A new cavernicolous Parobisium Chamberlin 1930 (Pseudoscorpiones: Neobisiidae) from Yosemite National Park, U.S.A.

TEXAS TECH UNIVERSITY

Natural Science Research Laboratory

Occasional Papers

Museum of Texas Tech University

Number 297 30 September 2010

A New Cavernicolous Parobisium Chamberlin 1930 (Pseudoscorpiones:
Neobisiidae) from Yosemite National Park, U.S.A.

James C. Cokendolpher and Jean K. Krejca

Abstract

A new species of troglobitic Parobisium pseudoscorpion is described from two caves de¬
veloped in granite talus slopes in the Yosemite Valley, U.S.A. The 16 species of the genus are
all from the northern hemisphere (western U.S.A., China, Japan, South Korea). A taxonomic
key to the genus in the U.S.A. is provided. The new species has only an anterior pair of pale
colored eyespots without tapetum and is blind. Extensive searching at other shallow nearby caves
and on the surface has not revealed any other specimens of this species, although it is common
within certain areas of the two known caves. This may be only the second troglobite described
from granite talus caves in North America, and suggests the potential for fruitful exploration in
regions not traditionally sampled for subterranean fauna.

Key words: cavernicolous, boulders, granitic cave, Parobisium , pseudoscorpions, talus,
troglobite, Yosemite National Park

Introduction

Yosemite Valley is glacially sculpted and bor¬
dered by iconic granite bedrock formations including
Half Dome and El Capitan. According to Huber (1987),
about a million years before present, a Sherwin-aged
glacier excavated and shaped the valley. At least
two later glaciations occurred (Tahoe and Tioga gla¬
ciations, which probably peaked about 130,000 and
15,000-20,000 years ago, respectively) but were of
much lesser areal size and height (Huber 1987). Thus,
the later events did little to further modify or smooth
the walls of Yosemite Valley. Above the ice surfaces of
those later glaciers, the valley walls had almost a mil¬
lion years to weather - joints widened, rock fractured
and crumbled, with water flows and falls eroding back

into ravines and less durable rocks. The Merced River
has further incised the one km deep valley, and along
either side of the valley walls are talus piles ranging
from smooth, gently sloping grades to steep boulder
fields. Over 500 documented rock falls in historic
times (Wieczorek and Snyder 2004) demonstrate the
ongoing nature of the talus slope formation. Within
these talus slopes are caves of varying sizes that simply
consist of spaces within the jumble of boulders, with the
largest boulders providing stable roofs for a few more
extensive caves. The two caves of primary interest are
likely only a few hundred years old. Talus caves are
created, eroded, and then most likely the fauna moves
into the more recent talus fall regions. Only the more

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Occasional Papers, Museum of Texas Tech University

recent areas are accessible to humans, but countless
voids large enough for invertebrates must exist in the
old and buried rubble piles.

Krejca (2007) performed a biological inventory
of two of the longer caves in Yosemite Valley, Spider
Cave and Indian Cave, Mariposa County, California, in
the summer of2006. During that study four specimens

of a new species of pseudoscorpion were collected
from Indian Cave for taxonomic identification. Two
subsequent visits (3-4 March and 7-8 July 2009) were
organized to collect additional material and attempt
to determine if the species is actually cave-limited by
sampling a variety of similar surface habitats. On these
later trips a specimen also was obtained in Elf Village
Cave, located less than 0.5 km from Indian Cave.

Materials and Methods

Field observations. —Surveyors used standard
caving techniques and performed searches by crawling
slowly along the entire zone in each cave, recording
all fauna that occurred on each surface including the
floor, ceiling, and walls. Search effort was measured
in terms of surveyor-minute, and reflects the time it
took to search the entire zone of the cave for pseu¬
doscorpions. Temperature and relative humidity (RH)
were measured using a fan-cooled wet and dry bulb
psychrometer (Psychro-Dyne®, Industrial Instruments
and Supplies). Atmospheric pressure was measured
using a hand held Tech 0 digital barometer. Measure¬
ments were taken inside the caves at each zone where
searches were performed. Relative humidity (RH)
values were calculated from wet bulb, dry bulb, and
barometric pressure measurements.

Specimen repositories. —All but one male and
two female specimens are at the Museum of Texas Tech
University (TTU-Z). Of the three other specimens, one
male paratype (formerly TTU-Z 51564) and one female
paratype (formerly TTU-Z 51536, Zara 3563) are re-
posited in the collection of the California Academy of
Sciences (CAS), and one female paratype (formerly
TTU-Z 51532) is reposited at the Florida State Collec¬
tion of Arthropods (FSCA). All specimens are detailed
under “type material and treatments.”

