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COMMITTEE ON PUBLICATION
Dr. GeEorcE E. Linpsay, Chairman

Dr. Epwarp L. KEsseEL, Editor Dr. Leo G. HERTLEIN

eg: mee T8 Li — A bed

OCCASIONAL PAPERS

OF THE

California Academy of Sciences

NOem92i, 129) PaAGesi O20 Ea Gures),, 3 itablest

SCORPIONS OF THE NORTHERN CALIFORNIA
COAST RANGES
(Arachnida: Scorpionida)

By

John T. Hjelle

SAN FRANCISCO

PUBLISHED BY THE ACADEMY
january 26, 1972

COMMITTEE ON PUBLICATION

Dr. George E. Lindsay, Chairman
Dr. Edward L. Kessel, Editor Dr. Leo G. Hertlein

OCCASIONAL PAPERS

OF THE

California Academy of Sciences

Now (92), 959 pages!, 52 £igures, 3 tables’.

SCORPIONS OF THE NORTHERN CALIFORNIA
COAST RANGES
(Arachnida: Scorpionida)

By

John. T. Hjelle
California Academy of Sciences

INTRODUCTION

Until recently, the scorpion fauna of the Nearctic region
was given very little attention by arachnologists, the scor-
pions of northern California having been almost completely
ignored. One of the earliest references to the scorpions of
these ‘areas is that by ‘Wood (1863), in which he“described
several new species of North American scorpions. In 1894,
Pocock listed the various genera of scorpions to be found in
each of the zoogeographical regions of the world. Many of
his observations have since proved to have been in error.

For example, the northern limit of scorpions in North America
is closer to the 50th parallel rather than between the 35th
and 40th parallels. Also, the genus Uroctonus is now known
to extend south into Baja California, Mexico.

In 1900, Banks published a key to 23 species of North
American scorpions, and later, in 1910, published a key to
14 species of California scorpions. Gertsch (1966) gives a
critical discussion of the species included in both of these
keys. .

A work of major significance is Pocock's (1902) synopsis
of the scorpion fauna of North and Central America. Although
for many years a prominent work, the synopsis by Ewing (1928)
of the scorpions of the western United States has been out-
dated by both synonomy and the discovery of many new species.
Hoffmann (1931) gave detailed descriptions of the scorpions
of Mexico. Werner (1935) reviewed the order Scorpionida on
a global basis. Gertsch (1958) reviewed the scorpion fauna
of northwestern Mexico and Baja California, describing three
new species from these areas. The more recent papers by

2 CALIFORNIA ACADEMY OF SCIENCES [Occ.Papers

Gertsch and Allred (1965), on the scorpions of the Nevada
Test Site, and by Gertsch and Soleglad (1966), on the Boreus
group of scorpions, have contributed greatly to our under-
standing of the scorpion fauna of North America.

Wildvams (96 8b),) 1968d)) L970a, L970 by Ao 70e) EoTOdy
1970e) and Williams and Hadley (1967) have described many new
species of North American scorpions. By his recent revision
of the genus Hadrurus, Williams (1970e) has made a prominent
contribution to North American scorpion systematics. Most
recently, Gertsch and Soleglad (personal communication, 1970)
have in press a revision of the genus Uroctonus, and Williams
(1971) has corrected the spelling of the genus Vejovis to its
original form, Vaejovis (likewise Vejovidae to Vaejovidae),
and also the spelling of the genus Broteas to Brotheas.

Because of the lack of collections and knowledge of the
scorpion fauna of the northern California Coast Ranges, and
the uncertain status of many specimens held by various indi-
viduals and institutions, it seemed worthwhile to undertake a
study of the scorpions of this area. The main purposes of
this study were to sample the scorpion fauna of the northern
Coast Ranges of California by selective field work, and to
prepare for publication a comprehensive paper including
taxonomic keys, new species descriptions, differential diag-
noses of all species, distributional maps, species illustra-
tions, and whatever other bionomical information could be
gathered.

It should be mentioned that there are three new species,
authored by Gertsch and Soleglad, known from the northern
California Coast Ranges in addition to those discussed below.
However, since the descriptions of these species are now in
press, they have been omitted from this paper, although they
are in my unpublished master's thesis on which this paper is
based and which is on file at San Francisco State College.

ACKNOWLEDGMENTS

This study is based on scorpions collected by the author
over the past two years and on specimens deposited in numer-
ous other institutions. I take this opportunity to express
my thanks to the following individuals and institutions for
making available important study material: Franklin Ennik,
Bureau of Vector Control, Department of Public Health,
Berkeley, California; Saul Frommer, University of California,
Riverside, California; William E. Ferguson and R. E. Main,
San Jose State College, San Jose, California; Charles L.
Hogue, Los Angeles Museum of Natural History, Los Angeles,
California; Charles R. Smith, Bureau of Vector Control,
Department of Public Health, Redding, California; Robbin W.
Thorp, University of California, Davis, California; Marius
S. Wasbauer, Bureau of Entomology, Department of Agriculture,
Sacramento, California; Thomas S. Briggs, Galileo High School,
San) Prancisco, California; Paul’ H. Arnaud),.ar. ,. Calattomnaua
Academy of Sciences, San Francisco, California; and Mont A.
Cazier and Joe Bigelow, Arizona State University, Tempe,

No.92] HJELLE: NORTHERN CALIFORNIA SCORPIONS 8)

Arizona. I would especially like to thank Torbjorn Kronestedt
of the Naturhistoriska Riksmuseet, Stockholm, Sweden, and M.
Moritz of the Zoologisches Museum der Humboldt-Universitat,
East Berlin, Germany, for arranging the loan of very important
type specimens. Special thanks go to Stanley C. Williams for
his guidance and encouragement throughout this study, to
James R. Sweeney and Joel F. Gustafson for their reading of
the manuscript and for their suggestions and criticisms, to
Robert D. Beeman for his help in the taking of macrophoto-
graphs, and to Willis J. Gertsch and Michael Soleglad who
kindly made descriptions of species recently described by
them available to me prior to publication. Much appreciation
is due Frank H. Koehler, Jr. for making several of the draw-
ings, which allustrate, important) taxonomic; characters. The
following are gratefully acknowledged for their assistance

in the field: Thomas M. Glimme, Benjamin E. Proeres, Frank
H. Koehler, Jr., Carl S. Koehler, Mel Bolander, and Vincent
F. Lee. Finally, I would like to thank my wife, Martha, for
clerical assistance and for her help and encouragement
throughout the term of this study. This study was partially
supported by the National Science Foundation through Research
Grants GB-7679 and GB-23674.

MATERIALS AND METHODS

This study is based on an examination of about 2000
specimens of immature to adult scorpions, mainly from the
northern California Coast Ranges. Most collecting was done
at night with the aid of an ultra-violet light, although some
specimens were collected by rock-rolling during the day.

Each specimen was fixed and preserved by the methods
recommended by Williams (1968a), that is, injection with
Dietrich's fixative and preservation in 70 percent ethyl
alcohol.

All measurements are in millimeters and were taken with
the use of a dissecting microscope equipped with an ocular
micrometer. The measurements used in the paper are, for the
most part, the conventional ones for scorpion systematics as
detailed by Williams (1968b), with the following exceptions:
the width of the palm is taken at its widest point dorsally
between the lateral and inner dorsal keels (or where these
keels would be if absent) (fig. 1); the depth of the palm is
taken at its widest point between the median dorsal and
Peery while looking at the inferior median surface

Gio y)))/2))!s

Macrophotographs were taken on a Wild dissecting micro-
scope equipped with a standard 35 mm. film box, using KB14
film. Illustrations and photographs are based on one speci-
men, those of type specimens being indicated separately with
each figure. Type specimens are deposited, along with all
other material, in the California Academy of Sciences.

4 CALIFORNIA ACADEMY OF SCIENCES [Occ.Papers

RESULTS

For the purpose of this study, the northern California
Coast Ranges are designated as those ranges present in the
area bounded by the southern Monterey County line to the
south, the Klamath Mountains to the north, and the Central
Valley to the east. In addition, records from both the
Klamath and Cascade mountains are included.

The general framework of the Coast Ranges was probably
established about 20 million years ago, although it was not
until about one million years ago, at the beginning of the
Quaternary, that the Coast Ranges region was uplifted and the
sea receded from the Central Valley (Durrenberger, 1968).

The area under study is heterogeneous in many respects.
The northern Coast Ranges are composed largely of sandstones,
shales, and limestones laid down in the Jurassic. In the
more northern areas, particularly in the Klamaths and Cas-
cades, Mesozoic-Paleozoic metamorphic and granitic rocks are
the major geologic units. The central portion of the study
area is most diverse, with combinations of Cretaceous,
Tertiary, and Quaternary sedimentary rocks and some areas with
the more typically northern granitic rock complex.

The flora of this area is also quite diverse, falling into
the Oregonian, Californian, and Nevadan (in part) Biotic
Provinces of Munz and Keck (1965), or into the Pacific Mari-
time, Suwerra Nevadan (frilniwtian Diseriece)!,) and Californian
Biotic Provinces of Schick’ (1965))2" Within ‘these jprovainicesi,
the major plant communities associated with the scorpion
fauna of this area are: Northern Coastal Scrub, Coastal Sage
Scrub, North Coastal Coniferous Forest, Closed-cone Pine
Forest, Redwood Forest, Douglas-Fir Forest, Yellow-Pine For-
est, Mixed Evergreen Forest, Northern Oak Woodland, Foothill
Woodland, Chaparral, and Coastal Prairie.

Life zones from Lower Sonoran to Boreal are represented
in various areas. The climate is also variable. Along the
coast, it is cool, without extreme temperature fluctuations,
while along the eastern slopes there is a greater range in
temperature, being modified locally by topography. Also,
the occurrence of fog is an important climatic factor along
the coast and foothills.

Because of the diversity of geologic, floral, and climatic
conditions, one would expect, and indeed finds, considerable
diversity in the faunal elements.

SCORPION FAUNA

The scorpion fauna of the northern California Coast Ranges
includes representatives from two families, the Chactidae and
the Vaejovidae. The Family Chactidae includes one genus and
a Single species, while the Family Vaejovidae predominates
with four genera, nine species, and three subspecies. The
following is a key to the scorpion taxa now known to occur in
the northern Coast Ranges of California and peripheral areas.

No.9 2) HJELLE: NORTHERN CALIFORNIA SCORPIONS 5

Keygitoy the) Species Of Scorpions) Of. the
Northern California Coast Ranges

2 lateral eyes on each side of carapace;

principal teeth of pedipalp fingers

arranged in oblique, discontinuous rows

(family Chactidae). . . Superstitionia donensis Stahnke
Not with 2 lateral eyes on each side, but

with 3 or 4 lateral eyes on each side;

principal teeth of pedipalp fingers

arranged in a single continuous row (fig. 4)

(fama yi RV aed OV Gael) 2) ely Si el ia Ae) NPC) awe eating 2

Four lateral eyes on each side (fig. 5);
male with swollen bulb on basal part of
aculeus (fig. 6).(genus Anuroctonus). . . 5 ‘
Se Le le RAMUrOCEOnUS phaiodactylus (Wood)
Three lateral eyes on each side; male
without swollen bulb on basal part of
EKORDUI SHO USWA msn tee utes HMAN yesh bs Meera Mave ae eS MUC GM Semler seca Pa Ct LaMar nica 3

With single enlarged dark tooth on inferior

margin of movable cheliceral finger (genus

Hacirurus))jn. “ - e Hadrurus obscurus Williams
Inferior margin oe movable cheliceral finger

without such a tooth, this margin toothless

or with a series of eigen dentenediess | (Eig) 8) ie rs ee 4

Ratio of palm width/palm depth 1.05 or

greater; genital operculum of female not

completely fused longitudinally, free

distally up to 1/4 length; 9 or fewer

muddle) dliamedilac) i(iitg.)3):.. (genus \Uroctonus) ij) 4, 21% 5
Ratio of palm width/palm depth usually 1.00

or less (if greater, either genital

operculum of female completely fused

longitudinally, or male and female with

10 or more middle lamellae, or fixed

finger of chela more than twice palm

Wad ehianiGenush ,ViaeG gOS) ris, ie! ite smelt ulay peat eeeN stelinles veh eau cameras d)

Inferior median keel of metasoma V un-

basanche did steadliye Wes cht onclumreliel « 5 5
Wien emu SOCEONUS glimme i Haeduiap new species
Inferior median keel of metasoma V
branche dias ial yin a a siis ike. dichMioppnsueied Werkelcu: ‘eteehs iamemrenapeh tiie 6

Supernumerary teeth number 7 and 8 on
fixed and movable fingers of chela,
BESDeGEL Velivis. - »- Uroctonus mordax mordax Thorell
Supernumerary teeth number Sivand ion
fixed and movable PARI ESS of eeus
respectively
7s ULOCEOnUS mordax pluridens Hyedilion,, new “subspecies

6 CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

fio Eixed) finger of chela’ twaceron mone ‘ast long
as palm width; both fingers of chela
with elongate distal tooth (fig. 45);
two distinct dark lateral longitudinal
stripes on mesosoma; pectine teeth 13
tomlGr in maler ai toned 4rainy fenallcnny é cee
Vaejovis gertschi striatus Hoedneee new subspecies
Fixed finger of chela less than twice as
long as palm width, both fingers without
elongate distal tooth; pectine teeth 18
Or Moreuine Female, =20- or mMOreo nema len i isnt ane =rsemEs 8

8. Terga of mesosoma with pale band along
posterior margins; teeth of chela
strongly amen MOTGeS Osama mye eee eA
5 : 56 to) WelSrOwas boreus | (Girard)
Terga of mesosoma with dark pattern
continuous to posterior margin over
most of range (in areas detailed in
text, color variable, some individuals
resembling V. boreus); teeth of chela

ecw pool renee in Orn SexeSn eet
SE oP aR I Ags AA ha eee Tk Vaejovis ea eseia Bowens

Family CHACTIDAE
Genus SUPERSTITIONIA Stahnke

Superstitionia donensis Stahnke.
(GAIQUEES i 2i7 Walls Pea S 2) 2)

Superstitionia donensis Stahnke, 1940, p. 102; 1949, p. 243.
GereEsechicw Ali eayelo Gio, “ppl lan) oy ae
Diplops desertorum Mulaik and Higgins, 1944, p. 238.

