Systematic Observations on North American Scorpionida with a Key and Checklist of the Families and Genera

SCOTT
Department of Entomology,

A. STOCKWELL
of California, Berkeley, California 94720

University

J. Med. Entomol. 29(3): 407-422 (1992) The phylogenetic associations of North American scorpion genera are examined. Several taxonomic changes are supported by new hypotheses of relationship. The subfamily Syntropinae is synonymized under Vaejovinae. Chactidae and Vaejovidae are restricted to their nominate subfamilies. Superstitioninae, Euscorpiinae, and Scorpiopsinae are raised to family rank. The subfamily Megacorminae is synonymized under Euscorpiinae. Anuroctonus Thorell and Ndibrotheus Williams are removed from Vaejovidae and placed in Iuridae and Chactidae, respectively. TrogEotuyosicus Lourenco and Belisarius Simon are transferred from Chactidae to Superstitionidae. Twelve species are transferred from Vaejovis Koch to Pseudouroctonus Stahnke. The diplocentrid genus Bioculus Stahnke is resurrected to contain four species from Baja California Sur, Mexico, that were traditionally placed in Didymocentrus Kraepelin. Paruroctonus (Smeringurus) Haradon is raised to the rank of genus. An illustrated key and annotated checklist are provided for the identification of the seven families and 25 genera of North American scorpions.

ABSTRACT

KEY WORDS

Arachnida,

North American scorpions, systematics, classification

CLASSIFICATION of North American scorpions (in this work, North America is taken to mean all of Mexico, the 48 contiguous United States, Alaska, and Canada) has remained nearly unchanged since its inception. As a result, our understanding of the relationships between genera has degenerated to the point that the familial assignment of a given specimen cannot be clearly ascertained without first identifying it to genus, then matching it against its traditional placement in the classification. This is because the North American scorpiofauna is composed primarily of the so-called chactoid genera, the relationships of which are extremely poorly understood (Francke 1976, Soleglad 1976, Francke & Soleglad 1981, Sissom 1990a). It is the purpose of this contribution to offer alternative hypotheses of phylogeny that in turn allow for an intuitive reconstruction of the classification of the scorpions of North America. The changes to higher classification proposed in the next section follow a cladistic analysis of all nonbuthid genera (Stockwell 1989) and are built on similar rearrangements at lower levels. These changes involve raising subfamilies that have recognized phylogenetic meaning (Sissom 1990a) to family rank, and synonymizing those that lack meaning. Although the traditional familial compositions (such as in the Chactoidea) must be cast aside, the new families retain their traditional (subfamilial) diagnoses and do not THE HIGHER 1 Current address:U.S. Army Natick RD&E Center, Biotechnology Division (STRNC-YM), Natick, Mass. 01760.

need to be redefined. The overall result is a more stable and logical family-level classification that better lends itself to phylogenetic interpretation. A comprehensive key to the scorpion genera of North America is also long overdue. Early keys to the scorpions of the United States (Banks 1900; Comstock 1912, 1940; Ewing 1928) and Mexico (Hoffmann 1931, 1932) are outdated and contain many errors. More recently, the scorpiofauna of North America has been treated by smaller regions; e.g., Baja California, Mexico (Williams 1980), Arizona (Stahnke 1940), Nevada (Gertsch & Allred 1965), Florida (Muma 1967), Utah (Johnson & Allred 1972), Idaho (Anderson 1975), and California (Hjelle 1972, Williams 1976). Many authors have worked vigorously describing members of particular North American families and genera (see the following checklist for specific citations). The description of many new genera and new understandings of the familial relationships as described in the following section has necessitated and facilitated the production of a new key to the North American scorpion genera.

Systematic Observations The terminology used here generally follows that reviewed by Hjelle (1990) and Sissom (1990a). Persons new to scorpion taxonomy should also consult Stahnke (1970b) for carinal terminologies of the carapace and opisthosomal segments, Vachon (1973) for trichobothriotaxy,

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and Francke (1978a) for pedipalp carinal terminology. As currently composed, the families Chactidae Pocock, 1893 and Vaejovidae Thorell, 1876 cannot be characterized and are taxonomically and phylogenetically meaningless (Stockwell 1989, Sissom 1990a). To clear up some of the confusion in these families, we must examine the accepted phylogenetic affinities of the genera contained therein. The Vaejovidae has traditionally comprised three subfamilies, Vaejovinae Thorell, 1876; Syntropinae Kraepelin, 1905; and Scorpiopsinae Kraepelin, 1905. The subfamily Syntropinae was originally monotypic, but now contains Syntropis Kraepelin, 1901 and Vejovoidus Stahnke, 1974. This grouping is based on the presence of a single, ventromedian keel on the metasomal segments in both genera. Despite the fact that several authors have expressed doubts about both the strength of this particular character and the validity of the generic association (Soleglad 1976, Francke 1982a, Sissom 1990a), the subfamily has never been dissolved. Vejovoidus shares several potential synapomorphies with Paruroctonus (Paruroctonus) Werner, 1934, Paruroctonus (Smeringurus) Haradon, 1983, and Paravaejovis Williams, 1980 (all traditionally placed in Vaejovinae). These include the presence of a convex anterior margin on the carapace, the presence of distinct setal combs on the tarsi, and the absence of a ventral serrula on the movable finger of the chelicera. Vejovoidus also exhibits the absence of a sclerotized mating plug in the spermatophore of males, a feature shared with Paruroctonus (Paruroctonus) and Paruroctonus (Smeringurus). This is adequate justification for transferring Vejovoidus from the Syntropinae to the Vaejovinae. Among the Vaejovinae, Syntropis appears to share at least one potential synapomorphy with the punctipalpi, eusthenura, and intrepidus groups of Vaejovis Koch, 1836 (i.e., the presence of numerous spines on the margin of the distal barb of the mating plug, a feature not present in other vaejovids). Syntropis thus appears to be a slender, single-keeled Vaejovis. For these reasons, Syntropinae is synonymized under the subfamily Vaejovinae Thorell, 1876; both Syntropis and Vejovoidus are thus placed in the Vaejovinae. The subfamily Scorpiopsinae, traditionally placed in the Vaejovidae because of the presence of three pairs of lateral eyes, is a compact group of genera and subgenera that, based on potential synapomorphies in trichobothrial patterns and paraxial organ-hemispermatophore morphology, are more closely related to Euscorpius Thorell, 1876, the Megacorminae Kraepelin, 1899, and the Chactinae Pocock, 1893, as was once recognized by Francke (1976). Scorpiopsines definitely do not represent the sister group of the

