nigrescens " group

P. coahuilanus Haradon, 1985 P. gracilior (Hoffmann, 1931) P. hirsutipes Haradon, 1984 P. luteolus (Gertsch et Soleglad, 1966) P. maritimus Williams, 1987 P. marksi Haradon, 1984

P. nitidus Haradon, 1984

P. pecos Sissom et Francke, 1981 P. pseudopumilis (Williams, 1970) P. shulovi shulovi (Williams, 1970) P. shulovi nevadae Haradon, 1985 P. silvestrii (Borelli, 1909) P. simulatus Haradon, 1985 P. stahnkei (Gertsch et Soleglad, 1966) P. surensis Willliams et Haradon, 1980 P. utahensis (Williams, 1968) P. variabilis Hjelle, 1982

P. ventosus Williams, 1972 P. williamsi Sissom et Francke, 1981 P. xanthus (Gertsch et Soleglad, 1966)

Soleglad & Fet: Smeringurinae and Syntropinae 77

Distribution. Canada (southern edges of Alberta and British Columbia, only P. boreus ), Mexico (Aguascalientes, Baja California, Baja California Sur, Chihuahua, Coahuila, Sonora, Zacatecas), USA (Arizona, California, Colorado, Idaho, Montana, New Mexico, Oregon, South Dakota, Texas, Utah, Washington, Wyoming).

This genus is one of the most widely spread in family Vaejovidae (see map in Fig. 197 ), but most of this is due to the very large range of species P. boreus , which extends from southern Nevada to the extreme southwestern parts of Canada. Only species P. gracilior extends any distance into mainland Mexico, its type locality being Tepezalá, Aguascalientes. Genus Paruroctonus has a disjunct range in Baja California, Mexico; only two species are found in Baja California Sur, P. pseudopumilis and P. surensis , both isolated in the Vizcaino Desert. The closest most southern species in Baja California are P. arnaudi , P. ventosus , P. silvestrii , and P. luteolus ; none are found beyond El Rosario on the east or Luis Gonzaga on the west. P. arnaudi and P. ventosus are psammophiles found in isolated inland sand dunes on the Pacific side of the peninsula.

Diagnosis. Metasomal segments I–IV with a pair of ventromedian (VM) carinae; metasomal segment I usually as wide as long in male and wider than long in female, segment III never twice as long as wide, segment IV never three times longer than wide; setal pairs found on ventromedian (VM) carinae of segments I–IV, intercarinal area of VM carinae without setation; lamina of hemispermatophore with squared or rounded distal tip, inner base of lamina without small protrusion; vesicular tabs well developed, equipped with conspicuous distal granule.

Taxonomic history. Two names, Paruroctonus Werner, 1934 and Hoffmanniellius Mello-Leitão, 1934 were proposed as replacement names for Uroctonoides Hoffmann, 1931 , a subjective junior homonym of Uroctonoides Chamberlin, 1920 (= Teuthraustes Simon, 1878 : Scorpiones , Chactidae ). The name Paruroctonus was conserved by the International Commission on Zoological Nomenclature in 1999 due to its prevalent usage (Sissom et al., 1998; ICZN, 1999b). Paruroctonus was treated for some time as subgenus of Vaejovis until reinstated as genus by Williams (1972). The genus was revised by Haradon (1983, 1984a, 1984 b, 1985) who established two subgenera ( Paruroctonus and Smeringurus ). Stockwell (1992) elevated Smeringurus Haradon, 1983 to the genus level (see below). No subgenera are currently recognized in Paruroctonus , although several informal assemblages of species (“infragroups” and “microgroups”) exist ( Haradon, 1984a, 1984 b, 1985; Sissom, 2000).

Discussion. In a series of detailed papers, Haradon (1984a, 1984 b, 1985) presented a complete revision of the genus Paruroctonus , defining, informally, no less than three “infragroups” and eight “microgroups” (also see Sissom, 2000). In these contributions ten species and subspecies were defined. Although many diagnostic characters were used in these treatments, the emphasis and most illustrations were placed on the setation of the pedipalp and the leg basitarsus and tarsus. Consequently, the question arises: do these setal patterns exhibit consistency across the various groups in Paruroctonus such that these groups can be elevated to separate genera?