DIAGNOSIS. Small species, base color yellow to orange-
brown, with an overlying black pattern as follows: Carapace
with irregular dark markings over most of surface; mesosoma
and metasoma dorsally with three distinct black stripes, a
Single median stripe running the full length to the aculeus,
and one on each side laterally, somewhat broken on the meta-
soma; metasoma ventrally with a single dark median line ex-
tending the full length and onto the posterior half of the
last segment of the mesosoma. Metasoma without keels, with
heavy granules only on posterior portion of segment V. Seg-
ments I, II, and III of metasoma broader than long. Pedi-
palps with short fingers; fingers of male with large gap when
fingertips touching. Subdistal teeth of chelicerae usually
minute, two in number; characteristic patch of long, whitish
hairs ventrally along middle of movable cheliceral finger.
Usually six pectinal teeth; genital operculum of female
completely fused longitudinally; genital papillae of male
Visible externally.

TYPE DATA. Syntypes ‘of Superstitionia donensis from the

No.92] HJELLE: NORTHERN CALIFORNIA SCORPIONS 7

Superstition Mountains, Arizona. Depository: Arizona State
University (Stahnke collection). Holotype of Diplops
desertorum from 16 miles east of Tucson, Arizona. Depository:
University of Utah.

DISTRIBUTION. Known from Arizona, New Mexico, Sonora,
Baja California, Nevada, southern and central California.

RECORDS. CALIFORNIA: San Benito County: 3.5 miles west
of the Fresno County line on Road J-1l between Mendota and
Panoche, 14 August 1969 (S. C. Williams, M. M. Bentzien, V.
F. Lee), 2 males.

REMARKS. This record from San Benito County, California,
extends the range of this species northward by about 100
miles. This species is much less common than any others of
the coast ranges.

Family VAEJOVIDAE
Genus ANUROCTONUS Pocock

This genus is represented by a single species which is
easily identified by the presence of four lateral eyes on
each side of the carapace and by the swollen bulb on the
aculeus of mature males.

Anuroctonus phaiodactylus (Wood).
(haigiisssii5i) 6) lS * 6) 49.7) 5:2 «)

Centrurus phaiodactylus-Wood, 1863a, p. 111; 1863b, p. 372.

UsockEonus phiacodactylus) Karsch,, 18/19), ips 102.) (Kraepe lini,
SOY, pple 196-198"

Uroctonus phaiodactylus Banks, 1900, p. 424; 1904, p. 365.

Oncocentrus phaeodactylus PhoKel ly liSSa wy pra Siar

Anuroctonus phaiodactylus Banks, 1910, p. 188. Ewing, 1938,
Pp lk4 ei

Anuroctonus phaeodactylus Pocock, 1893, p. 309. Kraepelin,
SOO pe Pao esi lo 247 psa Hor EMan\,, ho Sie ops
AVAZ—A 05.5) Werner, 19357. 284. Gertsich), 1958, pe 14);
OGD. DP lal 20 Dacaz Nayemay,.1970),, ps 216.

DIAGNOSIS. Medium to large size; base color yellowish
brown to brown. Anterior portion of carapace elevated; four
lateral eyes per group. Inferior median keels of metasoma
distinctly ‘granular on ald, but segment) IV, where obsolete:
Vesicle of telson large; aculeus of male with bulbous swell-
ing at base. Inferior margin of movable cheliceral finger
bears up to four denticles. Genital operculum of male large
and unlike Vaejovis in that the genital papillae may pro-
trude ventrally as well as posteriorly. Pectine teeth number
5 to 8 in females, 7 to 10 in males.

TYPE DATA. Male type of Centrurus phaiodactylus Wood

rom yeah 4) Grom ioniginal jdesicription,,) Ut ahi temnaitony ,7 Mi:
McCarthy, .Esq."). Depository: United States National Museum,
Soul Sjene. 2S

DISTRIBUTION. Known from Nevada, Utah, Baja California,

8 CALIFORNIA ACADEMY OF SCIENCES [Occ.Papers

southern and central California.

RECORDS. CALIFORNIA: Ventura County: Casitas Spriing,
1956 (via Ventura Health Department), 1 male; Quatal Canyon,
Cuyama Badlands, 9 April 1966 (R. R. Montanucci), 2 males.
San Luis Obispo County: 1.3 miles west of Highway 101 on
West Cuesta Road, elevation 2400 feet, 4 April 1970 (J. T.
Hjelle, F. H. Koehler, Jr., B. E. Proeres), 1 female; 2 miles
east of San Luis Obispo, 26 December 1966 (K. Hom), 1 male,
1 female. San Benito County: 3 miles east of Pinnacles
National Monument, elevation 1100 feet, 5 May 1968 (S. C.
Williams), 1 male, 3 females; 1 mile north of Pinnacles
National Monument, 4 May 1968 (B. Butterworth), 1 male, 3
females; Pinnacles National Monument, elevation 1200 feet,

4 May 1968 (S. C. Williams, V. F. Lee), 5 females.

REMARKS. In 1863, H. C. Wood described Centrurus phaio-
dactylus as a new species of North American scorpion (Wood,
1863a). Later, in 1879, F. Karsch amended the spelling of

"“phaiodactylus" to "phaeodactylus" without stating his rea-
sons for doing this eceehi, 1879). In studying Wood's
Original description, and a description appearing in another
journal later in the same year (Wood; 1863b), it is clear
that he definitely intended to use the spelling “phaiodactyl—
us," as the specific name is spelled this way uniformly in
the text. Therefore, Karsch's emendation must be considered
as an unjustified emendation according to Articles 32 and 33
of the International Code of Zoological Nomenclature. As
such, Karsch's "phaeodactylus" must be considered as a junior
objective synonym of Wood's "phaiodactylus." The new records
listed above extend the range of this species northward by
about 100 miles. This species is an obligate burrower, with
only one scorpion in a burrow. The burrows can be readily
identified by their) characteristic Shape’ (fig! \16)i\ the
females probably spend their complete pre-adult and adult
life in and around one burrow, while the male, upon reaching
maturity, appears to leave his burrow in search of a female.
A discussion of the burrowing activities of Anuroctonus
phaiodactylus is given by Williams (1966).

Genus HADRURUS Thorell

A genus of large hirsute scorpions which may be easily
identified by the presence of a single, large, dark, con-
Spicuous tooth on the inferior margin of the movable cheli-
ceral finger. Only one species of Hadrurus has been reported
from the northern California Coast Range.

Hadrurus obscurus Williams.

(Exiguire’s) INS MIS 2)
Hadrurus obscurus Williams, 1970e, pp. 28-30, 61-62.

DIAGNOSIS. Large, hirsute scorpion, base color pale

No.92] HJELLE: NORTHERN CALIFORNIA SCORPIONS 9

yellow with dark pigmentation on carapace and mesosoma; anter-
ior region of carapace light yellow; females with carapace
longer than metasoma V, and with metasoma I broader than long;
inferior median keels of metasomal segments I to III densely
covered with stout bristles; total length averages about 71
millimeters in males, females averaging about 67 millimeters;
pectine tooth counts 34 to 37 in males, 24 to 30 in females.
Sexual dimorphism exhibited by the following characters: male
carapace shorter than movable finger of pedipalp; carapace
longer than metasoma V in males, shorter in females; movable
finger of pedipalp longer than metasoma V in female, subequal
in male.

TYPE DATA. Holotype male and allotype female from 3.5
miles west of the Fresno County line along Road J-1 between
Panoche and Mendota, San Benito County, California, 14 August
1969 and 18 May 1968, respectively (S. C. Williams and par-
ties). Depository: California Academy of Sciences.

DISTRIBUTION. Found in California along the southeastern
base of the Sierra Nevada, across the southern Central Valley.

RECORDS. CALIFORNIA: San Benito County: 3.5 miles west
of the Fresno County line, along Road J-1l between Panoche and
Mendotay, ks May 968 (Sir C. wililvams, Wie Rv Gabel, Ks) €.
Schroen), allotype and 2 paratopotypes, and 14 August 1969
(S. C. Williams, V. F. Lee, M. M Bentzien), holotype and 20
paratopotypes (9 males, 11 females).

Genus UROCTONUS Thorell

This genus can be distinguished from most vaejovids in
that the genital operculum of the female is not completely
fused) Jongitudinally. (in) addition, the: hand is heavy,’ £lat—
tened, and wide, the ratio of palm width/palm depth is usual-
iveegmeates) thanid.05 i(caneibe: as: haghas’ 1 .:45)",- and: there are
nine or fewer middle lamellae.

Uroctonus glimmei Hjelle, new species.
Paqurese107025=29),° 315,449, 523)

DIAGNOSIS. Medium-sized species of Uroctonus closely
related to Uroctonus mordax subspecies. Base color of cuti-
cle reddish brown to brown with few contrasting markings.
Carapace with shallow anterior median emargination; carapace
lightly granular; median eyes slightly anterior to middle of
carapace. Inferior median and inferior lateral keels of
metasoma all distinctly granular; first metasomal segment
broader than long, succeeding segments longer than broad.
Bectinal teeth VOMeoy 2 an mailles 79" tok Mi ane remaies); sicher —
cerae with inferior border of movable finger smooth. Tarsi
each with a median row of at least fifteen spinules on the
inferior surface.

May be distinguished from U. mordax in the following
ways: distinctly smaller size; the lack of dark markings on

10 CALIFORNIA ACADEMY OF SCIENCES [Occ.Papers

most body regions; presence of keels on inferior median sur-
faceo&upalm\ (fig). 315)};) presence Of iventnal kee lion yoatin
(fig. 31); inferior median keels of metasoma not smooth or
obsolete on segments I and II; inferior median keel of meta-
somal segment V not branched posteriorly; dorsal and dorso-
lateral keels end posteriorly in a sharp point.

HOLOTYPE. Maile. Coloration: “Base color of ‘casapaee,
pedipalps, and metasoma light reddish brown; mesosoma and
telson lighter yellowish brown to brown; walking legs lighter
yellow to yellowish brown; carapace with lighter brown areas
posterolaterally and anterior to median eyes; dark contrast-
ing markings on mesosoma absent or very pale, more obvious in
juveniles; metasoma without dark contrasting markings; pec—
tines yellowish white; tips of pretarsal claws light reddish
brown.

Carapace: Anterior margin with shallow median emargina-
tion, with three pairs erect reddish bristles; lateral eyes
three per group, anterior two eyes about equal size, poster-
ior eye smaller; median eyes situated 0.7 mm. anterior to
middle of carapace, each eye 0.2 mm. in diameter, separated
from each other by 0.2 mm.; low median ocular tubercle,
raised 0.1 mm. from carapace (taken from most elevated point) ;
most of carapace finely granular, agranular just behind an-
terior margin.

Mesosoma: Terga 1-6 finely granular, essentially bare,
without obvious dark markings; on some specimens a hint of a
lighter median longitudinal band; tergum 7 with two pairs of
distinct, granular lateral keels, the inner keel projecting
laterally, joining the outer keel anteriorly; intercarinal
spaces finely granular. Sterna agranular except sternum 7
with one pair distinct granular keels and a few scattered
granules; each sternum set with row of reddish bristles on
posterior margin; stigma short, slit-like.

Metasoma: All dorsal and dorsolateral keels granular
except dorsal keels of segment V absent; dorsal keels I to IV
each end posteriorly in sharp upturned point; dorsolateral
keels I to IV each begin anteriorly as distinctly broadened
wing (quite enlarged in some) and end posteriorly in sharp
point; lateral keels granular on full length of segment I
posterior 1/4 segment II, posterior 1/6 segment III, absent
on segment IV, granular on anterior 2/3 segment V; inferior
lateral and inferior median keels distinct, granular on seg-
ments I to V; single inferior median keel of segment V dis-
tinct, granular, unbranched posteriorly; inferior median
keels}\on; segments! ty, Li bi, land) EVicare: sebawath 22) Somes
erect, reddish bristles, respectively; intercarinal spaces
essentially agranular, except segment V with two groups gran-
ules on either side of inferior median keel, each group set
with one erect, reddish bristle.

Telson: Ventral aspect of vesicle irregular, granular
along darker longitudinal bands; this surface with six pairs
of reddish bristles; very subtle subaculear tubercle; acule-
us deeply curved with one pair of reddish bristles ventrally.

Pectines: 12/12 pectinal teeth; fulcra triangular; mid-
dle lamellae with six subequal pieces distal to basal piece;
anterior lamellae hirsute; middle lamellae, fulcra, and

No.92] HJELLE: NORTHERN CALIFORNIA SCORPIONS ieIL

pectinal, teeth hairless on with a few hairs, mainly on basal
area. Sternite between pectines deeply concave anteriorly,
2/3 as broad as long, set with several stout, reddish bris-
Gilesr

Genital operculum: Completely divided longitudinally;
genital papillae visible externally.

Chelicerae: Superior margin of movable finger with five
strong teeth, distal tooth elongate; inferior margin of mov-
able finger smooth, without denticles, with one strong,
elongate, distal tooth. Superior margin of fixed finger with
four strong teeth, proximal two forming compound tooth, dis-
tal tooth elongate; inferior margin rounded, toothless.

Basal segment covered ventrally with fine whitish hairs,
these hairs extending onto inferior margins of movable and
fixed fingers; basal segment dorsally without dark markings.