Vaejovinae and should be transferred from the Vaejovidae to a position closer to the Chactidae. The family Vaejovidae is therefore restricted to its nominate subfamily. Considerable controversy surrounds the placement of Anuroctonus Pocock, 1893 in the Vaejovidae. Originally placed in the Vaejovinae (Kraepelin 1905), the genus was placed by Stahnke (1974) in the subfamily Hadrurinae Stahnke, 1974 based on similarities in trichobothrial numbers and patterns. In their resurrection of the family Iuridae, Francke & Soleglad (1981) dismissed the characters cited by Stahnke (1974) as not indicative of phylogenetic relationship and declined to assign Anuroctonus to any group. Sissom (199Oa) recognizes the genus as incertae cedis. Anuroctonus clearly belongs to either the Vaejovidae or the Iuridae because it shares at least one potential synapomorphy with both groups (i.e., the displacement of one or more distal v trichobothria to the external side of the patella). With one exception (Uroctonus Thorell, 1876), all Vaejovidae and Iuridae share this feature. There are several elaborations on this basic vaejovoid plan. One variation is exhibited by Uroctonus (Vaejovidae), in which the distal trichobothrium has moved back to the ventral side. In the iurid genera Iurus Thorell, 1876 and Calchas Birula, 1917, both the median and distal v trichobothria are displaced to the external side of the patella. The iurid genus Hadrurus Thorell, 1876 possesses numerous (>20) additional v trichobothria, several of which are displaced to the external side of the patella. This particular arrangement is also exhibited by Anuroctonus, although there are fewer (13-15) extra trichobothria involved. Among recent scorpions, this arrangement of ventral trichobothria is seen only in Hadrurus + Anuroctonus and is considered synapomorphic. Further support for this association is seen in the external (e) series of trichobothria on the patella. The majority of vaejovids and iurids have an orthobothriotaxic number of e trichobothria on the patella. A few (e.g., some species of Vaejovis, and the iurid genera Hadruroides Pocock, 1893 and Caraboctonus Pocock, 1893) possess one additional e trichobothrium. Hadrurus and Anuroctonus, however, possess at least 15 additional e trichobothria. This trichobothrial arrangement is unique among Vaejovidae and Iuridae and is also considered synapomorphic. Anuroctonus also shares several similarities in its reproductive system with Hadrurus, Hadruroides, and Caraboctonus that it does not share with the vaejovids. These synapomorphies include the presence of fully separated genital opercula in the female and a very large, greatly coiled testis and an enlarged seminal vesicle in the male. Based on this evidence, Anuroctonus is clearly an iurid and the sister genus of Hadrurus. The

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genus is therefore transferred to the subfamily Hadrurinae in the family Iuridae. Nullibrotheus Williams, 1974 was transferred from the subfamily Chactinae Pocock, 1893 to Vaejovidae by Williams (1974). This monotypic genus does not share any synapomorphies with the Vaejovidae (i.e., the displacement of the distal ZJtrichobothrium to the external face of the patella and a hemispermatophore with the floor of the sperm duct developed into a narrow, invaginated pouch). It does have clear affinities with the Chactinae, Scorpiopsinae, Megacorminae, and Euscorpius, with which it shares several potential synapomorphies: five or more ti and 17 or more e trichobothria on the patella (13 v and 114 e are found in all vaejovids), and the seminal vesicle of the paraxial organ enlarged and adnate to the distal lamella sheath (seminal vesicle is of normal proportions and free in vaejovids). Of these groups, Nullibrotheas appears more closely related to the Chactinae by virtue of its cheliceral serrula (not present in Scorpiopsinae, Megacorminae, or Euscorpius) and very similar hemispermatophore morphology. For these reasons, Nullibrotheas is transferred back to the subfamily Chactinae. We are thus left with a very compact group of genera forming the Vaejovidae: Vuejovis, Syntropis, Serradigitus Stahnke, 1974, Pseudouroctonus Stahnke, 1974, Uroctonites Williams & Savary, 1991, Uroctonus, Paravaejovis, Vejovoidus, Paruroctonus (Paruroctonus), and Paruroctonus (Smeringurus). Distinct groupings within this family are clear enough to recognize subfamilies, but I will defer on this subject until a later time. The differences between P. (Paruroctonus) and P. (Smeringurus) are as significant as differences between any of the other genera; therefore, I give Smeringurus full generic status. The transfer of three species formerly maintained in Uroctonus to a new genus, Uroctonites, by Williams & Savary (1991) renders the former genus, as interpreted by Gertsch & Soleglad (1972), Soleglad (1973), and Sissom (1990a), polyphyletic and difficult to characterize. Williams & Savary (1991) attempt to remedy this situation by restricting Uroctonus to three species (U. mordax Thorell, 1876, U. frunckei Williams, 1986, and U. gruhami Gertsch & Soleglad, 1972), and transferring the remaining species to the minimus group of Vaejovis. I agree that Uroctonus must be restricted to the three species named above and that the remaining species should be transferred to the minimus group. However, retaining the minimus group in the genus Vaejovis is undesirable because it confuses the taxonomic definitions of both the uroctonoids and Vaejovis and does not indicate any phylogenetic position for the minimus group species. By extending the arguments of Williams & Savary (1991), it is possible to assign the minimus group on a phylogenetic basis.