After studying the three papers, we see that Haradon (1984a, 1984 b, 1985) utilized landmark seta (lms) for the pedipalp (femur and patella [addressed as tibia]), and leg III (basitarsus and tarsus). These were, in a sense, defined at a group level and therefore appear to be potential synapomorphies for these groups. However, except for the leg, the utilization of lms character set by Haradon was minimal (i.e., in the pedipalp), and did not encompass many species. In addition, lms were not always illustrated in all pedipalp femur figures, and for the patella, lms were never identified (albeit, the standard trichobothria were shown in all cases). Therefore, we limit this discussion to only leg III; in Table 10 we summarize the setation for both the basitarsus and tarsus (referred to as the telotarsus in Haradon), as were illustrated and/or discussed by Haradon (1984a, 1984 b, 1985). Utility of setation (beyond trichobothria) remains largely unexplored in scorpion systematics; see some preliminary information of pedipalp finger landmark setae in our recent work on constellation array in Smeringurinae ( Fet et al., 2006c).

For the leg III basitarsus three distally positioned lms were identified by Haradon in contrast to the superior row of large elongated setae, or the “setal combs” of Stockwell (1989); the arrangement and number of latter is considered diagnostic (Graeme Lowe informed us that this was an excellent species-level diagnostic character, pers. comm. 2007). Haradon illustrated the basitarsus III for 14 species and supplied additional information on four species in the text. We consider the number of superior setae to be a species-level character and therefore do not necessarily believe they are of importance at the group level (except, maybe, if stated as a range). It does appear that the three lms of the basitarsus are consisently placed and numbered, providing a potential diagnostic character for the genus Paruroctonus , though, of course, not germane at lower group levels.

For the leg III tarsus, Haradon identified five categories of setae: superinterminal (st), a large seta, which he always designated as lms; retrosuperior (rs) (i.e., the “setal combs” of Stockwell, 1989); retromedial (rm); retroinferior (ri); and retroinferior terminal (rit). As seen in Table 10, all these setal groups are designated as landmark groups in one context or another, depending on the microgroup. I.e. the “baergi ” and “borregoensis ” microgroups both declare ri and rit setal groups as lms, and the “shulovi ” and “borregoensis ” microgroups stipulate the rm setal group as lms. Therefore, it is clear that the lms designation was independently based on species subsets, and therefore, providing no consistency across the genus. For example, in the “boreus ” microgroup, lms are not utilized at all in the diagnosis of its species set. Similarly, landmark seta were not declared for the “williamsi ” microgroup. We also see the same number of setae in setal groups across different microgroups where the setal group is or is not declared lms. For example, P. marksi , of the “baergi ” microgroup, has two rs and rm setae, the same number of setae as in the “shulovi ” microgroup where these setal groups are designated as lms; P. shulovi , of the “shulovi ” microgroup, has two ri and rit setae, the same number as in the “borregoensis ” and “baergi ” microgroups where these same setal groups are declared as lms. Finally, species P. luteolus , of the “borregoensis ” microgroup, P. marksi , of the “baergi ” microgroup, and P. shulovi , of the “shulovi ” microgroup, all exhibit two setae each in the rs, rm, ri, and rit setal groups. In fact, based solely on these setal groups, the only difference between these three species, assigned to three separate microgroups each, is one less superior seta on the basitarsus in P. luteolus . Clearly, other diagnostic characters must be used to differentiate these three species and the groups they occupy. This is reflected in the keys provided by Haradon (1984a, 1984 b, 1985) where all lms characters are combined with other diagnostic characters such as pectinal tooth counts, coloration, metasomal setation, morphometric ratios, etc.

Based on these data, where we only consider the setation of the leg III basitarsus and tarsus, it appears that the lms as used in Haradon’s infra- and microgroup definitions do not provide, by themselves, useful cladistic characters for the further breakdown of the genus Paruroctonus . Possibly, using a combination of many characters may provide a basis for these group definitions.

Figures

Figure 197: General distribution of scorpion subfamily Smeringurinae in North America with the ranges of genera Paravaejovis (tribe Paravaejovini) and Smeringurus, Paruroctonus, and Vejovoidus (tribe Smeringurini) delineated.