Pedipalps: Humerus almost three times as long as broad,
anterior surface with several sharp granules along 3/4 of
its length; brachium almost three times as long as broad; all
keels of humerus and brachium granular; hands moderately
heavy and thickened, with various keels developed as follows:
ventral keel lightly granular, directed slightly inward dis-
tally .(£1g. 31); ventrolateral keel, distinctly granular,.ex-
tending distally straight toward superior margin of movable
finger (fig. 31); lateral, dorsolateral, and median dorsal
keels distinct, raised, lightly granular; inner dorsal keel
raised, granular; inferior median surface with two raised
granular keels, forming a shallow groove between them - this
surface otherwise agranular or with very fine granules. In-
ferior margin of fixed finger with six supernumerary teeth;
principal teeth of fixed finger divided into six groups by
enlarged teeth; inferior margin of movable finger with seven
supernumerary teeth; principal teeth of movable finger di-
vided into seven groups by enlarged teeth.

Walking legs: Femora I to IV with double row of granules
extending length of inferior face of each femur, all rows
equally distinct; all tarsi with median claw, immediately
behind which are a pair of spinules followed by a single row
of approximately 15 or more similar spinules, this row
flanked by two pairs of light spines on each side (fig. 10).

Standard measurements: Table l.

ALLOTYPE. Female. Essentially the same as holotype in
color and morphology with the following exceptions: slightly
greater in total body length and most other body proportions;
pectines with fewer teeth (10/11 instead of 12/12); genital
papillae absent; genital operculum divided longitudinally
only along distal 1/4.

Standard measurements: Table l.

PARATYPE VARIATION. Paratypes showed little significant
variation from the descriptions of the holotype and allotype.
Adult males varied in carapace length from 5.2 to 5.6 milli-
meters, while females varied from 5.6 to 6.2 millimeters.
Samples contained adult and juvenile instars, the juveniles
being generally lighter in color and with more contrasting
markings. Little sexual dimorphism occurs. Males and fe-
males have overlapping pectine tooth counts, males varying
from 10 to 12 (predominantly 11) and females varying from 9

12 CALIFORNIA ACADEMY OF SCIENCES [Occ.Papers

to 11 (predominantly 10 and 11). Males can be distinguished
by externally visible genital papillae and a genital opercul-
um which is completely divided longitudinally.

TYPE DATA AND ETYMOLOGY. The holotype, allotype, and 10
paratopotypes were collected 5 miles north of Rayhouse Road,
at the’ junction of Davis and: Cache creeks, Lake’ County,” Cala—
fornia, 15 June 1969 (3.0.7. Hjyelle, M. Bolandex)): “ALL speci—
mens were collected at night under ultraviolet light along a
dirt roadbank with both eastern and western exposures. The
area is characterized by chaparral vegetation and is hot and
dry during the summer months. The holotype and allotype are
permanently deposited in the California Academy of Sciences.
The species is named "glimmei" for my good friend, Thomas M.
Glimme, in appreciation for valuable help in the field.

DISTRIBUTION. Known from coastal Monterey County and
north along the slopes of the inner coast ranges into Mendo-
Cine) County.

RECORDS. In addition to the holotype, allotype, and 10
paratopotypes, an additional seven paratypes were studied.
Paratypes were collected from the following localities in
Californias Mendocino County: 13 miles east of Covelovon
Mendocino Pass Road, 30 August 1969 (J. T. Hjelle, B. E.
Proeres), 1 male; 3 miles up Robinson Creek Road (off High-
way 253), elevation 1000 feet, 28 August 1969 (J. T. Hjelle,
B. E. Proeres), 1 male. Lake County: 5 miles north of
Rayhouse Road, at the junction of Davis and Cache creeks, 9
August 1969) (je°T. Hjyelle, DT. Farris), limale,. i, temateyas
subadult. Napa County: 1/4 mile southwest of the Lake Coun-
ty line on Butts Canyon Road, elevation 750 feet, 11 Septem-
bery1969° (ail Te) Hyelle, 1. Ms. Glimme) oi tmale. |) Stanwsiaws
County: Frank Raines Park, 18 miles west of Patterson, 27
September 1969 (S. C. Williams, J. T. Hjelle, M. M. Bentzien,
W. E. Azevedo), 1 female.

REMARKS. This species occurs in the drier inland areas
of the northern California Coast Ranges along with Vaejovis
gertschi striatus. Most of the specimens were collected at
night along road cuts or areas with little or no grass cover
but with brush cover nearby.

Uroctonus mordax mordax Thorell.
(RGuBesShl=3)Ani7—9V Me sONNs2— 366 SOW sp oie)

Uroctonus mordax Thorell, 1876, p. 11 (original description) ;
LSVC=7i7)" pp L96—1918.... Manni, L676) peu2l2 -2 harsh econo
Pp) LO2-103..0 Keacpelin, (1894) pao Ai E899 ono
Pocock, (1902), p # 309) .) Banks; 719010), ps 14245 1904 ores Gere
L910; pp. L8e, Ess. Borel la). 1908-09, p. 2242 Motes,

EO Zea WLS EWELNG), (O28 piwik .te HOt fman)y rosea.
pp. 402-403. Werner, 1935, pp. 283-284. Mello-Leitao,
19457 -p. 228.0 Henry, 1949) pps 120-125... \Gertsch &
Allred, 1965, ps. 4. Gertsch & Soleglad; 1971, in press
(personal communication). Parrish, 1966, pp. 12-13.
Diaz WNajenayy LOO; pe llGr

No.92] HJELLE: NORTHERN CALIFORNIA SCORPIONS 13

In the years which have elapsed since Thorell first des-
cribed Uroctonus mordax, all dark scorpions from the northern
California Coast Ranges that bore some resemblance to U. mor-
dax have been identified as that species. Thorell's original
description (1876) was limited to a few sentences both in
English and Latin, but later in the same year, he published
a more extensive description of U. mordax. The measurements
given in the later paper (none were included in the original
description) were made in an obsolete style and did not seem
wholly reliable. Also, the sex of the holotype was not men-
tioned in either the original or later description.

DIAGNOSIS. Medium-sized, robust, dark scorpion with
thick, swollen pedipalps. Base color of cuticle dark reddish
brown to brown, with contrasting patterning on most body
regions; carapace with deep anterior median emarginate notch;
carapace heavily granulated; median eyes slightly anterior to
middle of carapace; first metasomal segment broader than long
succeeding ones longer than broad; inferior lateral and in-
ferior median keels of metasoma granular, variable in appear-
ance; pectine teeth 8/9; lower margin of movable finger of
chelicerae with four and five denticles on right and left
chelicerae, respectively.

RESESCRIPTION OF FEMALE LECTOTYPE. Coloration: General-
ly dark reddish brown with intricate contrasting markings on
Carapace, mesosoma, metasoma, and telson. Pedipalps, anteri-
or 3/4 carapace, and metasoma dark reddish brown, darkest
along keels and areas of heavy granulation; chelicerae with
dark markings on dorsal basal segment; carapace with darker
markings radiating in a pattern anterolaterally from median
groove, base color lighter on posterior 1/4; mesosoma brown
with essentially same lighter pattern on each segment (except
segment 7), consisting of a pair of oval spots along midline
and a series of spots and elongate areas extending laterally;
metasomal segments I to IV with dorsal "arrowhead-like"
markings, the point facing posteriorly; sterna of mesosoma,
the pectines, and genital operculum pale brownish yellow;
telson light reddish brown with darker longitudinal bands, one
on each side of ventral midline, one on each lateral side, and
one median dorsal band.

Carapace: Anterior margin with deep median emargination,
with two pairs of erect reddish bristles (others probably
lost); median ocular tubercle high, 0.2 mm. above surface of
carapace (taken from most elevated point); median eyes situ-
ated 0.6 mm. anterior to middle of carapace, each eye 0.25
mm. diameter, separated from each other by 0.4 mm.; lateral
eyes three per group, anterior two equal in size, posterior
eye smallest; entire carapace finely to coarsely granular,
coarser granules in pattern from middle of carapace outward
to lateral eyes.

Mesosoma: Terga finely granular, essentially bare, with
a few erect bristles along posterior margins; tergum 7 with
two pairs of granular lateral keels posteriorly. Sterna
smooth, with erect reddish bristles on each segment laterally,
finer whitish bristles in rows across each segment on anterior
margin, middle of segment, and posterior margin; one pair fine
granular keels posterolaterally on sternum 7.

=

14 CALIFORNIA ACADEMY OF SCIENCES [Occ.Papers

Metasoma: Dorsal and dorsolateral keels granular, den-
tate to serrate in appearance, on segments I to IV; dorsal
keels absent on segment V; dorsolateral keels granular on
segment V; lateral keels weakly granular on posterior 1/2 of
I, as row of granules on posterior 1/6 of II, as two or three
granules on posterior end of III, absent or up to four gran-
ules on IV, showing as irregularly spaced granules on anter-
ior 1/2 of V, most prominent along anterior 1/6; inferior
lateral keels obvious on all segments, raised to crenulate on
I, crenulate on II and III, granular on IV and V, appearing
serrate to dentate; inferior median keels weakly crenulate,
but not obvious, on segments I and II, crenulate on III,
crenulate to granular on IV, single median keel granular on
V, this keel branching laterally at distal end. Segments I,
Iv, and V widest in middle, segments II and III widest
slightly posterior to middle; metasomal segment I broader
than long, segments II to V longer than broad.

Telson: Aculeus with moderate curve, 1/2 vesicle in
length; vesicle wider than metasoma V, equals metasoma IV in
width; slightly concave above, ventral surface irregularly
covered with small granules, granules darker, larger, and
more conspicuous at basal end. Subtle subaculear tubercle;
ventral surface of vesicle with seven pairs of reddish bris-
tles; aculeus with one pair of hairs ventrally.

Pectines: 8/9 pectinal teeth; length of sternum between
pectines slightly more than twice its breadth, shallowly
concave; anterior lamellae of three segments, basal sclerite
longest; middle lamellae composed of basal sclerite and six
rounded sclerites, each rounded sclerite with one erect,
reddish bristle, a few sclerites also with one smaller bris-
tle; fulcra subtriangular, each with one or two erect, red-
dish bristles.

Genital operculum: Divided longitudinally along distal
1/4; genital papillae absent; each valve with two elongate,
reddish bristles along lateral margin, one shorter bristle
on posterior margin, and one short bristle on medial margin.

Chelicerae: Superior margin of right movable finger with
five strong teeth, distal tooth elongate; inferior margin
with elongate distal tooth and bearing four small, evenly
spaced denticles. Superior margin of right fixed finger with
four strong teeth, the proximal two forming compound tooth,
distal tooth elongate; inferior margin rounded, toothless.
Left chelicera the same, with the following exceptions:
first subdistal tooth of superior margin of movable finger
appearing as a small extra tooth on the elongate distal tooth
(fig. 7); inferior margin movable finger with five small
denticles (fig. 8). Basal segment and both fingers covered
entirely with brush of fine, whitish hairs on ventral surface.

Pedipalps: Humerus slightly more than twice as long as
broad, anterior surface with 5 to 6 sharp granules; brachium
about twice as long as broad(including projecting spurs) ;chela
heavy and thickened, with various keels developed as follows:
ventral keel absent; ventrolateral keel pronounced and granu-
lar, curved inward distally; (fig. 32) lateral keel obsolete
or indicated by faint dusky markings or fine scattered gran-
ules; dorsolateral keel solidly raised, smooth; median dorsal

No.92] HJELLE: NORTHERN CALIFORNIA SCORPIONS BS

keel raised only very slightly, with occasional granules,
indicated primarily by dusky markings; inner dorsal keel
raised and granular; inferior median surface with one obso-
lete keel, indicated only by faint dusky markings, this sur-
face irregularly covered with granules. Inferior margin of
fixed finger with seven supernumerary teeth; principal teeth
of fixed finger separated into seven groups by enlarged
teeth; inferior margin of movable finger with eight super-
numerary teeth; principal teeth of movable finger separated
into eight groups by enlarged teeth; principal teeth of each
finger aligned in single continuous row.

Walking legs: Femora I, II, and III with double row of
granules extending the length of inferior face of each femur,
the more inferior row being larger and most obvious; femora
IV showing only a short row of granules along distal 1/4; all
tarsi with median claw, immediately behind which are a pair
of spinules, followed by a row of five to seven similar
Sspinules (fig. 9).

Standard measurements: Table 2.

TYPE DATA. Female lectotype presumably Een oan Ehnan—
cisco, California." Depository: Naturhistoriska Riksmuseet,
Stockholm, Sweden (vial number 59b). Information on the
Original label reads as follows: "Uroctonus mordax Thor.,
SCOmpAO/; MAME Sn SCDE. EUSen Su Hrancrsco, Cala forn. a Mert hicg
There are two males in the same vial, one subadult, the other
a young juvenile. The question of whether or not Eisen
actually collected these specimens in San Francisco is still
open to speculation, although Banks (1904) lists another San
Francisco record. At the present time, it seems unlikely
that U. mordax mordax or U. m. pluridens are now to be found
in the northern end of the peninsula, Since recent efforts to
collect here have been unsuccessful.

In a second vial (59a), there are four adult females.

The original label reads as follows: "Uroctonus mordax Thor.,
America: Guatemala, Eisen." Research on Eisen showed that

he arrived in the United States from Sweden in 1873 and did
not travel’ to Guatemala until 1882, several years after
Thorell"s description of U. mordax. Being that the specimens
in this vial definitely belong to U. mordax mordax, and that
this subspecies does not extend south of California, it seems
most likely that Eisen mistakenly gave Thorell the wrong
locality data, resulting in this spurious Guatemalan record.