Williams & Savary (1991) cite the presence of an elevated lamellar hook on the spermatophore as a synapomorphy uniting the minimus group with the eusthenuru, punctipalpi, and intrepidus groups, Serrudigitus, and Syntropis. Pseudouroctonus (not considered a valid genus by Sissom [ 19911) and Parclvclejovis also exhibit this apomorphy. As stated earlier, the presence of a spiny distal barb margin on the spermatophore mating plug is a potential synapomorphy uniting the eusthenuru, punctipalpi, and intrepidus groups, Syntropis, and Paravaejovis. This places the minimus group in a polytomy with the clade eusthenura group + punctipulpi group + intrepidus group + Syntropis + Puruvuejovis, as well as Pseudouroctonus and Serrudigitus, two very dissimilar genera. The minimus group species share several characters with Pseudouroctonus, as well as Uroctonus and Uroctonites, that they do not share with Serrudigitus. These characters include the reduction of the ventral internal and internal carinae of the pedipalp chela and patella, respectively; the presence of inferior accessory cheliceral teeth in many species; a densely granular and noticeably hydrophilic integument; and strong, densely granular carinae. Two species, V. iviei Gertsch & Soleglad, 1972 and V. glimmei (Hjelle 1972), d o not possess the first two characters but are undoubtedly minimus group species based on hemispermatophore morphology, which is unique in several smaller, definable groups within this complex. These two species serve to illustrate that the inflation of the chela is not necessarily correlated with the loss of chelal and patellar carinae. The minimus group species share no synapomorphies with Serrudigitus that are not also shared with other groups. The minimus group is thus more closely related to Pseudouroctonus than to any other vaejovid group. To make Vaejovis and the uroctonoid genera more easily distinguishable, I advocate the the transfer of the the so-called minimus group of Vaejovis to Pseudouroctonus. Pseudouroctonus is therefore recognized as a valid genus with a revised diagnosis using the characters listed above. The following species are transferred to Pseudouroctonus: Vaejovis rufulus (Gertsch & Soleglad, 1972) = Pseudouroctonus rufulus (Gertsch & Soleglad, 1972); Vaejovis bogerti (Gertsch & Soleglad, 1972) = Pseudouroctonus bogerti (Gertsch & Soleglad, 1972); Vaejovis williumsi (Gertsch & Soleglad, 1972) = Pseudouroctonus williumsi (Gertsch & Soleglad, 1972); Vuejovis andreas (Gertsch & Soleglad, 1972) = Pseudouroctonus andreas (Gertsch & Soleglad, 1972); Vaejovis angelenus (Gertsch & Soleglad, 1972) = Pseudouroctonus angelenus (Gertsch & Soleglad, 1972); Vuejovis Zindsayi (Gertsch & Soleglad, 1972) = Pseudouroctonus Zindsuyi (Gertsch & Soleglad, 1972); Vaejqvis

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chicane (Gertsch & Soleglad, 1972) = Pseudouroctonus Chicano (Gertsch & Soleglad, 1972); Vuejovis apucheunus (Gertsch & Soleglad, 1972) = Pseudouroctonus upucheunus (Gertsch & Soleglad, 1972); Vuejovis glimmei (Hjelle, 1972) = Pseudouroctonus glimmei (Hjelle, 1972); Vuejovis montcuzieri (Williams, montcuxieri (Wil1980) = Pseudouroctonus liams, 1980); Vuejovis iviei Gertsch & Soleglad, I972 = Pseudouroctonus iviei (Gertsch & Soleglad, 1972); Vuejovis minimus minimus Kraepelin, 1911 = Pseudouroctonus minimus minimus (Kraepelin, 1911); Vuejovis minimus custuneus Gertsch & Soleglad, 1972 = Pseudouroctonus minimus custuneus (Gertsch & Soleglad, 1972); and Vuejovis minimus thompsoni Gertsch & Soleglad, 1972 = Pseudouroctonus minimus thompsoni (Gertsch & Soleglad, 1972). The family Chactidae is in slightly better taxonomic condition than the Vaejovidae, but it still requires some revision. The subfamily Superstitioninae Stahnke, 1940 is of particular concern. It was originally monotypic (Stahnke 1940) and was placed in the Chactidae because Superstitioniu Stahnke, 1940 has only two pair of lateral eyes. As genera were added to the subfamily, it became clear that the placement of these species in the Chactidae was unsatisfactory (Mitchell 1968, Francke 1982b). The lateral eye number criterion for assignment of genera to families has been discredited elsewhere (Mitchell 1968, Gertsch & Soleglad 1972, Gonzalez-Sponga 1977, Francke & Soleglad 1981, Sissom 199Oa) and deserves no further discussion here. Considering that the placement of the superstitionines into the Chactidae was based on overall similarity of plesiomorphies, it is easy to determine the correct affinities of the group by examining shared apomorphies with other Chactoidea. The absence of apomorphies that are not troglobitic features among the genera of Superstitioninae make their evaluation somewhat difficult. Furthermore, there are very few specimens of these interesting scorpions, and particular characters are often not known for all species. In superstitionine males, the dorsal trough of the hemispermatophore (a keel on the external side of the hemispermatophore) is produced into a sclerotized hook at its anterior (sperm duct) end. This may represent a basally displaced lamellar hook or an entirely separate structure, but it nonetheless represents a unique condition among Recent scorpions and is considered a synapomorphy for the group. Francke (1982b) provides several other compelling arguments for maintaining the group. The first clue as to the phylogenetic affinities of the Superstitioninae may be found in their trichobothrial patterns. Superstitioniu, the only fully epigean member of the group, possesses a more or less orthobothriotaxic arrangement of patellar trichobothria with one exception: the distal

v trichobothrium is displaced to the external side of the segment. This is a potential synapomorphy and Vaejovidae + Iubetween Superstitioniu ridae. The remaining genera have neobothriotaxic arrangements that do not readily provide evidence of relationships to Vaejovidae + Iuridae. However, if we accept the hypothesized relationship of the group as outlined above, then we can interpret the differing trichobothrial patterns as follows. Alucrun Francke, 1982, Typhlochuctus Mitchell, 1968, and Sotunochuctus Francke, 1986 exhibit a character reversal in which the distal v trichobothrium of the patella migrates back to the ventral surface of the segment. These genera also have developed one or more additional external trichobothria on the patella. Typhlochuctus and Sotunochuctus shared one derived pattern in which either the second or third v trichobothrium is completely absent (leaving two ventrally), and one additional trichobothrium has appeared in the esb series (labeled sb by Mitchell [1968]). Alucrun, on the other hand, still has all three v trichobothria but shows 21 external trichobothria, all but obliterating the primitive pattern. An alternate hypothesis has Typhlochuctus and Sotunochuctus losing the third v trichobothrium directly from its external position, but in light of the apparently close relationship between these genera and Alucrun based on other characters (Francke 1982b), the former character state hypothesis is preferred. Superstitionines share no synapomorphies with the Chactinae, Scorpiopsinae, Megacorminae, or Euscorpius. In contrast, however, these groups all exhibit the presence of five or more ventral patellar trichobothria, none of which has been displaced to the external side, and 17 or more external patellar trichobothria. These are hypothesized synapomorphies for the group Chactinae + Scorpiopsinae + Megacorminae + Euscorpius. The subfamily Superstitioninae is thus more closely related to the Iuridae and Vaejovidae than it is to the Chactidae and is raised to family rank to reflect this hypothesized phylogenetic position. These three families form a monophyletic clade and are herein referred to as vaejovoids. Although not part of the North American fauna, it is appropriate at this point to treat the genera Troglo tu yosicus Lourenco, 198 1 (currently Chactinae) and Belisurius Simon, 1879 (currently Euscorpiinae). Both exhibit troglobitic characters, primarily the loss of the median, and in Belisurius, the lateral eyes as well. In most other characters, these genera are plesiomorphic. TrogZotuyosicus has a trichobothrial pattern identical to that of the Vaejovidae, with the distal v trichobothrium of the patella displaced to the external side. This is a potential synapomorphy shared