Specimen preparation. —The specimens were
collected directly into 75% or 95% ethanol. The ap¬
pendages from one side of the body of some of those
in 95% ethanol were removed and placed in a -80°C
freezer. The remainder was transferred to 75% ethanol.
Most specimens were then transferred to lactophenol
on a shallow depression slide and cleared. After study

with light microscopy, the specimens were returned to
75% ethanol. For SEM study the appendages were re¬
moved and dehydrated through a graded ethanol series,
transferred to hexamethyldisilazane (HMDS) through
an ethanol/HMDS series to pure HMDS, dried in a vent
hood overnight at room temperature, and mounted on
a stub without metal coating.

Image and examination. —Fight microscopy
(FM) included use ofNikon SF-3D and Feica MS5 mi¬
croscopes with photo attachments and an 8 megapixel
Nikon Coolpix 8700. Photoshop was used to enhance
these and other photographs, some with the use of an
i-pen Mouse (FingerSystemU.S.A. Inc.). Z-axis mon¬
tages of FM images were created with Helicon Focus
(Helicon Soft Ftd.). Scanning electron microscopy
was achieved by the use of a Hitachi S-4300SE/N.
Digital images presented herein as well as others to
photographically document the specimens are retained
in the Museum of Texas Tech University digital catalog
under each specimen’s individual number (TTU-Z).
Unless indicated otherwise, all illustrations are by the
first author.

Measurements were made following the direc¬
tions in Chamberlin (1931, p. 32-25), using an ocular
micrometer with FM, or read directly off SEM scales.
Trichobothria names follow those used by Chamberlin
(1931). Harvey (1992) noted that the leg basifemur
and telofemur were not both femoral segments, and
are correctly termed the femur and patella. The cheli-
ceral “flagellum” has been renamed the rallum (Judson
2007). The terminology for the female genitalia follows
Judson (1993).

COKENDOLPHER AND KREJCA-CAVERNICOLOUS PSEUDOSCORPION

3

Taxonomy

Muchmore (1990) provided an annotated key to
the various taxonomic levels down to and including
genera of pseudoscorpions in the U.S.A., and this key is
still useful, although slightly outdated. Harvey (1992)
remains the modern standard for higher classification
of pseudoscorpiones.

Family Neobisiidae Chamberlin 1930

Diagnosis. —Apex of palpal coxa (manducatory
process) rounded and with three or more setae. Venom
apparatus absent from moveable chelal finger (Harvey
1992).

Subfamily Neobisiinae Chamberlin 1930

Diagnosis. —Galea reduced to a sclerotic knob
(Harvey 1992).

Genus Parobisium Chamberlin 1930

Neobisium (Parobisium ) Chamberlin 1930:17;
Beier 1932:84; Morikawa 1960:112-113; Hoff
1961:427.

Parobisium Chamberlin: Chamberlin and Mal¬
colm 1960:112-113; Chamberlin 1962:123; Harvey
1991:394; Mahnert 2003:744-745.

Diagnosis. —Fixed chelal finger with a compact
subterminal cluster of only three tactile setae ( et, it,
est ) and a more diffuse subbasal to basal cluster of
five tactile setae ( isb, ist, ib, esb, eb)', both clusters of
setae number four (et, est, it, ist/esb, eb, isb, ib) in other
genera, and seta ist is subterminal instead of subbasal
in position (Chamberlin 1962).

Subordinate taxa and distribution.—Parobisium
now consists of 16 described species and four additional
subspecies (Harvey 2009): East Asia: China (three spe¬
cies), Japan (four species), South Korea (two species);
North America: western U.S.A. (seven species).

North American (U.S.A.) taxa:

Parobisium charlotteae Chamberlin 1962. Type
locality: Redmond Lava Cave, Deschutes County,
Oregon.

Parobisium hastatum Schuster 1966. Type local¬
ity: 6 miles S El Dorado, El Dorado County, California.
Other reported localities: Carson Ridge, Marin County;
Bear Valley, Mariposa County; 10 miles S Monticello,
Napa County; 4 miles W Newcastle, Placer County;
Dutch Flat, Placer County; Placer Flat, Placer County;
7 miles NE Santa Rosa, Sonoma County; 6 miles N
Rumsey, Yolo County, California.

Parobisium hesperum (Chamberlin 1930). Type
locality: Cannon Beach, Clatsop County, Oregon.
Other reported locality: Dunsmuir, Shasta County,
California.