DISTRIBUTION. Northern California and Oregon (figs. 50,
Siac

RECORDS. OREGON: Lane County: 3.5 miles east of Oak-
Budge, 25 April 1970 (RR. (M2. Winokur),, 4 females.) Wasco
County: 8 miles west of Rowena along U. S. Highway 80N, 16
June 1968 (J. Davidson), 2 males, 1 female. CALIFORNIA: Del
Norte County: Junction of Dunn Creek and the East Fork of
thew llinousi River, -Ousune L960) (2. Rs Hag) 2 males... ‘Siis=
kiyou County: Yreka, 2 May 1969 (J. R. Grisham), 1 male;
12.7 miles north-northeast of Happy Camp, elevation 2000 feet,
(Oe vUnee IG 9 (Re Aw eG Re Re sonel Ling) 7.2 emale, 3 tema lies:
Humboldt County: Willow Creek, 25 August (year not given),
(N. Elmore), 1 male; Williams Grove, 3 July 1948 (S. F. Wood),
1 female; 1 mile northeast of Willow Creek, elevation 590 feet,

16 CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

28 April 1968 (W. Strickland), 1 female; Orleans, Hoopa,
Wildow Greek Area, 1969 “(W. Strickland)),, 2 males, 15 females,
8 miles south of Honeydew, 28 September 1969 (P. Wilson), 1
female. Trinity County: Junction of Mumbo Creek and Trinity
River (East Fork near Clair Engle Lake), 30 May 1965 (G.
Brooks), 1 male; 8 miles south of Highway 36 on Van Duzen
Road, elevation) 3000" feet, 29 August 1960 (or. Hyciley ar
E. Proeres), 2 females; 5 miles northeast of Zenia on zZenia
Roady ‘ellevativon 4000 feet, 29 August 1969) (olen nyelle ae.
ES 'Proeres), 1 male, 8 females. “Shasta County: “Hat Greeks
Ranger Station, 7 May 1965 (R. R:. Pinger). T female Biedey
25 May 1967 (collector unknown), 1 male; 2 miles north, 7
miles east of Shingletown, elevation 4000 feet, 26 April 1968
(Smith and Hazeleur), 3 males. Tehama County: 32 miles east
of Covelo on Mendocino Pass Road (toward Paskenta), elevation
5200teet "30 August 1969 (oc. Toy hyeilic Bon Precres) lame.
females Mild Creek 7 duly 1970) "(Re Dietz, Pe RUC) ie en eee
Mendocino County: Albion Ridge Road, 9 miles south of Albion
Road, 18 June 1969 (F. J. Radovsky), 2 females; Ukiah, 24
January LG2 (Tf. S.) Briggs), 2 females, U2 miles nor eawesie

of Cloverdale on Highway 128, elevation 400 feet, 28 August
1969 (dS. T. Hyelte, Bee. Proeres) I male, i female same
east of Covelo on Mendocino Pass Road, elevation 4000 feet,
30 Augusit® 1969! (ae. Te Hyelle:” BL yh 2 eProeres)iyeaL2 alates asian
females; Riley Ridge, Hopland Field Station, elevation 1900
feet, 8’ May 1970 (M. M. Bentzien), 5 males, 2 females; Men=
docino Woodlands Camp, 15 November 1970 (J. R. Gabel), 1 male.
Lake County: 3.15 miles north of Soda Creek Camp, Mendocino
National Forest; 9 duly?) (K- Hom, Vor. Lee), 2) females are
mile north Glen Eden School, Scott Valley, 17 July 1965 (K.
Hom), 1 female; 5 miles north of Rayhouse Road, at junction
of Davis and) Cache ‘creeks,’ ellevation, 900 feck, (>) dune ibe
(jo, Tee hjelve, MA Bollander)”, 2 males, (3, females ; ;abouims
miles east of Bear Creek Ranger Station on Bear Creek Road,
elevation ss00) feet, si) August: 1969 (Gna ijeisie Besar
Proeres), 6 males, 5 females; 2 miles east of Bear Creek
Ranger Station on Bear Creek Road, elevation 2300 feet, 31
August A969 (Gl Te Hveitel Boake Proere’s 7) i female, mscile auee
2400 feet up Mt. Konocti on Konocti Road, 9 September 1969
(2° TR? Hjelle roms) Glamme)”,* 4 males; 2 females > one hacitende
Springs Road, 5 miles southwest of intersection with Bell
Hill’ Road, “elevation W600" feet, 9 August L969 (ol Toone,
T. M. Glimme), 2 females; 1/4 mile southeast of Highway 175
on Adams Springs Road, elevation 2600 feet, 11 August 1969

(J. T. Hjelle, T. M. Glimme), 1 female; 1/4 mile south High-
way 175 on Anderson Springs Road, elevation 1300 feet, 11
August’ 1969 8(o 9 T. Hielle, T.°9M. Glaimme) , 1) male, 3) fematest
1 mile east of Loch Lomond on Siegler Canyon Road, elevation
3000 feet, 11° August 2969 (3. T..Hjelle, T. M. Glimme), £
male, 5 females; Rattlesnake Island, Clear Lake, elevation
1350) feet, t2yAuGgust 1969= (oO. Tt hjetle on: Mo Glaimme) sy io
males, 12°females: Golusa County: Erom 2° to 5 miles along
unnamed road connecting Letts Valley Road and Twin Valley
Road, elevation 4000 feet, 31 August 1969 (J. T. Hjelle, B.
EB. Proeres), 7 mates, 7 females. Woonoma County: 72 un) Siciateis
Woods," between Forestville and Guerneville, 30 November 1968

No.92] HJELLE: NORTHERN CALIFORNIA SCORPIONS iL 7/

(S. Bridge), 1 female; Summerhome Park, Russian River, 13
August 1969 (L. Lagomarsino), 1 female; 1/2 mile off Highway
116 on road connecting Highway 116 with River Road (approx.
1.8 miles SouthwestJ/o£ Rio Nido), 11 October 1969 (co. T.
Hyedakey) 7B).snt 3) Peoerelsi)i,. 2) iImalies;ap2'itemales.) (Napa ‘County:

1/4 mile southwest of the Lake-Napa County line on Butts
CanyonjpRoad,; elevation: i750; ereet; ell (September 19169) 3(d/. Wri
Hyelle, TeeM., Glimme), 2: males... “Yolo. County:) Monticello Dam
Spi llway),/0, November 1967 (F.! Ennik), 1:male.) Marin’ County:
near entrance to Muir Woods, 10 January 1964 (V. D. Roth,

P. R. Craig), 1 male; Novato, 11 December 1964 (H. Mathis),

1 female cast skin; White's Estate, 3 March 1965 (K. Hom), Ll
Maver tahong, Bolianas, Road.,{713) Apri: 1965, A(Tew<See Briggs). /3
males, 1 female; 2 1/2 miles west of Alpine Dam, 7 July 1965
(K. Hom), 1 male; near Tiburon, 11 December 1965 (K. Hom, T.
S. Briggs), 2 males, 3 females; 1/4 mile north of Laurel Dell,
Mt. Tamalpais, 8 September 1969 (M. M. Bentzien), 1 male; 1
mile south of Fairfax on Bolinas Road, elevation 200 feet, 16
September 1969 (J..T. Hjelle, T..M. Glimme), 3 males, 8 fe-
males; 5 miles south of Fairfax on Bolinas Road, elevation
600 feet, 16 September 1969 (J. T. Hjelle, T. M. Glimme),

4 males, 13 females; along fire trail to Kent Lake,
elevation 50 feet, 18 September 1969 (J. T. Hjelle, F. H. &
C. S. Koehler), 25 males, 26 females; Lily Lake, north of
Alpine Reservoir, 1 December 1969 (J. Mackey), 1 female; near
Lily Lake (just north of Alpine Lake), 18 April 1970 (M. M.
Bentzien), 2 males, 3 females; Phoenix Lake, north shore, 2
May 1970 (J. Collins), 1 female; San Rafael, 21 May 1970 (W.
L. Blackwell), 1 female; 1/4 mile north of Lake Lagunitas, 17
September 1970 (A. D. Bacon), 1 female; south shore of
Phoenix Lake; 1: Octeber-1970. (3. T:; Hjelle);, lomale, 5 fe-
males. Solano County: 2.2 miles south of Cordelia (Cerada
Raneh ey 28 March l96i) (Re. Ee Doty) 7a@de mates, Ba:temaliest) dk15
miles, south of) Cordelia; 3¢Mayii1961. {(R: i Ee: Doty),52 males;
on Cordelia Road, 0.9 miles north of junction with Pittman
Road 9 shebruary 1964e(Ds Lei Dailey; P.@R.o€raig);:1 female;
5 miles southwest of Winters, 19 July 1964 (collector un-
Known) pcedomales vs Contra Costa: County: -yCastlewRock: Park;%5
miles west of Mt. Diablo, no date (F. Ennik), 3 males, 4 fe-
males; Lafayette, May 1958 (R. F. Peters), 1 female; El
Cerrito, November 1959 (W. Anderson), 1 female. Alameda
County: 4Oakland,;*Maytl957. (Gl Salter); 11 female; oskyiine
Boulevard, 0.1 mile south of intersection with Colton Road
anGdgSnake Road, GyOcEobers 1962 (Bi R.oCralg),2D.2 Les Darley) v2
male; Berkeley Hills, Marlborough Terrace below Grizzly Peak
Bivids ,;0241Februaryel9632(P. (RebiCrailg, Dich. eDavlley) mila te-
male; near Niles, 3 January 1964 (V. D. Roth), 1 female;
OakVand pl cAprily 9630 (Weidt Hott) Vemate; sSunol Regional
Pawks re Sdanuary 7 1965.\(Ke (Hom; ls) S. Buiggs) 7 .lemaleyulste-
male; Sunol Regional Park, 7 August 1965 (K. Hom), 1 female;
Oakland, 30 August 1965 (W. W. Sampson), 1 female with 29
-first-instar young; Berkeley Hills, Strawberry Canyon, 17
ApEIL 1966 )(Piy.R. Craig); 2. females; northisideso£k Albany
Hill, Albany, 29 March 1968 (F. Ennik), 8 males, 3 females;

18 CALIFORNIA ACADEMY OF SCIENCES [Occ.Papers

Montclair Area, Oakland, 20 September 1968 (D. Pomerantz), 1
male. Santa Clara County: Gilroy Hot Springs, 13 February
1966 (R. W. Cahill), 2 males; Uvas Canyon County Park, 26
February 1966 (K. Hom), 1 female; Arroyo Mocho, Mt. Hamilton,
28 April 1968 (collector unknown), 1 female; Alum Rock Park,
8 May 1968 (R. Pisciotto), 1 female; Alum Rock Park, 30
January 1969 (J. Smith), 1 male. San Mateo County: Redwood
Gitty ;ol2) August 19518). (De) Wee Ozenbaugh)s, |i eremadtie:

REMARKS. Other specimens which were examined showed
Itteleljsvgnifican’ variation in) most characters) Aswa supe
species, U. mordax mordax ranges from 20 feet to over 7000
feet above sea level. In the past, arachnologists tended to
associate this species primarily with the redwood communities.
However, as would! be expected from les) extensive mange, Uj.
mordax is found in other plant communities as well - from
coastal scrub to predominantly pine areas, to the more un-
likely inner coastal chaparral areas. Their relative abun-—
dance in the latter areas seems, from collection records
available, to be much lower than in the other areas mentioned.

A certain degree of genetic isolation appears to manifest
itself in three characters, these being color, the relative
numbers of pectinal teeth of both sexes, and the number of
denticles on the inferior margin of the movable cheliceral
finger.

Various populations, particularly from the inner coastal
areas east of Covelo, Mendocino County, have a predominance
of individuals of a lighter color than is normally the case
in U. m.mordax. Lighter individuals are present in) mest) orcker
populations also, but are a minority. The inner coastal
ranges at the edge of the central valley represent the
fringes of U. mordax distribution, thus these lighter popula-
tions may be a result of habitat differences or of some de-
gree of ecological isolation.

In the case of pectine tooth counts, as a subspecies,
those of the female vary from 16 to 23, while those of the
male vary from 22 to 29. Generally, these counts seem to
form a mode at about 20 in the female and at about 25 or 26
in the male. However, within these ranges, various popula-
tions have different modal tendencies - toward either high
or low modes, or toward single, double, or even triple modes.
For example, in an area in Glenn County, California (fig.
35a), females ranged from 17 to 21 with two modes, one at
18, the other at 20 (based on 21 specimens), while males
Kanged: £rom 22) £027, also with) two modes, one at 24) the
other at 26 (based on 19 specimens). At Bootjack Camp,

Marin County, California (fig. 35b), females ranged from 17
to 23 with a single mode at 19 (based on 41 specimens), while
males ranged from 24 to 28 with a large mode at 24 anda
smaller mode at 26 (based on 40 specimens). In a population
from Sonoma County, California (fig. 35c), females ranged
from 18 to 22 with modes at 18, 20 and 22 (based on 45
specimens), while males ranged from 23 to 27 with a single
mode at 26 (based on 30 specimens). Finally, from Kent Lake,
Marin County, California (fig. 35d), females ranged from 16
to 21 with a single mode at 18 (based on 26 specimens), while
males ranged from 24 to 27 with a single mode at 25 (based on
23 specimens).

No.92] HJELLE: NORTHERN CALIFORNIA SCORPIONS 19

These data tend to show that, though there is a genetic
continuum which maintains the range of variation for this
species character, there is sufficient geographic or ecologi-
cal isolation between the populations to allow the formation
of distinctly different subpopulation tendencies.