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with the vaejovoids. Belisarius is completely orthobothriotaxic and has all three v trichobothria located ventrally on the patella in an arrangement similar to that of AZacran. Hemispermatophores are unknown for these two genera, so a more definitive association with the superstitionines cannot be demonstrated at this time; however, the genera Troglotayosicus and Belisarius are here transferred to the Superstitionidae. This relationship, suggested by Vives (1981) and Dumont (1986), is an improvement over the current classification scheme because neither species is closely related to the Chactidae. The Chactidae can thus be narrowed to the subfamilies Chactinae, Scorpiopsinae, Megacorminae, and Euscorpiinae, which, as mentioned above, all share two hypothesized synapomorphies. These chactoid groups, however, are widely separated geographically, and, with the exception of Megacorminae and Euscorpiinae, are fairly distinct morphologically. In the interest of long-term stability and taxonomic continuity, I advocate the recognition of the Chactinae and Scorpiopsinae as families (i.e., Chactidae [restricted] and Scorpiopsidae). A similarity between Euscorpius and the megacormines was first recognized by Birula (1917) and later by Soleglad (1976). Euscorpius and Megacormus Karsch, 1881 do share a unique hemispermatophore morphology (i.e., the outer [anteriormost] lobe bears a row of strong spines along the sperm duct opening). The hemispermatophore of Plesiochactas Pocock, 1900 is unknown but is probably like that of Megacormus. Megacormus, Plesiochactas, and Euscorpius also have a single ventromedian keel on the metasomal segments. However, Troglocormus Francke, 1982, has paired ventromedian keels on the metasoma and a completely different hemispermatophore, which resembles that of scorpiopsines and chactines and is presumably plesiomorphic. Troglocormus does share a unique trichobothrial arrangement with Euscorpius, Megacormus, and Plesiochactas; the displacement of chelal trichobothrium V, from the ventral to the external side of the segment. Euscorpius thus appears to be more closely related to Megacormus and Plesiochactas than does Troglocormus. Therefore, the subfamily Megacorminae Kraepelin, 1899 must be considered a junior synonym of Euscorpiinae Laurie, 1896, which is raised to family rank to correspond with Chactidae and Scorpiopsidae. Together, these three families form a monophyletic group recognized here as the chactoids. I have chosen to resurrect the genus Bioculus Stahnke, 1968, for the four diplocentrid species known from Baja California Sur, Mexico (Stahnke 1968, Williams & Lee 1975, Williams 1980). This genus was synonymized under Didymocentrus Kraepelin, 1905 by Williams & Lee (1975), but the Baja species continue to be prob-

lematic in their morphological characters and geographic distribution (Stockwell 1988). These four species share one synapomorphy with other Didymocentrus in which the external or dorsal secondary carina (or both) is more prominent than the digital carina, but lack other synapomorphies found in Central American and Antillean species. Unfortunately, I am reluctant to assign a synapomorphy to these four species. Bioculus appears to represent an early branch of the Didymocentrus line that became isolated on the Baja California peninsula and subsequently underwent speciation. Bioculus, therefore, appears to be the sister genus of Didymocentrus. Based on the original description and figure (Penther 1913), Pentheria Francke, 1985, a replacement name for Parubroteas Penther, 1913, is considered a junior subjective synonym of Vuejovis. Sissom (1990a) apparently also considered this genus to be of doubtful taxonomic status because he left it out of his review of scorpion systematics. The buthid genus Darchenia Vachon, 1977, described from Merida, Yucatan, Mexico, most closely resembles Old World genera such as Compsobuthus Vachon, 1949 and Hottentotta Birula, 1908, and may not be a natural part of the Mexican fauna. I have included it in the key on the chance that it may be collected again. The genus Tityus Koch, 1836 (Buthidae) is not found in North America but is included in the key because it is occasionally introduced on produce and other goods imported from Central and South America. Of the vaejovoid families, only the Vaejovidae is entirely North American in distribution. Three of the 10 genera in this family are monotypic and known only from Baja California. The family Iuridae has two genera that are endemic to North America, but others are known from South America (two genera) and the northeastern Mediterranean area (two genera). The family Superstitionidae is known mostly from North America, but the monotypic genera Troglotayosicus from Colombia and BeZisarius from Spain and France also belong to this group. Curiously, all the genera of Superstitionidae except Superstitionia are troglobitic. The family Chactidae is represented in North America by one monotypic genus endemic to Baja California Sur. The remaining genera of this family are distributed in South America. The family Euscorpiidae contains four genera, three of which occur in North America. The nominate genus is widespread in the Mediterranean region. A third chactoid family, Scorpiopsidae, is distributed from the Himalayas to Indonesia in southeastern Asia. The family Diplocentridae contains eight genera and has a disjunct distribution in the New World and southwestern Asia (two genera). The genus Diplocentrus Peters, 1862 is very diverse