Parobisium hesternum Schuster 1966. Type lo¬
cality: Riverton, El Dorado County, California. Other
reported localities: Bear Valley and Yosemite National
Park, Mariposa County, California.

Parobisium utahensis Muchmore 1968. Type
locality: Blacksmith Fork Canyon, Cache County,
Utah.

Parobisium vancleavei (Hoff 1961). Type local¬
ity: Mesa Verde National Park, Montezuma County,
Colorado.

Parobisium yosemite Cokendolpher and Krejca,
n. sp. (this paper). Type locality: Indian Cave, Yosem¬
ite Valley, Yosemite National Park, Mariposa County,
California. Also reported from: Elf Village Cave,
Yosemite Valley.

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Occasional Papers, Museum of Texas Tech University

Key to Adults of the Genus Parobisium of North America

L Galea absent (Chamberlin 1962: Fig. 7H); large species, palpal femur

and tibia each longer than 2.0 mm. Parobisium charlotteae Chamberlin 1962

(troglophile, Deschutes County, Oregon)

Galea present, small rounded sclerotic knob (Figs. 25-26, 28); small

to medium sized species, palpal femur and tibia each 0.5-1.8 mm long.2

2. Anterior eyes reduced in size or absent, posterior pair absent; palpal femur

longer than 1.4 mm.3

Two pairs of eyes well developed; palpal femur shorter than 0.85 mm.4

3. Epistomal process a small extension (Fig. 5), palp patella with bulge on
mesal margin (Figs. 14-15), palpal hand elongated in dorsal and ventral

view (Fig. 15). Parobisium yosemite Cokendolpher and Krejca, n. sp.

(troglobite, Mariposa County, California)

Epistomal process absent, palpal patella smooth on mesal margin, palpal

hand rounded in dorsal and ventral view (Schuster 1966: Fig. 12).

. Parobisium hesperum (Chamberlin 1930)

(humus, Clatsop County, Oregon; Shasta County, California)

4. Smaller robust species, palpal femur 0.57-0.64 mm long, California.5

Medium sized species, palpal femur 0.72-0.84 mm long, Utah, Colorado.6

5. Galeal knob a long, low crest (Schuster 1966: Fig. 9); patella II with
11-12 setae; coxa IV with about 12 setae; anterior genital area (sternite II)

of male with about 20 setae. Parobisium hastatum Schuster 1966

(humus, El Dorado, Marin, Mariposa, Napa, Placer, Sonoma, Yolo counties, California)

Galeal knob a short, rounded crest (Schuster 1966: Fig. 11); patella II
with 18 setae; coxa IV with about 16 setae; anterior genital area (sternite II)

of male with about 30 setae. Parobisium hesternum Schuster 1966

(humus, El Dorado, Mariposa counties, California)

6. Posterior margin of carapace with 10-12 setae; palpal femur of male
3.9-4.1 times as long as broad, chela 3.3-3.4 times as long as broad;

chelal fingers with 45-59 marginal teeth. Parobisium vancleavei (Hoff 1961)

(beneath rocks in pinyon-juniper woodland, Montezuma County, Colorado)

Posterior margin of carapace with 8-10 setae; palpal femur of male 4.2-4.4 times as
long as broad; chela 3.7-3.8 times as long as broad; chelal fingers with 62-82

marginal macrodenticles. Parobisium utahensis Muchmore 1968

(moss on rocks, Cache County, Utah)