The third character under consideration is that of the
number of denticles present on the inferior margin of the
movable cheliceral finger. For many years, U. mordax, as a
species, was diagnosed as having five denticles on this sur-
face. However, this character is variable within any one
local population and is even more variable for the sub-
Species, U. m. mordax, as a whole. The range of variation
for this character, based on analysis of several hundred
specimens from the entire range, was 0 to 7 denticles with a
mean Of 4.2. Most populations showed values that clustered
around this over-all mean, for example: in the Fairfax Area
of Marin County (fig. 36a), the range was 2 to 6 with a mean
of 4.1, and in the Bear Creek Area of Lake County (fig. 36b),
the range was 3 to 6 with a mean of 4.3. However, various
populations showed quite different tendencies, for example:
13 miles east of Covelo, Mendocino County (fig. 36c), the
range was from 0 to 4 denticles with a mean of 2.0 (this
again may be due to habitat differences or ecological iso-
lation along the periphery of the distribution), and on
Rattlesnake Island, Clear Lake, Lake County (fig. 36d), the
range was from 0 to 5 with a mean of 3.3. There were few
scorpions on the island, so the main factor involved in
altering the typical mode would appear to be genetic drift
in a small population. Geographic isolation may also be a
factor, since the island has been isolated from the mainland
for several thousand years.

Uroctonus mordax pluridens Hjelle, new subspecies.
(Figures 37-40, 50, 52.)

In examining various specimens of Uroctonus from coastal
northern California areas, it was noticed that several speci-
mens differed from typical U. mordax in the presence of a
greater number of supernumerary teeth on the fixed and mov-
able fingers of the chela. Because of the homogeneity of
this important and otherwise nonvariable character in large
clearly defined geographic areas, I consider this group a
subspecies of U. mordax.

DIAGNOSIS. Essentially the same as U. m. mordax except
for the following differences: mesosoma more of a solid dark
brown, the characteristic U. m. mordax pattern of lighter
spots less obvious; metasomal segments I to IV with dorsal
“arrowhead-like" markings less distinct than on U. m. mordax,
absent on some individuals (on the allotype these are faintly
marked on segments I and II, absent on III and IV); length
of humerus much greater than twice its width; length of
brachium much less than twice its width; 8 and 9 supernumer-
ary teeth on the fixed and movable fingers of the chela,
respectively.

20 CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

Standard measurements: Table 3.
TYPE DATA. Holotype male from near Corralitos, Santa
Cruz County, California, 8 March 1969 (J. Smith). Allotype

female from Scotts Valley, Santa Cruz County, California,
5 October 1968 (G. Bianconi). Depository: California Acade-
my of Sciences.

DISTRIBUTION. Santa Clara and Santa Cruz counties, Cali-
fornia.

RECORDS. California: Santa Clara County: Near Loma
Prieta Weather Station, 10 November 1968 (J. Smith), 1 female;
Summit above Los Gatos, 4 March 1969 (3.. Smith), 3 males, 2
females;, Alum Rock Park, 8 March 1969) (a. Smith) ,2l male; a
female; Uvas Meadows, near Sveadahl, 15 March 1969 (J. Smith),
1 male, 1 female; Summit above Los Gatos, just off Summit
Road, 204 March 1969... Smith), Jofemailie;) Mess Char liemRoad,

13) Apri 1969) (Ri MeGehee). 74 malie; Lomay Pre tay, 9) November
L969eKd.* Smath) ,.2 tCemakes.4i Santa Cruz County: 720i miles
SOUEn! Of Holy City? dO. April 96 8in(AC ReeiGae lo qilsy)) ale eemarer,
Big Basan State) Park, 28. Apri Wol96 8: (Ran omShe imam) pa
female; 10 miles southwest of Saratoga on Highway 9, 27 March
1969 (Bob Agnoletti), 2 females; Camp Loma, 23 November 1969
(J. Smith), 1 male, 1 female.

Uroctonus mordax (mordax x pluridens hybrids).

In the Alum Rock Park area of Santa Clara County, Cali-
fornia, U. mordax mordax appears to hybridize with U. mordax
pluridens. These individuals may be recognized by the
presence of 7 and 9 supernumerary teeth on the fixed and
movable fingers of the chela, respectively.

RECORDS. Santa Clara County: Alum Rock Park, 3 March
1970 (R. P. Swartzell), 1 female.

Genus VAEJOVIS Koch

This genus can be distinguished from other genera of the
family Vaejovidae through the following combinations of
characters: the genital operculum of the female of most
species is completely fused longitudinally; the inferior mar-
gin of the movable cheliceral finger is completely smooth, or
crenulate, or with several small denticles; there are three
lateral eyes per group; the ratio of palm width/palm depth is
usually 1.00 or less (if greater, either the males and fe-
males each have 10 or more middle lamellae, or the fixed fin-
ger of the chela is more than twice the palm width).

Vaejovis gertschi striatus Hjelle, new subspecies.
(Figures 41-45, 49.)

DIAGNOSIS. Differs from Vaejovis gertschi gertschi

No.92] HJELLE: NORTHERN CALIFORNIA SCORPIONS 2p

Williams in the following characters: females less than 30
millimeters in total length; pectine teeth 13 to 16 in males,
My jo, 24, am. females (eather than; 16to 1:8 ain; males 44 to: 6
in females); subtle subaculear tubercle in the male; two
distinct dark lateral longitudinal stripes on mesosoma. Base
color of cuticle brownish yellow with contrasting dark mark-
ings on most body regions. Carapace with median emarginate
notch; inferior median and inferior lateral keels of metasoma
all distinctly developed, these serrate, crenulate, or inter-
mediate. Chela elongate, palm with lightly crenulate keels
medially and smooth keels dorsally and laterally (fig. 45);
fixed finger longer than palm, at least twice palm width,
terminal tooth on each finger elongate. Chelicerae with
inferior border of movable finger smooth.

TYPE DATA AND ETYMOLOGY. The holotype, allotype, and 59
paratopotypes were collected at Frank Raines Park, 18 miles
west of Patterson, Stanislaus County, California, 27 Septem-
bea 969 ..0S.. C.jWiddliaams, Jefe; Hpelle,;:M.wiM. Bentzien, W:

E. Azevedo). All specimens were collected at night under
ultraviolet light along a dirt roadbank and on a hillside
with a northwest exposure and light grass cover. The holo-
type and allotype are permanently deposited in the California
Academy of Sciences. The subspecies is named "striatus"
because of its characteristic dark lateral stripes.

PARATYPE VARIATION. Paratypes showed little significant
variation from the descriptions of the holotype and allotype.
Matesiwvanied in total length from 11.8 to 22.1 millimeters
while females varied from 12.5 to 30.0 millimeters. Males
Varied: in carapace length from 2.1 to 2.9«millimeters while
females varied from 2.6 to 4.0 millimeters. Samples con-
tained both adult and subadult instars but younger instars
were totally lacking. Little sexual dimorphism occurs.

Males and females have overlapping pectine tooth counts.
Males can be distinguished by the presence of genital papil-
lae and by having the genital operculum completely divided
longitudinally.

DISTRIBUTION. Known from the slopes of the inner (east-
ern) coast ranges from San Benito County to Mendocino County.

RECORDS. In addition to the holotype, allotype, and 59
paratopotypes, an additional 104 paratypes were studied.
Paratypes were collected from the following localities in
California. Mendocino County: 3 miles north of Highway 253
along Robinson Creek Road, elevation 1000 feet, 28 August
1969 (J. T. Hjelle, B. E. Proeres), 2 males, 1 female. Lake
County: 4 miles east of Kelseyville on Konocti Road, Mt. Kon-
octi, elevation 2800 feet, 9 September 1969(J. T. Hjelle, T.
M. Glimme), 1 male; Kugelman Ranch Road, Lower Lake, elevation
1200 feet, 1 September 1969 and 7 September 1970(J. T. Hjelle,
T. M. Glimme), 17 males, 3 females, and 8 males, 1 female,
respectively; 6 miles north of Upper Lake (Witter Springs), 2
February 1970 (T. Toren), 1 female; Bachelor Valley, 12
December 1970 (T. Toren), 3 females; approximately 5 miles
north of Rayhouse Road, at junction of Davis and Cache creeks,
15 June 1969 (J. T. Hjelle, M. Bolander), 8 males, 17 fe-
males, and 9 August 1969 (J. T. Hjelle, T. Farris), 13 males,
10 females. Napa County: 1/4 mile southwest of the Lake

ae CALIFORNIA ACADEMY OF SCIENCES [Occ.Papers

County line on Butts Canyon Road, elevation 750 feet, 11
September 1969 (J. T. Hjelle, T. M. Glimme), 9 males, 6 fe-
males; 3 miles south of junction of road to Pope Valley on
west side of Lake Berryessa, elevation 550 feet, 11 November
NI6I (Voor. Lee) /el kemales -(Sittanistaus (County:n “Del! Puerue
Canyon, 4 April 1970 (W. E. Azevedo), 1 female, 2 subadults.
Additional specimens have been recorded from the following
localities: Solano County: Vacaville, 29 October 1964
(AN) Mendes), 12 femadies. * Contra’ Costa County :)) (Walnuey Grecia,
4 June 1956 (Leo Farr), 1 male; Concord Area, 30 August 1965
(Fv Ennik))) Ll males Me. (Diablo, sh] April io 65° n(R: PRo i Pakacgena,
R. W. Pinger), 1 male. San Joaquin County: Stockton, 9
April 1970 (Walt Bauer), 1 female. Stanislaus County: la
Grange), 32 gully (19169 (Reyes Allien), 1! tkemad ell /7 4 maslemciaisit
of Frank Raines Park, elevation 1000 feet, 3 April 1970
(A. Dean Bacon), 1 female. San Mateo County: San Carlos
(burned chaparral hillside), 20 December 1970 (C. Rees), 1
male. San Benito County: Pinnacles National Monument, 25
AUGUSEMLIG So (Ee Ennwk) 2) males

REMARKS. This subspecies appears to be relatively com-
mon in the drier areas of the northern California Coast
Ranges, particularly on the slopes of the inner ranges. Most
of the specimens were collected at night along road cuts and
in areas with light grass cover. Several specimens have been
collected in houses.

Vaejovis boreus (Gasca)
(Figures 46, 51.)

Scorpio (Telegonus) boreus Girard, 1854, p. 257.

Vejovis boreus Banks, 1900, p. 424; 1910, pp. 187, 189.
ComsEOC 1 Ok ts (LON Si pie Sls S4 Ore pi Sa | Webstecrs,
HOA Sips 248, Chamberlin; 19/247 i pew 64s, sKumata elo ia
Pen28en, Werner, 1935), p- 2825 \ Stahnker,) 1940 iy toe ieesOnee
Gereschiy 1958) ip.) Gs) Gertsch es Allreds| 1965 pia ore
Gertsch & Soleglad, 1966, pp. 6-14.

Vaenovaiswboreus. Ewing, 92'8:,0 pps) Or, 2)

DIAGNOSIS. Medium-sized scorpion with base color pale
yellow to orange-brown. Carapace with V-shaped patch at
median eyes, with dark bands running to lateral eyes;
anterior portion of interocular triangle pale. Mesosomal
terga 1 to 6 darkly colored, each with a series of yellow
spots extending laterally and each with a transverse pale
stripe along the posterior margin. Carapace finely to
coarsely granular. Metasoma I to III with inferior median
keels essentially obsolete, indicated by diffuse, brown
stripes; metasoma V longer than carapace in both sexes.
Pectine teeth 18 to 23 in females, 25 to 31 in males; middle
lamellae with two rows of sclerites, the anterior row some-
what indistinct. Genital operculum of female not completely
fused longitudinally, usually free over distal 1/4. Superior
margin of movable cheliceral finger with five stout teeth,
the inferior margin with pale crenulations or more distinct,

No. 9/21] HJELLE: NORTHERN CALIFORNIA SCORPIONS 23

dark teeth. Pedipalps with inner keel of fixed finger
strongly scalloped, more so in male; fixed and movable fingers
with six and seven supernumerary teeth, respectively.

TYPE DATA. Type specimen from the valley of the Great
Salt Lake, Utah, collected by Howard Stansbury. Depository:
United States National Museum.

DISTRIBUTION. This is a common widespread species occur-
ring in the southern parts of Alberta and British Columbia in
Canada, the northwestern United States, the Rocky Mountain
States, northern Arizona, Nevada, and eastern California.

RECORDS > //iDAHOe TButte County: 10 milesseast) of Anco,
31 July 1966 (E. Slansky), 1 male. OREGON: Harney County:

2 miles northwest of Crane, elevation 4200 feet, 11 June 1967
(R. R. Snelling), 1 female. CALIFORNIA: Modoc County: Lava
Beds National Monument, 28 August 1966 (R. C. Gardner & S. E.
Hasruson)i, 1 female. (Siskiyou County: 7S miles northeast of
Bray, 26 July 1967 (collector unknown), 1 male. Alpine
County: (ePleasant \V.) (2); 8 October: 1962); (WR. Bauer), oi
male. Inyo County: Westgard Pass, 1961 (R. Hock), 2 males;
dry wash of the North Fork of Crooked Creek, opposite Bucks
Peak, Ancient Bristlecone Pine Area, Inyo National Forest,
elevation 9270 feet, 10 September 1968 (C. R. Smith, F.
Ennik), 1 female. Mono County: McGee Creek, 29 June 1938
(Fleshner), 1 male, 1 female; near Hot Springs east of de
Chambeau Ranch, northwest end of Mono Lake, 26 June 1965 (K.
Hom), 2 males, 4 females. Riverside County: Mt. San Jacinto,
23 August 1938 (W. Warren), 1 male.

REMARKS. This species has the northernmost distribution
of all North American scorpions. Of all the specimens from
the Coast Ranges which I have seen, those identified as V.
boreus have actually been specimens of V. silvestrii of either
DESMEYpI Cal), form) or juts lightly colored racial form. Not
having seen all of the specimens from this area recorded as
V. boreus, I cannot make a positive statement, but from the
records available, it appears that V. boreus occurs only to
the east of the Sierra Nevada and does not occur in the Coast
Ranges. I have retained this species in the key because of
this uncertainty.

Vaejovis silvestrii Borelli.
(hatguise’s Sea 47 V 487 Sid 2)

Vaejovis silvestrii Borelli, 1908-1909, pp. 225-227 (original
description). Ewing, 1928, p. 14.