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in Mexico but also has representatives in Belize, Guatemala, and Honduras. The genus Bioculus is restricted to Baja California Sur, Mexico. The genus Didymocentrus occurs in Central America and the Antilles. The remaining diplocentrids (four genera) are found in South America and the Antilles. Only one of the ~50 genera contained in the Buthidae is found naturally in North America. Centruroides Marx, 1890 is also found throughout Central America and the Antilles, and parts of northern South America. It is the only medically important genus in North America. Key to the Families and Genera of North American Scorpions The key has little utility outside the context of North America. To facilitate species level identifications, useful revisionary works have been annotated in the checklist. These citations, however, are by no means exhaustive. The numbers of species given in the checklist refer only to the North American species of each genus. The figures are not drawn to scale. 1. Pedipalp patella lacking ventral trichobothria (Fig. 1); inferior margin of movable finger of chelicera bearing two large ac. .2 cessory teeth (Fig. 8) . . BUTHIDAE Pedipalp patella with two or more ventral trichobothria (Fig. 2-7); inferior margin of movable finger of chelicera variable, but never with two large accessory teeth (Fig. 9-15) . . . . . . . . . . . . .4 2. Dentate margin of pedipalp chela fixed finger bearing seven to nine rows of primary granules flanked internally and externally by rows of supernumerary granules (Fig. 16) . . . . . . . . . . Centruroides Dentate margin of pedipalp chela fixed finger bearing lo-17 rows of primary granules, supernumerary granules lacking (Fig. 17) . . . . . . . . . . . . . . . . .3 3. Telson bearing a distinct rhomboid subaculear tubercle; tibia1 spurs absent. Tityus Telson lacking a well-developed subaculear tubercle; tibia1 spurs present on legs III and IV . . . . . . . . . Durcheniu 4. Telson always bearing a papillate subaculear tubercle covered with whitish microsetae (Fig. 18); movable finger of chelicera with one superior subdistal tooth, ventral serrula never present (Fig. . . . . .5 9) . . . . . . DIPLOCENTRIDAE. Telson usually without a subaculear tubercle (Fig. 19), or if subaculear tubercle is present, it is usually spinoid and never bears numerous whitish microsetae; movable finger of chelicera bearing two superior subdistal teeth, serrula variable (Fig. lo-15), or if only one subdistal tooth is present, then a

5.

6.

7.

8.

9.

10.

ll.

conspicuous serrula is also present (Fig. lo)..........................6 Digital keel of pedipalp chela more than prominent secondary keels (Fig. 20) . . . . . . . . . . . . . . Diplocentrus Secondary keels of pedipalp chela more prominent than digital keel (Fig. 21) . . . . . . . . . . . . . . . . . Bioculus Pedipalp patella bearing three or fewer ventral trichobothria (Fig. 2-4) . . . . . .7 Pedipalp patella bearing six or more ventral trichobothria (Fig. 5-7) . .20 Dentate margins of pedipalp chela fingers armed with five to seven oblique subrows of primary granules (Fig. 22); spiracles small, round to short-ellipsoid . SUPERSTITIONIDAE. .8 (Fig. 24-25). Dentate margins of pedipalp chela fingers armed with one to seven continuous subrows of primary granules (Fig. 23); spiracles long-ellipsoid to slitlike (Fig. 26) . . . . .VAEJOVIDAE . . . . -11 Carapace with median and lateral eyes (Fig. 27); two basalmost teeth of fixed finger of chelicera forming a distinct bicusp (Fig. 11); telotarsus with a ventromedian longitudinal row of setal tufts (Fig. 29); all legs bearing prolateral and retrolateral pedal spurs (Fig. 29) . . . . . . . . . . . . . . . . . . . Superstitionia Carapace completely lacking eyes (Fig. 28); two basalmost teeth of fixed finger of chelicera not forming a distinct bicusp (Fig. 10); telotarsus lacking a ventromedian longitudinal row of setae (Fig. 30); all legs lacking retrolateral pedal spurs (Fig. 30) . . . . . . . . . . . . . .9 Pedipalp patella with three ventral (Fig. 31) and 20-21 external trichobothria (Fig. 32) . . . . . . . . . . . . . . . . . . Alucran Pedipalp patella with only two ventral (Fig. 3) and 14-15 external trichobothria (Fig. 33) . . . . . . . . . . . . . 10 Pedipalp chela finger with internal terminal trichobothrium (it) situated near mid-finger, external trichobothrial series located in distal half of finger (Fig. 34) . . . . . . . . . . . . . . . . . Sotanochactas Pedipalp chela finger with trichobothrium it situated at base of finger, external (e) trichobothrial series located on middle area of finger (Fig. 35). . . . . . . . .*. . . . . .*. . . . . Typhlochactas Anterior margin of carapace emarginate to nearly straight (Fig. 36); tarsi of legs I-III lacking distinct setal combs (Fig. 38); inferior surface of movable finger of chelicera bearing a serrula of variable development (Fig. 11, 50-51) . . . 12 Anterior margin of carapace straight to convex (Fig. 37); tarsi of legs I-III usually with well-developed setal

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Diagnostic features of North American scorpion genera. (l-7) Ventral aspect of right pedipalp Fig. 1-17. (2) DipZocentrus, (3) Typhlochactas, patella showing positions of ventral trichobothria; (1) Centruroides, (4) Vuejouis, (5) Anuroctonus, (6) Ndibrotheus, (7) Megacormus. (8-15) Ventral aspect of right chelicera; (8) Centruroides, (9) Diplocentrus, (10) Sotanochuctas, (11) Vaejozjis, (12) Pururoctonus, (13) Hudrzsrus, (14) Nullibrotheus, (15) Megacormus. (16-17) Dentate margin of fixed finger of right pedipalp chela showing configuration of denticles; (16) Centruroides, (17) Tityus. Abbreviations: u, accessory teeth; b, basal tooth; m, median tooth; pg, primary granules; s, subdistal tooth; sg , supernumerary granules.

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Fig. 18-37. Diagnostic features of North American scorpion genera. (lS-19) Lateral aspect of telson; (18) Diplocentrus, (19) Serrudigitus. (20-21) External aspect of right pedipalp chela showing positions of external keels; (20) Diplocentrus, (21) Bioczdus. (22-23) Dentate margin of fixed finger of right pedipalp chela showing configuration of denticles; (22) Superstitioniu, (23) Vuejocis. (24-26) Shapes of spiracles; (24) Alum-an, (25) Superstitioniu, (26) Vuejoois. (27-28) Dorsal aspect of carapace showing positions of (and lack of) median and lateral eyes; (27) Superstitioniu, (28) Typhlochuctus. (2%30) Ventral aspect of right telotarsus; (29) Superstitioniu, (30) Typhlochactus. (31) Ventral aspect of right pedipalp patella of Alucrun showing three ventral trichobothria. (32-33) External aspect of right pedipalp patella showing positions of external trichobothria; (32) Alucrun, (33) Typhlochuctus. (34-35) Fixed finger of right pedipalp chela showing positions of dorso-external trichobothria; (34) Sotanochuctus, (35) Typhlochuctus. (36-37) Dorsal aspect of carapace showing contour of frontal margin; (36) Vuejovis, (37) Pururoctonus. Abbreviations: /I)IS, prolateral pedal spur; rps, retrolateral pedal spur; d, digital keel; Ze, lateral eyes; me, median eyes; se, secondary keels; st, subaculear tubercle; tt, telotarsus; eb, external basal trichobothrium; esb, external suprabasal trichobothrium; est, external subterminal trichobothrium; et, external terminal trichobothrium; it, internal terminal trichobothrium; ib, internal basal trichobothrium.