COKENDOLPHER AND KREJCA-CAVERNICOLOUS PSEUDOSCORPION

5

Parobisium yosemite Cokendolpher and Krejca,
new species

Figs. 1-29

Type material and treatments [75% ethanol —►
lactophenol^-75% ethanol = 75elpe; 95%
ethanol^>lactophenol^>75% ethanol = 9 5 elpe ].—
U.S.A.: California: Mariposa County: Indian Cave
(type locality): Holotype male, TTU-Z 51528 (Zara
3524), Journal Room, floor, 4 March 2009, leg. J.
Krejca, K. McDermid, 95% ethanol, appendages
-80°C, remainder 95elpe. Paratype male, TTU-Z 51540
(Zara-940), Journal Room, 15 August 2006, leg. J.
Krejca, G. Stock, 75elpe. Paratype female, TTU-Z
51548 (Zara-915), Journal Room, 16 August 2006,
leg. J. Krejca, G. Stock, elpe, appendages dried and
mounted on SEM stub. Paratypes 2 females, TTU-Z
51584, 51588 (Zara 3532), Journal Room, 4 March
2009, leg. K. McDermid, J. Krejca, 95% ethanol,
appendages -80°C, remainder room temperature. Not
exposed to lactophenol. Paratypes 1 male, 1 female,
CAS (formerly TTU-Z 51564 and 51536, Zara 3563),
1 female paratype, FSCA (formerly TTU-Z 51532),
Journal Room 3 March 2009, leg. J. Krejca, K. Mc¬
Dermid, G. Stock, R. Sas, 95elpe. Male paratype,
TTU-Z 51576, West Branch, 7 July 2009, leg. J. Kre¬
jca, poorly preserved - died in captivity, 75elpe. Male
paratype, TTU-Z 51572, East Branch, 7 July 2009, leg.
G. Stock, died in captivity - abdomen missing, poorly
preserved, elpe. Female paratype, TTU-Z 51568, West
Branch, 7 July 2009, leg. J. Krejca, died in captivity,
100% ethanol, -80°C, not exposed to lactophenol.
Female paratype, TTU-Z 51556 (Zara 3549), East
Branch, floor, 3 March 2009, leg. G. Stock, J. Krejca,
K. McDermid, R. Sas, 95elpe. Tritonymph paratype,
TTU-Z 51544 (Zara-920), Eastern Arm, 16 August
2006, leg. J. Krejca, G. Stock, elpe. Deutonymph
paratype, TTU- 51552 (Zara 3549), East Branch, floor,
3 March 2009, leg. G. Stock, J. Krejca, K. McDer¬
mid, R. Sas, 95elpe. Deutonymph paratype, TTU-Z
51564 (Zara 3563), Journal Room, 3 March 2009, leg.
J. Krejca, K. McDermid, G. Stock, R. Sas, 95elpe.

U.S.A.: California: Mariposa County: ElfVillage
Cave: Paratype female, TTU-Z 51580 (Zara 4175), 8
July 2009, leg. J. Krejca, left appendages in 100%
ethanol, -80°C, body elpe.

Etymology. —The specific name Yosemite is
used as a noun in apposition and is taken from the
known localities which are in Yosemite (pronounced
“yo-SEM-it-ee”) Valley in Yosemite National Park of
the U.S.A. The area was apparently named after the
Yosemite Indian peoples (http://theautry.org/yosemite/
[accessed 1 Sept. 2010]).

Vernacular name. —Yosemite Cave Pseudoscor¬
pion.

Diagnosis. —With the characters of the family,
subfamily, and genus. Posterior eyes absent, ante¬
rior pair reduced to a small, irregular shaped eyespot
without reflective tapetum (Fig. 5), epistomal process
a small extension (Fig. 5), cheliceral galea a small
rounded knob (Figs. 25-28), palpal patella with small
bulge mesally (Figs. 14-15), chelal fingers with 68-95
teeth, carapace 0.98-1.31 mm long.

Identification. —The new species can be told apart
from all described congeners from North America by
the key provided above. Parobisium hesternum was
recorded from the Yosemite National Park and Bear
Valley by Schuster (1966), but no habitat data were as¬
sociated with the collection. It is unknown if they were
from the bottom of a valley or on top of the mountains.
The new species and Parobisium hesternum differ pri¬
marily in the more elongated palp and loss of eyes in
P. yosemite. The new species differs also by having
a bulge on the mesal margin of the palp patella (Figs.
14-15) and in setation: P. yosemite (followed by P.
hesternum in brackets) patella II with 21-24 [18] setae;
coxa IV with about 11-12 [16] setae; anterior genital
area of sternite II of male with 58 [30] setae.

Description. —Coloration of abdomen and legs
light tan to amber; prosoma, coxa and trochanter
I-II, chelicerae, and palps reddish-brown (Figs. 1-4).
Nymphs are similarly colored, but darkest areas are
only light orange in color with lighter areas being cream
to yellowish-amber.

Male (holotype): Body length 3.46 mm. Carapace
a little longer than broad: widest in anterior fourth 0.99
mm; length 1.31 mm. Derm smooth; epistomal process
small and rounded (Fig. 5). Posterior eyes absent, an¬
terior pair reduced to a small, irregular shaped eyespot.

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Occasional Papers, Museum of Texas Tech University

Chaetotaxy 4-5-3-4-2-6 (24); 2 pairs of small, medial
lyrifissures at % and % carapace length (Fig. 5).

Abdomen elongated, 2.12 mm in length; widest
in middle, 1.36 mm wide; width greater than width of
carapace. Pleural membrane granulated (Fig. 9). Terg-
ites and sternites entirely smooth (Figs. 9-10). Tergites
wider than long. All tergites with setae in a single row:
chaetotaxy 8:13:12:13:12:12:12:12:10:11:6 (121).