Vejovis boreus Gertsch, 1958, p. 6.

Vejovis silvestrii Gertsch SiSoleqlad 49166), pps 2U5— 20).

DIAGNOSIS. Scorpion of moderate size resembling V. boreus
in general appearance. Base color yellow to tan; carapace
yellow with pattern of black stripes radiating from midline;
preocular area dark; mesosoma with dark coloration continuous
to posterior margin on segments 1 to 6, each tergum with pair
of oval spots near the midline and a series of spots radiat-—
ing outward laterally on each side. Local color races

24 CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

variable in color, from typical coloration to almost complete
absence of dark coloration, some individuals strongly resem-
bling V. boreus (fig. 48). Inferior median keels of metasoma
I to IV obsolete, indicated by dusky lines of varying inten-
sity. Pectine teeth of female 18 to 24, of male 25 to 29;
middle lamellae with two rows of sclerites. Genital opercu-
lum of female not completely fused longitudinally, free dis-
tally up to 1/4 length. Inferior margin of movable chelicer-
al finger with several irregularly formed teeth. Pedipalps
with inner keel of fixed finger more weakly scalloped than in
Vie wbeKneusK
" TYPE DATA. Female type from Sierra Madre, Los Angeles
County, California. }/ Depository: Museo) Zoologico; yturanay
Téaly.

DISTRIBUTION. Baja California and California:

RECORDS. CALIFORNIA: Los Angeles County:> Tanbark Flat,
20 June 1950 (J. W. MacSwain), 1 male. San Luis Obispo
County: }+Morro Bay, 24, September 71969) (fT. 1h. py Richards) py2Z
males, 1 female; 1.3 miles west of Highway 101 on West Cuesta
Road, elevation 2400 feet, 4 April 1970 (J. T. Hjelle, F. H.
Koehler, B. E. Proeres), 8 males, 10 females. Monterey
County: Arroyo Seco Campgrounds, 21 May 1968 (P. Cammer),
1 female; Los Padres National Forest, 4 miles southeast of
Nacimiento Summit Campground, 8 August 1970 (W. E. Azevedo,
V. F. Lee), 9 males, 8 females; Los Padres National Forest,
0.5 mile south of Nacimiento Summit Campground, 8 August
1970 (W. E. Azevedo, V. F. .Lee):,' 8 mahes;,)3 females; Los
Padres National Forest, 2 miles southeast of Nacimiento
Summit Campground, 8 August 1970 (W. E. Azevedo, V. F. Lee),
16 males, 5 females. San Benito County: Pinnacles National
Monument, 25 August 1968 (collector unknown), 1 female.
Stanislaus County: Adobe Creek, 26 February 1956 (D. Bur-
dick), 1 male; 1.3 miles east of the Atomic Energy Commission
Explosive Test Site, 12 April 1965 (K. Hom), 1 female; Car-
negte -Site, Corral, Hollow, 8 May b965a(Vii GF. obec) ; el ematies
1 1/4 miles east of the Atomic Energy Commission Explosive
Test) Site; jConnal -HollowzVahleys 73 sduly (965) (il anBElggs aaa
malles; along County Road J-Z, 1.5 miles east of sURE (3 imrles
east of San Joaquin-Alameda County line), 21 April 1970 (F.
Ennik), 1 male, 4 females. Santa Clara County: Santa Teresa
County /Pask),) 26) Rebsnuany 1966 ((ie Sei Braggs)) il femaiter
Alameda County: Conral Hollow, 30) IMayil960). (is 3S 4 0Basnaers) ee
2 females; in canyon north of Mitchell Ravine, Corral Hollow,
8 May 1965 (V. F. Lee, G. Wong), 1 male, 1 female; Sunol
Regional Park, 7 August 1965 (K. Hom), 1 female; 8 miles
north of Corral Hollow Creek, 3 October 1965 (collector un-
known), 1 male; 11 miles southeast of Livermore on Wagnor
Raneh, County Road wJi=2, 30 uly 19684 (C4 RR. Smilth; ka ean
8 males, 3 females; behind quarantine coops on Kimber Farms
Chicken Ranch, Niles district of Fremont, elevation 250 feet,
1 November 1970.5 (d., TH Hjelle, «Th. Mi vGlimme; jRSths Koehler,
F. Salazar), 3 females; Berkeley, 4 August 1966 (S. Jeffery),
I, male... Contra Costa (County: > Antioch, 15 April 1956 We
Wasbauer), 1 male, 1 female; Castle Rock Park, 5 miles west
of Mt. Diablo, no date (F. Ennik), 6 males, 3 females. Yolo
County: Putah Creek, near Zoological Field Station,

Norg2'] HJELLE: NORTHERN CALIFORNIA SCORPIONS 25

University of California, Davis, 1 November 1966 (R. R.
Montanucci), 1 female. Marin County: Carson Ridge, 2 Janu-
ary 1958 (D. D. Linsdale), 2 males; hillside directly behind
Boot nmack Camp), 1/2, mine up Matt "Tail, Mt. Tamalpaxs wl dieMay.
1968 (B. Butterworth), 1 female; 5 miles south of Fairfax on
Bolinas Road, elevation 600 feet, 16 September 1969 (J. T.
Hjelle, T. M. Glimme), 3 females. Colusa County: from 2 to
5 miles along unnamed road connecting Letts Valley Road and
Twin Valley Road, elevation 4000 feet, 31 August 1969 (J. T.
Hjelle, B. E. Proeres), 3 females; Century Ranch, 1 June 1970
(Lottle & Wickey), 1 female. Napa County: 1/4 mile south-
west of the Lake County line along Butts Canyon Road, eleva-
Elonnw/50" feet, ld) September: 1969..(d..T: Hjelle,;<T:.M. Glimme),
16 males, 43 females. Lake County: approximately 5 miles
north of Rayhouse Road at junction of Davis and Cache creeks,
elevation 900 feet, 15 June 1969 (J. T. Hjelle, M. Bolander),
ZA0pmatlles jy USiremailies, and; 9 ,August 1969) (diene Haedwier ams
Hateras) +17 males’) 121 females; from! 3 1/2: to..7 milessveast
of Bear Creek Ranger Station, elevation 3700 to 4500 feet, 31
August 1969 (J..T: Hjelle,; B. E. Proeres), 7 males, 14 fe-
males; along Clearlake Park Estates Road, Clearlake Park,
elevation 1400 feet, 8 September 1969 and 6 September 1970
(J. T. Hjelle, T. M. Glimme), 11 males, 40 females, and 3
females, respectively; Konocti Road, Mt. Konocti, elevation
LIOOF EO. 28002 feet, 9) September: 19695 (ds) T..Hjelle,) T.aMs
Glimme), 2 males, 4 females; 1/4 mile southeast of Highway
175 on Adam's Springs Road, elevation 2600 feet, 11 September
1969 (J. T. Hjelle, T. M. Glimme), 4 males, 7 females; 1 mile
east of Loch Lomond on Siegler Canyon Road, elevation 3000
feet, 11 September 1969 (J: T. Hjelle, T. M. Glimme) , 1 male,
1 female.

REMARKS. The above records extend the range of V. sil-
vestrii northward about 100 miles, from Marin County to Lake

County. In the area around Panoche Pass, San Benito County,
California, thereroccursia nearly nonmelanverrace ofnV, isaili=
vestrii (fig. 48). These scorpions are essentially the same

as the typical V. silvestrii except for this color variation.
In an extensive area around this locality are found scorpions
with varying degrees of dark coloration, from the typical V.
Ssilvestrii patterning, to patterning much like that of V.
boreus, ultimately to the nearly colorless forms. Most of

these variants show a dark V-shaped marking which connects
the median eyes with each lateral group of eyes. Records of

this race of V. silvestrii are as follows: San Benito County:
Panoche Pass, 3.5 miles west of Fresno County line on road
between Panoche and Mendota, 14 August 1969 (S. C. Williams,
V. F. Lee, M. M. Bentzien), 77 males, 139 females; Pinnacles
National Monument, 4 May 1968 (S. C. Williams & party), 22
males, 32 females; Panoche area, 3.9 miles southeast of
FUNGCELOn OF Road Wwath,,coad to. Ldsiay. on Jal) Ais May el96i8
(Si GC. eWalldlivams iKeuce ‘Schroen}: Jap RixGabel) 676 *malesye) 72
females. Stanislaus County: Frank Raines Park, approximate-
ly 18 miles west of Patterson, 27 September 1969 (S. C.
Williams, J.T. Hjelle,;/W..E. Azevedo, ;V.-F.,Lee) ;,,13-males,;
26 females. Fresno County: Panoche Canyon, 24 March 1967
(R. Snelling), 1 male. Monterey’County: Los Padres National

26 CALIFORNIA ACADEMY OF SCIENCES [Occ.Papers

Forest, Wagon Cave Campground (25 miles south of Arroyo Seco
Campground), 9 August 1970 (W. E. Azevedo, V. F. Lee), 22
males, 4 females.

DISCUSSION AND CONCLUSIONS

In separating the genus Uroctonus from the genus Vaejovis,
I believe that there are two characters of primary importance:
1) the ratio of palm width/palm depth. In Uroctonus, this
ratio is always more than 1.05, reaching a probable maximum
Of 1450 (actually; > in the more) than” 200) andivadual's” omvsewetss
al different species measured, the range of this ratio went
from: is Os tonle45 wath, ay mean) ot l12'8) \ Mose Species Of
Vaejovis show this ratio as 1.00 or less. In particular, in
the Husithenuxra Group, this ratio: ranged \Exrom)0)2/82i toy wavs
with a mean of 0.96; in the subgenus Paruroctonus, this ratio
Fanged trom 0)..67) tonl O0ews tha mean, Of 0SO 7; Mand) an ithe
Punctipalpus Group, this ratio ranged from 0.94 to 1.10 with
a mean of 1.02. In remaining groups, and in species of the
above-mentioned groups, where this ratio extends into the
range of Uroctonus, other important characters, which are
mentioned in the key, serve to distinguish Uroctonus from
Vaejovis. 2) the incomplete fusion of the genital operculum
in the female, the valves remaining free distally up to 1/4
of their length. All species of Uroctonus which I have had
the opportunity to examine have shown this condition, so this
appears to be a stable and important character of this genus.
Among the related vaejovids, only the subgenus Paruroctonus
exhibits this character. However, this subgenus may be dis-
tinguished by its greater numbers of middle lamellae (in some,
two rows of sclerites) and by the low palm width/palm depth
ratio. One must use these taxonomic characters in combina-
tion, since no single character is useful in determining all
species. However, these two characters appear much more
stable and reliable than those others previously considered.

The scorpion fauna of the northern California Coast Range,
as a whole, cannot be considered particularly rich in the
number of species present. Nine species and three subspecies
from two families were found to occur in an area of approxi-
mately 30,000 square miles. Gertsch and Allred (1965) re-
ported a North American record of nine species from two
families collected in the Nevada Test Site (an area of about
1000 square miles). Williams and Hadley reported five spe-
cies from two families from the Puerto Penasco area of
Sonora, Mexico in an area only 5 miles in diameter. However,
if the study area is broken up into two parts, with a San
Francisco-Oakland line dividing them, the picture changes
somewhat. In the northern area (about 24,000 square miles),
four species and 2 subspecies from one family are found,
while in the southern area, six species and three subspecies
from two families are found. To look even closer, in an
area from Alameda County to San Benito County (about 2500
Square miles), six species and three subspecies from two
families may again be found. This compares more favorably

No.92] HJELLE: NORTHERN CALIFORNIA SCORPIONS Di,

with the above-mentioned record reported by Gertsch and
Allred. These figures appear to correspond with the fact
that the geology, floral elements, and climatic conditions
are more diverse in the southern area. The spider fauna of
the coast ranges seems to be more diverse than the scorpion
fauna. Schick (1965) reported 59 species and 4 subspecies
of thomisid spiders from the northern California Coast Range
are (although his Idrian biotic district includes approxi-
mately 6500 more square miles).

In the study area north of San Francisco, the generally
predominant scorpions were Uroctonus mordax mordax and
Vaejovis silvestrii, both found in various situations ranging
from rather dry, inland chaparral areas to moist, foggy,
forested areas. Vaejovis silvestrii is generally more common
in drier areas. Vaejovis gertschi striatus was fairly common
along the inner coastal ranges, tolerating areas of little or
no brush cover. Uroctonus glimmei appears to follow the same
dustnibutivonalljpaths (as Vi. g-isitriatus, but. is,less:,common:.
In the study area south of San Francisco, Uroctonus mordax
plunidens predominates inthe western,coastal ranges of “Santa
Cruz and Santa Clara counties. Vaejovis silvestrii is most
closely associated with the chaparral and woodland-grass
aveas ‘south of) Santa Cruz County, and anland up through the
East Bay counties. Superstitionia donensis, Anuroctonus
phaiodactylus, and Hadrurus obscurus are restricted to San
Benito County in the warmer drier Lower Sonoran areas around
Panoche and Pinnacles National Monument. Anuroctonus phaio-
dactylus is common in localized areas, while S. donensis and
Hewobscunus are, much less common in) the Coast “Ranges, being
more closely associated with the southern Central Valley
desert regions.

The comparative distribution of various species of scor-
pions with different species of spiders suggests similar
zoogeographical distribution patterns. For example, the
distribution Of \U.. m.smomdaxs along wath U.) me. plunmadenisy aks
very similar to that of Xyst: Xysticus gosiutus Gertsch (Araneida:
ihomusudae) (Schuck, 1965). Also; /the (distributions, of OVE
silvestrii and the V. gertschi group are similar to that Fone
Misumenops aikoae Schick (Araneida: Thomisidae) (Schick,
LOGS)

In addition to the descriptions of a new species, Uroc-
tonus glimmei, and two new subspecies, Uroctonus mordax
pluridens and Vaejovis gertschi striatus, samples yielded
significant northern range extensions for three other species,
Superstitionia donensis, Anuroctonus phaiodactylus, and
Vaejovis silvestrii.