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Fig. 38-51. Diagnostic features of North American scorpion genera. (3!%39) Retrolateral aspect of right leg III; (38) Vaejovis, (39) Paruroctonus. (40-41) External (lateral) aspect of fixed finger of right pedipalp chela; (40) Serradigitus, (41) Vaejouis. (42-43) Ventral aspect of right pectine; (42) Serrudigitus, (43) Vaejovis. (44-45) Ventral aspect of metasomal segments I-IV showing ventral keels; (44) Syntropis, (45) Vuejouis. (46-47) Cross section of pedipalp chela showing positions of keels; (46) Vuejovis, (47) Uroctonus. (48-49) Internal aspect of right pedipalp patella showing development of internal median keel; (48) Vuejouis, (49) Uroctonus. (50-51) Ventral aspect of right chelicera; (50) Vuejouis, (51) Uroctonus. Abbreviations: bt, basal tubercle; d, digital keel; di, dorsointernal keel; dm, dorsal marginal keel; ds, dorsal secondary keel; es, external secondary keel; i, internal trichobothrium; im, internal median keel; pp, proximal pectinal tooth; SC,setal comb; ui, ventral internal keel; OTTX, ventromedian keel; usm, ventral submedian keel.

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JOURNAL OF MEDICAL ENTOMOLOGY combs (Fig. 39); inferior surface of movable finger of chelicera lacking a serrula (Fig. 12) . . . . . . . . . . . . . . . . . . . . .17 Metasomal segments I-IV with a single ventromedian carina (Fig. 44) . Syntropis Metasomal segments I-IV with paired ventral submedian carinae or completely lacking ventral submedian carinae (Fig. 45) . . . . . . . . . . . . . . . . 13 Pedipalp chela fixed finger with primary row of denticles serrate in side view, undivided or divided into two to five (never six) subrows by almost imperceptibly larger denticles (Fig. 40); females with one or more pairs of proximal pectinal teeth ovoid and lacking sensillae (Fig. 42) . . . . . . . . . . . . . . Serrtldigitus Pedipalp chela fixed finger with primary row of denticles not especially serrate in side view, divided into five or six subrows by larger denticles (Fig. 41); proximal pectinal teeth not ovoid or lacking sensillae (Fig. 43) . . . . . . . . .14 Pedipalp chela oval (with six or fewer complete carinae) or octagonal (with eight complete carinae) in cross section (Fig. 46); pedipalp patella with a welldeveloped internal median carina extending distally beyond the internal trichobothrium (Fig. 48); inferior surface of movable finger of chelicera with a moderate to very weak serrula and lacking accessory teeth (Fig. 50) . . . Vuejovis Pedipalp chela septagonal in cross section, with seven strong, granular carinae (Fig. 47); internal median carina of pedipalp patella obsolete or incomplete, not extending beyond the internal trichobothrium (Fig. 49); inferior surface of movable finger of chelicera with a long, well-developed serrula, with or without accessory teeth (Fig. 51) . . . . . . . . . . 15 Pedipalp patella with third ventral trichobothrium located ventrally (Fig. 52); external suprabasal trichobothrium (esb) of fixed finger proximal to external basal trichobothrium (eb) (Fig. 53) . . Uroctonus Pedipalp patella with third ventral trichobothrium located externally (Fig. 4); chelal trichobothrium esb at the level of or distal to eb (Fig. 54) . . . . . . . . . 16 Ventrolateral setae of telotarsi I-III spiniform (Fig. 55) . . . . . . . . . . . Uroctonites Ventrolateral setae of telotarsi I-III setiform (Fig. 29, 30, 38) Pseudouroctonus Mktasdmal seg*ments III and IV with a single ventromedian carina (Fig. 56); dorsolateral carinae of metasomal segments with greatly enlarged distal dentitles (Fig. 57) . . . . . . . . . . . Vejovoidus Metasomal segments III and IV

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12.

18.

13.

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20.

14.

21.

22.

23.

15.

24.

16.

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with paired ventral submedian carinae (Fig. 58 and 59); d orsolateral carinae of metasomal segments lacking enlarged distal denticles (Fig. 60) . . . . . . . . . . 18 Pedipalp chela with no more than six ventral trichobothria (Fig. 61); dentate margin of fixed finger of pedipalp chela with six subrows of primary denticles (Fig. 23) . . . . . . . . . . . . . . . . . . . . .19 Pedipalp chela with 210 ventral trichobothria (Fig. 62); dentate margin of fixed finger of pedipalp chela with four to five subrows of primary denticles (Fig. 63) . . . . . . . . . . . . . . Paravaejovis Ventral setae on metasoma generally paired and confined to the keels (Fig. 58) . . . . . . . . . . . . . . . . . Paruroctonus Ventral surface of metasoma with numerous small reddish setae in addition to the major setae along the keels (Fig. 59 and 60) . . . . . . . . . Smeringurus Pedipalp chela with >lO ventral trichobothria (Fig. 64); ventral trichobothrial series on pedipalp patella irregular and continuing on to the external side (Fig. 5). . . . . . . . IURIDAE. . . . . . . .21 Pedipalp chela with four ventral trichobothria (Fig. 65); ventral trichobothrial series on pedipalp patella in a single row and not continuing on the external side (Fig. 6) . . . . . . . . . . . . . . . .22 Pedipalp chela fingers with seven or eight continuous subrows of primary denticles (Fig. 66) . . . . . . . Anuroctonus Pedipalp chela fingers with nine or 10 oblique subrows of primary denticles (Fig. 67) . . . . . . . . . . . . . . . . Hadrurus Chelicera bearing a conspicuous serrula (Fig. 14) . CHACTIDAE . Nullibrotheas Chelicera lacking a serrula (Fig. . . . . . . .23 15) . . . . . . EUSCORPIIDAE Metasomal segments with paired ventral submedian carinae (Fig. 68); median eyes absent (Fig. 70) . . . . . Troglocormus Metasomal segments with single ventromedian carina (Fig. 69); median eyes present (Fig. 71) . . . . . . . . . . . .24 Dentate margin of fixed finger of pedipalp chela with external supernumerary granules only (Fig. 72); pedipalp patella with seven to eight et trichobothria (Fig. 74) . . . . . . . . . . . . . . . . . . Megacormus Dentate margin of fixed finger of pedipalp chela with internal and external supernumerary granules (Fig. 73); pedipalp patella with nine et trichobothria (Fig. 75) . . . . . . . . . . Plesiochactas

Checklist of the Families and Genera of North American Scorpions I. BUTHIDAE Simon, 1879.