Male genital opening with 58 microsetae on
sternite II. Stemite III with anteromedial cluster of 11
smaller setae near anterior margin, followed posteriorly
by row of 4 setae, then row of 18 setae; total 34-35
setae on sternite III (Fig. 11). Internally, the ventral
base of the genital operculum with 6 and 7 socketed
spines bounded by larger triangular denticle on each
side (Fig. 13).

Sternites V-VIII with 2 rows of setae, other ster¬
nites with single setal row posteriorly, plus anterior
pair near midline, chaetotaxy IV-X: 27-0+13-2+15-
2+15-2+13-12-10 (111). Anal plate with 2 anterior, 2
posterior setae.

Chelicera 1.05 mm long and 0.27 mm deep;
moveable finger 0.52 mm long, with 9 macrodenticles,
galea smooth, rounded, very short knob; with 2-4 pores
(see Figs. 28-29); palm with 7 setae and one lyrifissure
(Fig. 25: If); fixed finger with 13 (9 larger) macroden¬
ticles; rallum with two groups of blades: (1) with 5
setae (distal 2-3 with distal serrations) (Fig. 27, arrow
on right), (2) three shorter hirsute filaments (Fig. 27,
left arrow). Chaetotaxy of coxae: palp 6-6, legs 8-8:9-
10:9-9:11-12. Posterior maxillary lyrifissure curved
posteriorly at both ends (Figs. 7-8).

Palp long and slender, surface of each podom-
ere with granules (Figs. 15, 18-24), except for hand
smooth; granulations reduced or absent on dorsal side
of patella and distal end of femur. Palpal trochanter
0.99 mm long, 0.33 mm deep; femur 1.82 mm long,
0.33 mm deep; patella 1.79 mm long, 0.42 mm deep;
chela (tibia) 3.0 mm long, 0.64 mm deep, 0.68 mm
tall; moveable finger (tarsus) 1.705 mm long. Apex
of palpal coxa (manducatory process) curls around
dorsally and extends anteriorly, rounded, with 4 (other
type specimens with 4-6) setae on acute mesal edge.
Venom duct only present in fixed palpal finger; short,
not extending past A distance to et setae (Fig. 24). Palp

chelal trichobothriotaxy: st close to t, sb closer to st than
to on fixed finger it, et, and est near distal end, isb,
esb, and eb at base (Fig. 24), and ib and ist on dorsum
of hand (Fig. 18). Fixed chelal finger with 91 teeth;
moveable finger with 85 teeth.

Legs relatively long and thin. Patella II with 24
setae. Leg IV: (femur + patella) 1.13 mm long, 0.30
mm deep; tibia 1.07 mm long by 0.17 mm deep. Long
tactile setae on tibia and both tarsi of leg IV. Claws
long and smooth, flanked on each side with subter¬
minal tarsal seta deeply bifurcate (bifurcation with a
few further microtricia, Fig. 6), arolium less than half
length of claws.

Female (Indian Cave TTU-Z 51548, followed
by Elf Village Cave TTU-Z 51580 in parentheses or
brackets). Features as in male, except in details listed
below: Body length 4.17 (3.67) mm. Carapace a little
longer than broad: widest in middle, 0.92 (1.02) mm;
length 0.98 (1.22) mm. Chaetotaxy 4-5-4-1-6 (20) [4-
4-2-2-4-5 (21)]; pairs of lyrifissures in median position
at l A and 3 A distance of carapace.

Abdomen elongated 3.275 (2.65) mm in length;
widest in middle, 1.24 (1.205) mm. Tergites all with
setae in a single row: chaetotaxy 8-13-14-15-13-12-
13-12-11-12-9 (132) [8:11:12:12:13:11:11:12:13:14:
10(127)].

Female genital opening with a pair of lyrifissures
anteromedially (Fig. 12), 14 (7 right-7 left) [12 (4-8)]
microsetae on sternite II and single row of 34 (37)
setae (3-4 at each spiracle smaller than other setae) on
sternite III (Figs. 9-10); tracheae visible (Fig. 9); details
of internal genitalia as in Fig. 10.

Sternites VI (V in Elf Village Cave)-VIII with
2 rows of setae, others with single row posteriorly,
anterior pair near midline or % line and of similar size
and form to posterior setae: IV-X chaetotaxy: 29-0+15-
2+17-2+14-2+15-12-12 (120), [22-2+13-2+14-2+15-
2+12-14-13 (111)]. Anal plate with 2 (2) anterior, 2
(2) posterior setae. Genitalia with atrium between
sternites II and III; small median genital sac (Fig. 10,
arrow 5) covered with pores and extending posteriorly
from the cribiform gonosacs. The cribiform plate holes
are noticeably smaller in diameter than those ending as
pores on the medium genital sac.