Different groups exhibited varying degrees of genetic
isolation. Uroctonus m. mordax demonstrated, through various
characters, tendencies toward the formation of distinctly
different subpopulations. Also, V. silvestrii showed these
same tendencies in the formation of localized nonmelanic
races. These unique tendencies toward subpopulation forma-
tion may arise from such factors as habitat differences,
genetic drift in small populations, ecological and geographic
isolation.

Several species of scorpion have been reported from

28 CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

California which I consider to be spurious records. Banks
(1900, 1904, 1910) reported the following: Hadrurus hirsutus
(Wood), Vaejovis punctipalpus (Wood), and Brotheas alleni
(Wood) are all known only from southern Baja California;
Uroplectes mexicanus Banks is mentioned by Gertsch (1966) as
probably being of African origin; Opisthacanthus lepturus
(Palisot de Beauvois), Diplocentrus keyserlingi Karsch, D.
whitei (Gervais), Tityus tenuimanus Banks, and Isometrus
maculatus (DeGeer) are all tropical species and probably do
not extend into California unless as accidentals by transport
with food or other materials; finally, that Centruroides
californicus (Wood) occurs in Lake County, as reported by
Banks (1910), is doubtful, since Lake County has been inten-
Sively sampled, and no evidence of the occurrence of this
species (or genus) could be confirmed.

In his monograph, "Ueber Scorpione," Karsch (1879) des-
cribed a new species o£ Uroctonus from "California" which he
named"UToctonuspruvus: 2 Knaepelliny (E894) Waren teal ted
iPDELVUS al junItorssynonym or U. mordax. “Since! that, Erne, aem
other scorpion workers have accepted Kraepelin's placement.
However, Characters deemed unimportant by Kraepelin now
appear to have special significance. The type was, therefore,
borrowed from the Zoology Museum of Humboldt University,
Berlin, Germany. Close examination of the specimen (figs.
19-24) indicated that Kraepelin was mistaken in regarding
it as a synonym of U. mordax.

The original description was brief, not including some
important characters, and including no measurements. The
holotype examined agreed with the original description except
in the following ways: metasoma V shorter than metasoma III
and IV together; three denticles on the inferior margin of
the movable cheliceral finger. Karsch did not define the
sex which is clearly a juvenile female.

The characters which exclude this specimen from the genus
Uroctonus are: 1) the circular shape of the booklung open-
ings (stigma) (fig. 21); and 2) the presence of pectinal
teeth) on) only -the distal 3/4 of the (pectines Chag. 22)

These characters are not found on any known species of
Uroctonus’.: “Except for the presence of three. lateralveyes sper
group, this specimen appears closely related to Brotheas
alleni (Wood) (Scorpionida: Chactidae). In addition to the
Characters mentioned above, the structure of the pedipalps,
the relative lengths of the metasomal segments, and the
general coloration’ are very ‘stmelar to that Of 7B). a liveiman
Because the data needed for exact placement of this individ-
ual are unavailable at this time, I will only tentatively
suggest that this species may have its closest relationship
with the species now known as Brotheas alleni (Wood), from
Baja California Sur, Mexico. I consider the California
record to be spurious, this individual most probably being
from Baja California, Mexico.

Type data. Holotype female, "California" (there appears
to have been another word above "California" which could not
be made out). Depository: Zoologisches Museum der Humboldt-
Universitat, East Berlin, Germany (Graber collection). Type

No.92] HJELLE: NORTHERN CALIFORNIA SCORPIONS 29

specimen poorly preserved, segment 7 of mesosoma transparent,
third right walking leg missing, metasoma detached, one
chelicera missing.

The scoupvon fauna Of other, areas of California is still
relatively unknown. Data from the Modoc Plateau area, much
of the Sierra Nevada, and the southern coast ranges are par-
ticularly lacking. A complete picture of the scorpion fauna
of California is not only of scientific interest. From the
numbers of scorpions to be seen during much of the year, they
are obviously important predators of other arthropods, both
insect and arachnid. Also, there is a public health interest
in that almost nothing is known about the toxicity of the
venom of most of the species now known from California.

These and other topics, such as population and behavioral
studies, are only a few examples of needed future research.

REFERENCES

BANKS, N.
1900. Synopses of North American invertebrates. IX.
The scorpions, solpugids and pedipalpi. The
American’ Naturalist ,.ivol.° 34, ppi+421-427.
1904. Some arachnida from California. Proceedings of
the California Academy of Sciences, 3rd series,
VOM Sea lOve Deoipn2 Ons Ora ke

LOO The Scoupiens: Of California. 9 Pomona’Col lege
Journal of Entomology, vol. 27, ino.<2;. pp.=L385—
OS

BORELLI, A.

LIOS=e41 SCOrpienw waccoltie dali Prof: F.sSiilsvestri nell

USO America settentrionale e alle isole Hawaii.
Bollettino del Laboratorio di zoologica generale
€ agraria della R. Scuola superiore d'agricol-
UGA EP OG EMA, VO 1 913: ppak 222221.

CHAMBERLIN, R.V.

1923. .The northern range of the sicorpion. Science,*vol-

59 sh ps4 G4.
COMSTOCK Ube Ei.

"1912" (1913). The Spider Book. Doubleday, Doran and Co.,
inewsNewSYork.ui 72 pps

1940. The Spider Book, revised and edited by W. J.
Gertsch. Doubleday, Doran and Co., Inc. New
MONE eA) NAS oy ove

DIAZ-NAJERA, A.

1970. Contribucion al conocimiento de los alacranes de
Mexico (Scorpionida). Revista de Investigation
en: Salhidn Publica, <volvy 307 enov.2;8pp. 7 ll1l=122-

DURRENBERGER, R. W.

1968. Patterns on the land. National Press Books. Palo

Alto, ,calltornLrayivL09s pp.
EWING, H.

1928. The scorpions of the western part of the United
States with notes on those occurring in northern
Mexico. Proceedings of the United States Nation-
ateMuUSeUm VO 7S) tlOls 9, pple liar

30 CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

GERTISCH, W. J-
1958. Results of the Puritan-American Museum Expedition
to western Mexico. 4. The scorpions. American
Museum Novitates, no. 1903, pp. 1-20.
GERTSCH, W. J., and D. ALLRED
1965. Scorpions of the Nevada Test Site. Brigham Young
University Science Bulletin, Biological Series,
WO Opa Orn 149, Oe neal SY
GERTSCH, W. J., and M. SOLEGLAD
LI66en Bbhe Sscorprens Of Ene Vejovis boreus group (sub-
genus Paruroctonus) in North America (Scorpion-
ida: Vejovidae). American Museum Novitates,
NOW 228) pela oe.
GIRARD, C.
eS 4 Lr Marcy' s exploration of the Red River of
jmLouistana in 1852... -33rd) Congress, Ust\sessvon,
Senate Executive Document, pp. 251-259.
HENRY, L. M.
1949. The nervous system and the segmentation of the
head in a scorpion (Arachnida). Microentomology,
VOW 4s non Ai ppe. b2 025.
HOFFMAN, C. C.
1931. Monografias para la entomologia medica de Mexico.
Monografia num. 2, los scorpiones de Mexico.
Primera parte: Diplocentridae, Chactidae,
Vejovidae. Anales del Instituto de Biologica,
Universidad de Mexico, vol. 2, no. 4, pp. 291-
408.
JENKINS, O. P. (director)
1951. Geologic Handbook of the San Francisco Bay Coun-

ties. California Department of Natural Re=
sources, Division of Mines, Bulletin 154.
S92 Opis
KARSCH, F.
1879. Scorpionologische Beitrdge. Ueber scorpione.

Mittheilungen des Munchener Entomologischen
Vemeansi evo 1S),1 70pm mo iso Ole
KO CH Creare
W886.) Die Anachniden. Nurnberg.) )Vol..3) op 4y 5 0—ome
KRAEPELIN, K.

1894. Revision der skorpione. II. Scorpionidae und
Bothriuridae. Jahrbuch der Hamburgischen
Wissenschaftlichen Anstalten, vol. 11, pp. 1-248.

1899. Scorpiones und Pedipalpi. Das Tierreich, vol. 8,
De 82

RU RAGAy i livin iSie
1930. Notes on the northern scorpion, Vaejovis boreus
Girard, in British Columbia. Canadian Field-
Natunpailliasiey) Vole > 4a mpp ee. ons Ole
MANN, B. P. (editor)
1876. Synoptical tables for determining North American
IMSecks. — oCOnpDITONes a sPSVGhe, WVOlm a) nOmmSie,
pp. 209-214.
MELLO-LEITAO, C.
1945. Escorpioes Sul-Americanos. Arquivos do Museu
Nacional, vol. 4077p.) 228).

No.9 2] HJELLE: NORTHERN CALIFORNIA SCORPIONS Sill

MOLES, M.
OD a Amst Calla cormnsasay Ad achnada:s (Gla Coxe? iP aera s
The Scorpionida). Journal of Entomology and
HOO MKS) Gi NOLL Ga Sis! KOAY Dip joo talAoilss
MUGATK,.) S:.), sand) HH.) 1G.) HiGGINS
1944. A new genus of scorpions in the southwest.
Entomological News, vol. 55, no. 9, pp. 237-240.
MUNA eb.) Aang) DL) Di. KECK
UDOSiw WA Cala forntay Eloxra. 7 University of California
Press, 1681 pp.
RO COCK Ri. Liv.
e933. On the classification of scorpions. . Annals and
Magazine of Natural History, 6th series, vol.
LAA INO ALOK ol 4 SOLS
1894. Scorpions and their geographical distribution.
Natumala Scrence), volj.4, no. 24, pp. 253-364).
1902. Arachnida, Scorpiones, Pedipalpi and Solfugae.
Biologia Centrali-Americana. 71 pp., 12 plates.
SCRIBE AN) 1584) Ge
1965. The crab spiders of California (Araneida: Thomis-
idae). Bulletin of the American Museum of
Natural aston. VOl. L297 eno. a ppc thal oOr
STAHNKE, H.L.
1940. The scorpions of Arizona. Iowa State College
JOUENalTOt -Sevence,, “Vol 115i pp. LOL 033
1949) Diplops, desertorum a scorpion, Synonym. Entomo-
logically NewSey Vol. 160 > NO. 1)9);) Dis 24 Si.
1966. Some aspects of scorpion behavior. Bulletin of
the Southern California Academy of Sciences,
VOUS OD 7p MNOS 5 Jools
AMEKO URI Hi BiG PS aE
Loo. On. the classitacation, Of -SCcorp1ons. Annals) and
Magazine of Natural History, 4th series, vol.
LUTE Sh SOKO GS) Tips oj ole ila ys
1876- Etudes scorpiologiques. Atti della Societa
ILS italiana di scienze naturali, e del Museo civile
Giistormital matuisaer a MaiMlan@y sVvi@ueumlOs | ini@ malas
IIo) VO S30) Bi woul Gr ILE) A Malo AA eG LAYS
1893. Scorpiones exotici. Bollettino della Societa
PMeOnologiCa, kealllana,. VOl.) 2a; Ds (SOOT.
VACHON, M.
1963. De lL'utilite, en systematique, d'une nomenclature
des dents des chiliceres chez les scorpions.
Bulletin du Museum National d'Histoire Naturelle,
Pewelisne Geuegs Pi xrollsa siaipa eles Ap: joyoms LLGAL SILO
WEBSTER, R. Li.
23s ue SeCOrpLoOnG in, NOKthy Dakota.) | SCLeCNCe, aVOl 5S), aiDls
248.
WERNER, F.
1935 ju uSCOLpTons und -Pedipalipi. 2 In Bronny,) i. iG. Klassen
und Ordnungen des Tierreichs, Leipzig, bd. 5,
along. Ch, onblelal ts) pr dbalenee, Sipe yejoaq MSS,
WEELGAMS | Si. GC.
L966 5; BULKowIng acta vitles, (Of the scorpion Anuroctonus
phaeodactylus (Wood) (Scorpionida: Vejovidae).
Proceedings of the California Academy of Sci-
eniceswdthmserves Voll. 34, nO Sip pe 49 —4218%

32 CALIFORNIA ACADEMY OF SCIENCES [Occ.Papers

WihiitAMS See. (continued)

1968a. Scorpion preservation for taxonomic and morpho-
logical studies. Wasmann Journal of Biology,
Wola 2Opinoe dlp Vos kScodbsie

1968b. Scorpions from northern Mexico: Five new species
Of Vejovis from Coahuila, ‘Mexico: | \Occastomal
Papers of the California Academy of Sciences,
Oa Sip voy), bo Aaho

1968c. Methods of sampling scorpion populations. Pro-
ceedings of the California Academy of Sciences,
Ath SeELels, Vol. 3'6))) NO. @Si, MPs N22 Las Or

1968d. Two new scorpions from western North America.
Pan-Paciftic Entomologist, vol. 44, no. 4, spp.

Sl 3—= 32s

1970a. Three new species of Vejovis from Death Valley,
California, ) Pan-Pacific Entomologist; VO uno
IMO Gilby Owe / Lea leas

1970b. Scorpion’ ‘fauna, of Baja Calatorniia, Mexico: Eleven
new species of Vejovis (Scorpionida: Vejovidae).
Proceedings of the California Academy of Sci-
ences), 4th ‘series, (vols. Si, NO, 8,1 Ppa 21 o>eeee

1970c. New scorpions belonging to the Eusthenura group
of Vejovis from Baja California, Mexico
(Scorpionida: Vejovidae). Proceedings of the
California Academy of Sciences, 4th series, vol.
Sp WOn Wey jojo) BSIoaes}

1970d. A new species of scorpion belonging to the
pumilis group of genus Vejovis (Scorpionida:
Vejovidae). Pan-Pacific Entomologist, vol. 46,
MO Sy PO iol O se

1970e. A systematic revision of the giant hairy-scorpion
genus Hadrurus (Scorpionida: Vejovidae).
Occasional Papers of the California Academy of
SCm#encesy MO. Sila, MODs oye

1971. |) Clarifications) in ‘the! nomenclature) of some Neneh
American scorpionida. Pan-Pacific Entomologist,
VOL oi ThA sao Mk DIOS TASS YL)

Wii TbAMSY (Siu Cs). amc Nisgby en EADIE.