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et

Fig. 61-75. Diagnostic features of North American scorpion genera. (61-62) Ventral aspect of right pedipalp chela showing ventral trichobothria; (61) Paruroctonus, (62) Paruoaejoois. (63) Dentate margin of fixed finger of pedipalp chela showing configuration of denticles in Puruzjuejozjis. (64-65) Ventral aspect of right pedipalp chela showing ventral trichobothria; (64) Anuroctonus, (65) Nullibrotheus. (66-67) Dentate margin of fixed finger of pedipalp chela showing configuration of denticles; (66) Anuroctonus, (67) Hudrurus. (68-69) Ventral aspect of metasomal segments I-IV showing ventral keels; (68) Troglocormus, (69) Megucormus. (70-71) Dorsal aspect of carapace; (70) Troglocormus, (71) Megucormus. (72-73) Dentate margin of fixed finger of pedipalp chela showing configuration of denticles; (72) Megucormus, (73) Plesiochuctus. (74-75) External aspect of right pedipalp patella showing positions of external trichobothria; (74) Megucormus, (75) Plesiochuctus. Abbreviations: esg, external supernumerary granules; et, external terminal trichobothria; isg, internal supernumerary granules; Ze, lateral eyes; me, median eyes; pg, primary granules; o, ventral trichobothrium; om, ventromedian keel; usm, ventral submedian keel.

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1. Centruroides Marx in Howard, 1890. California, Nevada, Utah, Arizona, Colorado, New Mexico, Nebraska, Kansas, Oklahoma, Texas, Missouri, Arkansas, Tennessee, Mississippi, Louisiana, Georgia, Florida; all of Mexico. 21 spp. Additional species in Central America and the Antilles. See Stahnke & Calos (1977) for a key to the species of this genus. 2. Tityus Koch, 1836. Does not occur naturally in the United States or Mexico but is included here because it is an occasional stowaway on produce from South America. 3. Darchenia Vachon, 1977. Yucatan, Mexico. Monotypic. Closely resembles Old World genera and may not be a natural part of the North American fauna. II. CHACTIDAE Pocock, 1893. 4. Nullibrotheas Williams, 1974. Baja California Sur. 1 sp. See Williams (1974, 1980). III. EUSCORPIIDAE Laurie, 1896. 5. Troglocormus Francke, 1982. Tamaulipas, San Luis Potosi. 2 spp. See Francke (1982a). 6. Megacormus Karsch, 1881. TamauliPotosi, Queretaro, pas, San Luis Hidalgo, Veracruz, Oaxaca. 3 spp. See Soleglad (1976) for keys to the species of Megacormus and Plesiochactas. 7. Plesiochactas Pocock, 1900. Veracruz. 1 sp. One additional species known from Guatemala. See Soleglad (1976). IV. SUPERSTITIONIDAE Stahnke, 1940. 8. Superstitionia Stahnke, 1940. California, Nevada, Arizona; Baja California, Baja California Sur, Sonora. 1 sp. See Williams (1980). 9. Alacran Francke, 1982. Known only from very deep caves in northern Oaxaca. 1 sp. This genus and its phylogenetic position with respect to the other genera in this family are discussed by Francke (1982b). 10. Sotanochactas Francke, 1986. San Luis Potosi. 1 sp. See Francke (1986). 11. Typhlochactas Mitchell, 1968. Tamaulipas, San Luis Potosi, Veracruz, Oaxaca. 5 spp. See Mitchell (1968), Mitchell & Peck (1977), Francke (1982b), and Sissom (1989a). V. IURIDAE Thorell, 1876. 12. Anuroctonus Pocock, 1893. California, Idaho, Nevada, Utah, Arizona, New Mexico; Baja California. 1 sp. See Gertsch & Allred (1965) and Williams (1980). 13. Hadrurus Thorell, 1876. California, Oregon, Idaho, Nevada, Utah, Arizona;

VI.

Baja California, Baja California Sur, Sonora, Guerrero, Puebla, Oaxaca. 8 spp. See Stahnke (1969), Williams (1970b, 1980), and Soleglad (1975). VAE JOVIDAE Thorell, 1876. 14. Vaejovis Koch, 1836. California, Nevada, Utah, Arizona, New Mexico, Texas, Kentucky, Tennessee, Mississippi, Alabama, Georgia, North Carolina, South Carolina; all of Mexico. 70 spp. There are no comprehensive works dealing with all the species of this genus, which seems to be under revision. constant See Hoffmann (1931), Williams (1968, 197Oc, 1971, 1976, 1980, 1986a), Francke (1977a), Sissom & Francke (1985), and Sissom (1986b, 1989b, 1989c, 1990b, 1991). 15. Serrudigitus Stahnke, 1974. California, Nevada, Idaho, Utah, Arizona, Texas; Baja California, Baja California Sur, Sonora, Coahuila. 21 spp. The genus formerly comprised the wupatkiensis group of Vaejovis. There is no key to all of the species of this genus, but Williams (1980) covers those of the Baja California and Sissom & Stockwell (1992) cover those of Sonora, Mexico. See also Williams (1970a), Soleglad (1972, 1974), and Williams & Berke (1986). 16. Syntropis Kraepelin, 1901. Baja California Sur. 1 sp. See Williams (1980). 17. Pseudouroctonus Stahnke, 1974. California, Arizona, New Mexico, Texas; Baja California, Baja California Sur, Sonora, Chihuahua. 17 spp. See Gertsch & Soleglad (1972) and Hjelle (1972). 18. Uroctonites Williams and Savary, 1991. California, Arizona. 4 spp. See Williams & Savary (1991). 19. Uroctonus Thorell, 1876. Washington, Oregon, California. 3 spp. See Gertsch & Soleglad (1972), Williams (1986b), and Williams & Savary (1991). 20. Puravuejovis Williams, 1980. Baja California Sur. 1 sp. See Williams (1980). 21. Pururoctonus Werner, 1934 (contains the only species known from Canada). British Columbia, Alberta; Washington, Oregon, California, Idaho, Nevada, Utah, Arizona, Montana, Wyoming, Colorado, New Mexico, North Dakota, Nebraska, Texas; Baja California, Baja California Sur, Sonora, Chihuahua, Coahuila, Aguascaliente. 32 spp. See Haradon (1984a,b; 1985). 22. Smeringurus Haradon, 1983. California, Arizona, Baja California, Baja California Sur. 4 spp. See Haradon (1983). 23. Vejovoidus Stahnke, 1974. Baja California Sur. 1 sp. See Williams (1980).