COKENDOLPHER AND KREJCA-CAVERNICOLOUS PSEUDOSCORPION

7

Chelicera 0.71 (0.745) mm long and 0.32 (0.27)
mm deep; moveable finger 0.445 (0.46) mm long,
with 7 (6) macrodenticles (and few microdenticles),
galea a smooth short rounded knob; palm with 7 (7)
setae; fixed finger with 14 (13) macrodenticles; rallum
with 8 (7) setae, the first 2 (2) setae with short serra¬
tions (at or about width of setae), distal 3 (2) setae
with long (1.5 width of setae) serrations; 3 (3) shorter
hirsute filaments (Fig. 27). Chaetotaxy of coxae: palp
4-6 (4-5), legs 8-10-5-12-8-7-5-10 (65), [12-10-6-12-
11-9-5-14 (79)].

Palp relatively long and slender when compared
to other congeners, surface of each podomere with
granules, except for most (basal and distal) of hand
smooth (Figs. 23-24); granulations reduced or absent
on dorsal side of patella and distal end of femur. Me¬
dian maxillary lyrifissures of palpal coxae appearing
elongate oval in Figs. 7-8, because of the angle of view,
round in median view. Palpal trochanter 0.71 (0.825)
mm long, 0.26 (0.29) mm deep; femur 1.44 (1.625) mm
long, 0.265 (0.265) mm deep; patella 1.35 (1.56) mm
long, 0.37 (0.39) mm deep; chela (tibia) 2.42 (2.75) mm
long, 0.72 (0.63) mm tall; moveable finger (tarsus) 1.24
(1.415) mm long. Fixed chelal finger with 82 (74) teeth;
moveable finger with 77 (68) teeth (Figs. 18, 20-23);
teeth are more pointed anteriorly on fixed finger.

Legs relatively long and thin compared to other
congeners. Leg IV: (femur + patella) 1.18 (1.05) mm
long, femur 0.235 (0.18) mm deep, patella 0.18 (0.22)
mm deep; tibia 1.05 (0.995) mm long by 0.15 (0.13)
mm deep. Long tactile setae on tibia at A length and
both tarsi of leg at x h and !4, respectively.

Nymphs: As in adults, leg claws long and smooth,
flanked on each side with subterminal tarsal seta deeply
bifurcate, arolium less than half length of claws in
deutonymph and tritonymph. Eyespots not evident
because cuticle is not darkly pigmented as in adults;
no hint of a nerve fiber is evident.

Distribution .—This species has only been found
in two granitic talus caves located less than 0.5 km
apart, Indian Cave and Elf Village Cave in Yosemite
Valley, Mariposa County, California (Fig. 30). These
caves are located in a single expansive talus slope.
Extensive hand searching and Berlese extraction of

leaf litter at the second most extensive known cave
in the valley, Spider Cave (approximately 2 km to the
west), yielded no pseudoscorpions, except for a nymph
of Apochthonius sp. (Chthoniidae). Extensive hand
searching and Berlese extraction at other unnamed
minor talus caves and at nearby surface locations in
the talus piles yielded other pseudoscorpions, but no
specimens of Parobisium (Zara 2009).

Habitat .—Indian Cave and Elf Village Cave are
granitic talus caves (Figs. 31-32). The initial struc¬
ture of the talus caves was formed instantaneously as
a result of a rock avalanche (Wieczorek et al. 1999),
with subsequent weathering that may serve to make
the habitat more cave-like (e.g., stable temperatures
and humidities) as sediments and smaller rocks seal
off alternate entrances and surface water runoff creates
regular pathways for water flow. Indian Cave consists
of a series of passages and small rooms surveyed to 64
m long with a total depth below the entrances of 14 m.
The present cave probably formed relatively recently,
perhaps only a few hundred years ago (Wieczorek et
al. 1999); however earlier talus caves almost certainly
existed in the area as a result of rock falls from the
valley walls following retreat of the Last Glacial Maxi¬
mum glacier approximately 15,000-20,000 years ago
(Huber 1987). Even before the last glacial retreat, there
were talus slopes uphill that were formed after the first
glaciations about a million years earlier.