1967. Scorpions of the Puerto Penasco Area (Cholla Bay),
Sonora, Mexico, with description of Vejovis
baergi, new species. Proceedings of the Cali-
fornia Academy of Sciences, 4th series, vol. 35,
IMOree DH Dove, ALO Se IEG)

WOOD), WHC om:

1863a. Descriptions of new species of North American
pedipalpi. Proceedings of the Academy of Natur-
al Serence's' of /Phitadedphia 1863), pp. TOv—dak2e

VW863b." On’ the pédipalpi of North America: Journal) or, ehe
Academy of Natural Sciences of Philadelphia, 2nd
SESS), AVOle Up isa SS Io SOE

No.92] HJELLE: NORTHERN CALIFORNIA SCORPIONS 35,

TABLE 1. Measurements (in millimeters) of Uroctonus
glimmei Hjelle, new species, holotype and allotype.

Holotype Allotype
(male) (female)
Total length 34.8 40.2
Carapace, length 4.2 So7
width (at median eyes) 2450) 30
Mesosoma, length Oy0 11.4
Metasoma, length SKS pens
segment I (length/width) ZW 2ix3 DA 2a,
segment II (length/width) PRS) ye Zieh jie 0
segment III (length/width) Dr Of 2rer2 SIO /Q SS
segment IV (length/width) BA (2) 0 Sh], Lie 4
segment V (length/width) D2 210) GOV 2e0
Telson, length 4.8 Sis
Vesicle (length/width) SS Hie alee 4207/2..3
depth lee ILS)
Aculeus, length Eas) 13
Pedipalp
Humerus (length/width) Bi Ol/ dey 4.4/1.6
Brachium (length/width) 4.0/1.4 Aisle
Chela (length/width) 6.8/2.6 844/380
depth DS 2 «6
movable finger, length 3.3.6 4.2
fixed finger, length DG) 3,14
Pectines
teeth (left/right) Dale LOL
TABLE 2. Measurements (in millimeters) of Uroctonus
mordax mordax Thorell, lectotype.
Lectotype
(female)
Total length 5S
Carapace, length 6.4
width (at median eyes) Dye.0
Mesosoma, length 20;35
Metasoma, length 9K 4
segment I (length/width) 26/7 S52
segment II (length/width) BAA PAGS
segment III (length/width) 314/256
segment IV (length/width) BIB 214
segment V (length/width) 6.4/2)..2
Telson, length 6.8
Vesicle (length/width) 4.5/2.4
depth Diane
Aculeus, length BSS
Pedipalp
Humerus (length/width) 5c / Qi
Brachium (length/width) a6 8
Chela (length/width) MO 8/452
depth i 338 (0)
movable finger, length yous)
fixed finger, length 4.3

Pectines
teeth (left/right) 8/9

34 CALIFORNIA ACADEMY OF SCIENCES [Occ.Papers

TABLE 3. Measurements (in millimeters) of Uroctonus
mordax pluridens Hjelle, new subspecies, holotype and
allotype.

Holotype Allotype
(male) (female)
Total length D505) 572.0
Canapacey, Wengeh 0 TEAS
width (at median eyes) 549) B65
Mesosoma, length A Gyens Qik
Metasoma, length Def) 21.6
segment I (length/width) BG OY) Sone 3.0/3.4
segment II (length/width) BOSS 3473.0
segment III (length/width) ASW/ 35) BA ei Ais)
segment IV (length/width) 4.8/2.8 4.2/2.6
segment V (length/width) TE 3/256 Ti 27264
Telson, length 8.0 Si)
Vesicle (length/width) Bye ey Sio, 6) UN tBy// 24-6
depth Dee rae
Aculeus, length Os 2.2
Pedipalp
Humerus (length/width) 6 0/2055 610/26
Brachium (length/width) 569/350 630/738
Chela (length/width) W265 A/a 8 12 1/2
depth 309) S16
movable finger, length 6.4 6.8
fixed finger, length yo dl Diers
Pectines

teeth (left/right) ih3//10S 7a

No.92] HJELLE: NORTHERN CALIFORNIA SCORPIONS 35

FIGURE 1. Dorsal view of left chela of Uroctonus mordax
mordax Thorell, showing method of measurement of palm width.

FIGURE 2. Inferior median view of left chela of Uroctonus
mordax mordax Thorell, showing method of measurement of palm
déperEhr

FIGURE 3. Generalized view of left pectine of Uroctonus
mondax mordaxsThorelil.; \ Structures’ shown: | al”—) anterior
TamelITa; ml - middle lamellae; f - fulcra; pt - pectinal
Eeeithy.

FIGURE 4. Dorsal view of right’ movable pedipalp finger
of Uroctonus mordax mordax Thorell.

36 CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

am i

FIGURE 5. Dorso-lateral view of left half of carapace
Of Anuroctonus phaiodactylus (Wood). Structures shown:
am - anterior margin; le - lateral eyes (4); me - median eye.

FIGURE 6. Lateral view of telson of male Anuroctonus
phaiodactylus (Wood). Arrow indicates swollen bulb of
aculeus.

FIGURE 7. Left chelicera of type specimen of Uroctonus
mordax Thorell. Dorsal view, showing: mf - movable finger;
sm - superior margin of movable finger; ff - fixed finger.

FIGURE 8. Left chelicera of type specimen of Uroctonus
mordax Thorell. Ventral view, showing: mf - movable finger;
im - inferior margin of movable finger, with denticles;
fi.) faxed finger.

No.92] HJELLE: NORTHERN CALIFORNIA SCORPIONS a7

FIGURES 9 to 12. Ventral views of hind tarsi showing
characteristic spinule pattern. FIGURE 9. Uroctonus mordax
mordax Thorell. FIGURE 10. Uroctonus glimmei Hjelle, new
Species. FIGURE 1l. Vaejovis silvestrii Borelli. FIGURE

12. Superstitionia donensis Stahnke.

.

38 CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

13

FIGURES 13 and 14. Superstitionia donensis Stahnke, male.
FIGURE 13. Dorsal view. FIGURE 14. Ventral view.

No.92] HJELLE: NORTHERN CALIFORNIA SCORPIONS

FIGURE 15. Dorsal view of Anuroctonus phaiodactylus

(Woody) madie-
FIGURE 16. Typical burrow entrance of Anuroctonus

phaiodactylus (Wood).

39

40 CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

FIGURES 17 and 18. Hadrurus obscurus Williams, holotype
male. FIGURE 17. Dorsal view. FIGURE 18. Ventral view.

No.92] HJELLE: NORTHERN CALIFORNIA SCORPIONS 4l

FIGURES 19 and 20.

Uroctonus
female. FIGURE 19. Dorsal view.

FIGURE 20.

*

privus Karsch, holotype

Ventral view.

42

CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

FIGURES 21 to 24. Uroctonus privus Karsch, holotype
female. FIGURE 21. Representation of ventral surface of

mesosoma showing circular booklung Openangs. VE LCURE MOO.
Pectine’ structure. FIGURE 23. ,Exterio: lateral view of
Chela. FIGURE 24. Lateral view of metasoma III, IV, and

V, and telson.

No.92] HJELLE: NORTHERN CALIFORNIA SCORPIONS 43

25

FIGURES 25 and 26. Uroctonus glimmei Hjelle, new species,
holotype male. FIGURE 25. Dorsal view. FIGURE 26. Ventral
view.

44 CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

FIGURES 27 and 28. Uroctonus glimmei Hjelle, new species,
allotype female. FIGURE 27. Dorsal view. FIGURE 28.
Ventral view.

No.92] HJELLE: NORTHERN CALIFORNIA SCORPIONS 45

FIGURE 29. Exterior lateral view of left chela of
Uroctonus glimmei Hjelle, new species, holotype male.
FIGURE 30. Exterior lateral view of left chela of

Uroctonus mordax Thorell, lectotype female.

*

46 CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

FIGURE 31. Ventral view of chela of Uroctonus glimmei
Hjelle, new species, holotype male, showing characteristic
carination.

FIGURE 32. Ventral view of chela of Uroctonus mordax

Thorell, lectotype female, showing characteristic carination.

No.92] HJELLE: NORTHERN CALIFORNIA SCORPIONS

FIGURES 33 and 34.

Lectotype female of Uroctonus
mordax Thorell. FIGURE 33. Dorsal view. FIGURE 34.
Ventral view.

47

48 CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

9 o”

10

5

Numbers of individuals

5

10

3 tid
16 17 18 19 20 21 22 23

Number of pectinal teeth

24 25 26 27 #+28 29

FIGURE 35. Differences in pectine tooth count modes
between four different populations of Uroctonus mordax
mordax Thorell. a) Glenn County, California. b) Bootjack
Camp, Marin County, California. c) Sonoma County, Calli fornia
d)@ Kent hake, Marin County, Calla Fomniitar

No.92] HJELLE: NORTHERN CALIFORNIA SCORPIONS 49

4i

5

10

5

10

4.3

5
2.0

Numbers of individuals

15)

10

°
~
wv

3 4 5 6 7

Number of denticles

FIGURE 36. Differences in modes of counts of denticles
on inferior margin of movable cheliceral finger between four
different populations of Uroctonus mordax mordax Thorell.

a) Fairfax Area of Marin County, California. b) Bear Creek
Area of Lake County, California. c) 13 miles east of Covelo,
Mendocino County, California. d) Rattlesnake Island, Clear
Lake, Lake County, California.

50

CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

FIGURES 37 and 38. Uroctonus mordax pluridens Hjelle,
new subspecies, holotype male. FIGURE 37. Dorsal view.

FIGURE 38.

Ventral view.

No.92] HJELLE: NORTHERN CALIFORNIA SCORPIONS

FIGURES 39 and 40. Uroctonus mordax pluridens Hjelle,
new subspecies, allotype female. FIGURE 39. Dorsal view.
FIGURE 40.. Ventral view.

5) Ak

By

4]

CALIFORNIA ACADEMY OF SCIENCES [Occ sPapers

FIGURES 41 and 42. Vaejovis gents chan simiaus Hjelle,
new subspecies, holotype male. FIGURE 41. Dorsal view.

FIGURE 42.

Ventral view.

No.92] HJELLE: NORTHERN CALIFORNIA SCORPIONS

43

FIGURES 43 and 44. Vaejovis gertschi striatus Hjelle,
new subspecies, allotype female. FIGURE 43. Dorsal view.
FIGURE 44. .Ventral view.

53

54 CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

FIGURE 45. Exterior lateral view of chela of Vaejovis
gertschi striatus Hjelle, new subspecies.
FIGURE 46. Dorsal view of Vaejovis boreus (Girard).

No.92] HJELLE: NORTHERN CALIFORNIA SCORPIONS 55

FIGURES 47 and 48. Vaejovis silvestrii Borelli. FIGURE
Aj... Typical collor form. |) FIGURE Deus bughitly.wokored racial

form. :

56 CALIFORNIA ACADEMY OF SCIENCES [Occ.Papers

FIGURE 49. Distribution of Uroctonus glimmei Hjelle, new
species (©), Vaejovis gertschi striatus Hjelle, new subspecies
(@), and Anuroctonus phaiodactylus (Wood) ([]).

No.92] HJELLE: NORTHERN CALIFORNIA SCORPIONS

) 2 f Be = Aa
OO earl :
‘ ’
th A YRS !
@ \ a es
O OR ar pe
Or ae
©) aegis Cs Abr eae
t
» Ne }
- OS i rf
v aN /
Be
. ‘ ;
L if ‘ Eee
et Gi ie
N A ae ( i oe
” 7
AX ©) rte tee AOS ,
) he * vi \
y t ’
OVO ! oe. ; <>
(Naot S ey
i car vai Gack
s OO We SUG ee Al
eo teal an
\8O , - : i i
. 5 Be 7 eee ae
‘6% ed We a Ue ae
CaO 4 ne
et oA Be ne
\ 7 a4
ses fuwaee on |
~ \
L ENE i ReRicaaty ae
\ yy ia ‘
> ees
oe Y 1 S
( 4 ‘
Renee
a ey
Ete a ee res l
Udinese Ald SHAVE
Q ae
“oy
au !
0 5 ‘ oa : |
\P S
Miles ; oF |
SiGe ee ee
renee :
: \
\
\ |
Tes : |
BI Le Te tihs
i Ne
x
?

FIGURE 50. Distribution of Uroctonus mordax mordax
Thorell (Q) and Uroctonus mordax pluridens Hjelle, new

subspecies (@).

58 CALIFORNIA ACADEMY OF SCIENCES [Occ.Papers

Miles T-

Na i= es

FIGURE 51. Distribution of Vaejovis boreus (Girard) (@),
and Vaejovis Silvestri Borelli,,, eypiical icollon tor, \¢@))y,
lightly colored” racial form (@), and intergrades (@).

No.92] HJELLE: NORTHERN CALIFORNIA SCORPIONS 3)8)

C
/

FIGURE 52. Distribution of the following scorpions in
the East and South Bay counties: Uroctonus mordax mordax
-Thorell (©), Uroctonus mordax pluridens Hjelle, new
subspecies (A), Uroctonus mordax (mordax X pluridens
intergrades) (@), Uroctonus glimmei Hjelle, new species
(@), Anuroctonus phaiodactylus (Wood) (O), Superstitionia
donensis Stahnke (4), and Hadrurus obscurus Williams (@).