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DIPLOCENTRIDAE Pocock, 1893. 24. Bioculus Stahnke, 1968. Baja California Sur. 4 spp. See Williams (1980). 25. Diplocentrus Peters, 1862. Arizona, New Mexico, Texas; all of Mexico except Baja California. 22 spp. There are additional species known from Central America. No comprehensive works on this genus exist, but works covering the known species include those of Stahnke (1970a, 1976, 1981), Francke (1975; 197713, c; 1978b), Sissom (1986a), Stockwell & Nilsson (1987), and Stockwell (1988). Acknowledgment

I thank the following individuals and their respective institutions for the loan of specimens so that I could verify characters and make drawings: Norman Penny and Vincent Lee (California Academy of Sciences), Norman Platnick (American Museum of Natural History), and Herbert Levi (Museum of Comparative Zoology). Howell Daly (University of California, Berkeley) kindly allowed me the use of his microscope. Stanley Williams and Warren Savary (San Francisco State University) graciously provided me with a copy of their unpublished manuscript. Thanks also go to June Steele, Warren Savary, Steven Lombardi, and David Kaplan, who reviewed drafts of the manuscript and provided helpful comments. Publication of this paper is supported by the U.S. Army lMedical Research and Development Command. References Cited

Anderson, R. C. 1975. Scorpions of Idaho. Tebiwa 18: 1-17. Banks, N. 1900. Synopses of North American invertebrates. IX. The scorpions, solpugids and pedipalpi. Am. Nat. 34: 421-427. Birula, A. A. 1908. Ergebnisse der mit Sudvention aus der Erbschaft Treitl unternommenen zoologisthen Forshungreise Dr. F. Werner s nach dem ggyptischen Sudan und Nord-Uganda. XIV. Scorpiones und Solifugae, pp. 121-152. In Sitzungsberichte Akadamie Wisseuschaften Wien, tome 117, heft 2, abt. 1. 1917. Fauna of Russia and adjacent countries. Arachnoidea. Vol. I. Scorpions (in Russian, 1965 English translation by B. Munitz, E. Rabinovitz, ed. Israel Program for Scientific Translations, Jerusalem). Comstock, J. H. 1912 (1913). The spider book; a manual for the study of the spiders and their near relatives, the scorpions, pseudoscorpions, whipscorpions, harvestmen, and other members of the class Arachnida, found in America north of Mexico, with analytical keys for their classification and popular accounts of their habits. Doubleday, Garden City, N.Y. 1940. The spider book; a manual for the study of the spiders and their near relatives, the scorpions, pseuwhip-scorpions, harvestmen, and doscorpions, other members of the class Arachnida, found in America north of ,Mexico, with analytical keys for their classification and popular accounts of their

habits. (revised, ed. by W. J. Gertsch). Cornell University Press, Ithaca, N.Y. Dumont, F. 1986. Contribution a l tude des Score pions de France. Doctoral dissertation, Universitb Paris V (Rene Descartes). Ewing, H. E. 1928. The scorpions of the western part of the United States, with notes on those occurring in northern Mexico. Proc. U.S. Natl. Mus. 73(9): l-25. Francke, 0. F. 1975. A new species of Diplocentrus from New Mexico and Arizona (Scorpionida, Diplocentridae). J. Arachnol. 2: 107-118. 1976. Redescription of Paruscorpiops montanus Banks (Scorpionida, Vaejovidae). Entomol. News 87: 75-85. 1977a. Redescription of Vaejovis gEobosus Borelli (Scorpionida, Vaejovidae). Entomol. News 88: 4551. 1977b. Scorpions of the genus Diplocentrus from Oaxaca, Mexico (Scorpionida, Diplocentridae). J. Arachnol. 4: 145-200. 1977c. The genus Diplocentrus in the Yucatan Peninsula with description of two new troglobites (Scorpionida, Diplocentridae). Association for Mexican Cave Studies Bulletin 6: 49-61. Systematic revision of diplocentrid scorpions 1978a. from circum-Caribbean lands. Special Publications The Museum Texas Tech University 14: l-92. 197813. New troglobitic scorpion of genus Diplocentrus (Scorpionida, Diplocentridae). Entomol. News 89: 39-45. 1982a. A new genus of troglobitic scorpion from Mexico (Chactoidea, Megacorminae). Bull. Am. Mus. Nat. Hist. 170: 23-28. Studies of the scorpion subfamilies Supersti1982b. tioninae and Typhlochactinae, with description of a new genus (Scorpiones, Chactoidea). Association for Mexican Cave Studies Bulletin 8: 51-61 (Texas Memorial Museum Bulletin 28: 51-61). 1985. Conspectus genericus scorpionorum 17581982 (Arachnida: Scorpiones). Occasional Papers The Museum Texas Tech University No. 98. 1986. A new genus and new species of troglobite scorpion from Mexico (Chactoidea, Superstitioninae, Typhlochactini). Texas Memorial Museum, Speleological Monographs 1: 5-9. Francke, 0. F. & M. E. Soleglad. 1981. The family Iuridae Thorell (Arachnida, Scorpiones). J. Arachnol. 9: 233-258. Gertsch, W. J. & D. M. Allred. 1965. Scorpions of the Nevada Test Site. Brigham Young Univ. Sci. Bull. Biol. Ser. 6(4): 1-15. Gertsch, W. J. & M. E. Soleglad. 1972. Studies of North American scorpions of the genera Uroctonus and Vejovis (Scorpionida, Vejovidae). Bull. Am. Mus. Nat. Hist. 148: 551-608. GonzBlez-Sponga, M. A. 1977. Rectificacidn de1 caracter ojos laterales en varios gkneros de la familia Chactidae (Scorpionida) en Venezuela. Acta Biologica Venezuelica 9: 303-315. 1983. Smeringurus, a new subgenus Haradon, R. M. of Paruroctonus Werner (Scorpiones, Vaejovidae). J. Arachnol. 11: 251-270. 1984a. New and redefined species belonging to the Pururoctonus buergi group (Scorpiones, Vaejovidae). J. Arachnol. 12: 205-221. 1984b. New and redefined species belonging to the

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