The entrance to Indian Cave (Fig. 31) consists of
two discreet openings among boulders approximately
15 meters up a steep talus slope from a relatively flat
terrace in the valley (Fig. 33). After a short climbdown
from either opening is the first room of the cave, the
Twilight Room, typified by low level sunlight, mossy
walls, and a soil and leaf litter floor. After another short
climbdown the passage splits, with the large obvious
way being to the northeast, the Junction Room. This
room marks the end of the light zone and a series of
large steps downward and to the east in increasingly
smaller passage leads to the East Branch of the cave.
We found Parobisium yosemite in this branch. The
substrate here consists of granite sand, rocks, leaf litter,
and occasional rootlets with and without mold. From
the Twilight Room climbdown to the southwest, a
discreet smaller passage (West Branch, Fig. 35) winds
under the entrance itself and continues as a crawlway

8

Occasional Papers, Museum of Texas Tech University

with a granite gravel and sediment floor, past some
woody debris, over a perennial pool, past a couple nar¬
row restrictions and down another climbdown into the
terminal room of the cave, the Journal Room. We also
found Parobisium yosemite in this branch. The Journal
Room has a sediment floor with sparse woody debris
and leaf litter, and scattered loose rocks (Fig. 34).

The entrance to Elf Village Cave (Fig. 32) is a
small (0.5 m) opening at the base of a large (10 m+)
boulder. Approximately 4 m of crawlway passage leads
into a long and wide low room (over 10 m diameter,
1-2 m tall) that is situated at an approximately 30 de¬
gree slope. At the low end of that room is a moist area
with a rock, gravel, and sand floor that is scattered with
acorns and pine cones. This is the only part of the cave
where we found Parobisium yosemite. Caves where
the species were not found were shallower and shorter.
These sites often had 15 m or less of traversable cave
passage, had multiple entrances that may contribute
to a lack of stable temperature and humidity, and most
of the extent of these caves was in the twilight zone
rather than having true darkness like Indian and Elf
Village caves. Spider Cave was the exception, with a
true dark zone and extensive passage. At present the
species is limited to the rockslide that encompasses the
two known localities, but it is possible that the species
is less detectable at Spider Cave due to some other
factor such as frequent human visitation or a natural
environmental parameter.

Microhabitat (Figs. 34-35).—During visits in
August 2006, March 2009, and July 2009 we recorded
substrate and temperature data for 40 different Paro¬
bisium yosemite observations (not all specimens were
collected). We found most individuals on the under¬
sides of rocks (60%, or 24/40 observations), and some
of those rocks were on woody debris, on sand, or on a
gravel floor with rootlets and fungus. The next most
common microhabitat was wood (33%, or 13/40 obser¬
vations). Individuals were typically on the undersides
of pieces of wood, and occasionally on the top of the
wood. We found the remaining 7% (3/40 observations)
of individuals on a granite and sand floor; one was
under granitic rock on sand floor. Total available sub¬
strate type was not measured, however our qualitative
assessment of available habitat indicates a preference
for undersides of rocks and woody debris, considering

the majority of the floor area in the deep portions of the
cave is breakdown, gravel, and granitic sand.

Temperatures ranged from 8.0-15.8°C with a
mean of 10.5°C and standard deviation of 1,8°C. Rela¬
tive humidity ranged from 73.2-100% with a mean of
92.7% and standard deviation of 7.3%. Sources of
nutrients for this cave ecosystem include leaf litter and
woody debris washed into the cave, rootlets penetrat¬
ing into the cave, and possibly trogloxenes including
small mammals. We did not observe a large amount of
small mammal scat, however in some places it seemed
unlikely that acoms were washed in by rains, but more
likely carried in by mammals. Some of the woody
debris appears too large to have been carried in by
animals, thus it is possible that it was captured during
the rockslide event that formed the cave or carried into
the cave by early explorers (Fig. 35).

Abundance. —During visits in August 2006,
March 2009, and July 2009 we recorded survey effort
for 39 different Parobisium yosemite observations.
At Indian Cave, in the zones where the species was
found, one individual was found on average every
22 minutes (total seen = 36, total search effort = 784
person-minutes). At Elf Village Cave one individual
was found on average every 11 minutes (total seen =
3, total search effort = 33 person-minutes).

Prey. —On 7 July 2009 in the Journal Room of
Indian Cave we turned a rock over to find one Parobi¬
sium yosemite with a springtail in its chelicerae. The
entomobryid, on field identification, appeared to be the
fairly common small, white Sinella or Pseudosinella
springtail. Other potential prey items we observed
commonly included two species of free-living mites,
flies, small spiders (nesticid and pimoid), beetles, ants,
and millipedes. The larger silver Tomocerus springtails
are more commonly seen near the cave entrance, not
overlapping with the areas where Parobisium yosemite
was seen.

Three adult specimens we held in captivity in
Petri dishes that were half-filled with Plaster of Paris
and a small wax-lined hole for holding free water.
A variety of small to tiny invertebrates were offered
as food: snails, slugs, earthworms, nematodes, flies,
ants, aphids, collembolla, moths, small crickets, and

(text continues on page 23)

COKENDOLPHER AND KREJCA-CaVERNICOLOUS PSEUDOSCORPION

9

Figures 1-4. Parobisium yosemite, n. sp. from Indian Cave, Yosemite National Park. Males
in cave: 1, dorsolateral aspect; 2, dorsal aspect. Females: 3, TTU-Z 51568 in laboratory,
dorsal aspect; 4, in cave, anterodorsal aspect. Photographs not at same scales, but females are
larger. The animals with outstretched palps and fingers open are actively moving through the
environment. The female with palps pulled in closer to the body is in more of a resting position.
An animal at full rest will have the palps closed and pulled in close to the body (JCC, pers.
obs.). Photographs 1, 2, 4 by JKK, photograph 3 by JCC.

Figures 5-6. Parobisium yosemite, n. sp. from Indian Cave, Yosemite National Park. 5, holotype male
(TTU-Z 51528) carapace, dorsal (anterior is to left; posterior pair of lyrifissures added and indicated by
arrows - see text). 6, female (TTU-Z 51548) tarsus IV claw with bifurcate setae (indicated by arrows).
Scale, 5 = 0.25 mm, 6 = 0.05 mm.

10

Occasional Papers, Museum of Texas Tech University

Figures 7-8. Paratype female (TTU-Z 51548) Parobisiumyosemite, n. sp. from Indian Cave, Yosemite National Park.
7, palp and leg coxae. 8, left palp coxa. Rounded median maxillary lyrifissures (arrows pointing left - see text) and
curved posterior maxillary lyrifissures (arrows pointing right). Scale = 0.15 mm.

COKENDOLPHER AND KREJCA-CaVERNICOLOUS PSEUDOSCORPION

11

Figures 9-10. Paratype female (TTU-Z 51548) Parobisium yosemite, n. sp. from Indian Cave,
Yosemite National Park. 9, sternites III and IV with spiracles positions noted with arrows; the
trachea are easily visible, pleural membrane granulations indicted by circle. 10, enlarged view
of genitalia: 1 = atrium, 2 = left lyrifissure, 3 = left group of seven setae on genital sternite, 4 =
series of pores of lateral apodeme, 5 = medium genital sac below the pore covered gonosacs, la
= part of cribriform plate of the lateral apodeme. Scales, 9 = 0.15 mm, 10 = 0.1 mm.

12

Occasional Papers, Museum of Texas Tech University

12

Figures 11-12. Parobisium yosemite, n. sp. from Indian Cave, Yosemite National Park
(TTU-Z 51528). 11, male genital sternites. 12, paratype female (TTU-Z 51548) genital
sternites with seven pairs of setae, arrows point to pair of lyrifissures. Scales, 11 =0.1 mm,
12 = 0.05 mm.

COKENDOLPHER AND KREJCA-CaVERNICOLOUS PSEUDOSCORPION

13

Figures 13-17. Parobisium yosemite, n. sp. from Indian Cave, Yosemite National Park. 13, holotype TTU-51528,
internal male genitalia (internal piece of genital operculum) with six and seven socketed spines bounded by larger
triangular denticles. 14-17 male paratype TTU-Z51576. 14, enlargement of palp patella base showing medial buldge,
ventral aspect; 15, palp, ventral aspect; 16, leg I, posterior aspect; 17, leg IV, posterior aspect. Scales, 13 = 0.063 mm,
14 = 0.167 mm, 15-17 = 0.5 mm.

14

Occasional Papers, Museum of Texas Tech University

Figures 18-19. Paratype female (TTU-Z 51548) palp chela of Parobisium yosemite, n. sp. from Indian
Cave, Yosemite National Park. Lateral aspect; 18, basal portion of hand broken off; 19, detail of hand
showing granulation.

COKENDOLPHER AND KREJCA-CaVERNICOLOUS PSEUDOSCORPION

15

Figures 20-21. Paratype female (TTU-Z 51548) tips of chelal fingers of Parobisiumyosemite,
n. sp. from Indian Cave, Yosemite National Park. 20, lateral aspect; 21, mesal aspect showing
longitudinally grooved distal tips (in yellow) and opening to the venom duct (red arrow).
Teeth more flattened on moveable (ventral) finger of palp.

16

Occasional Papers, Museum of Texas Tech University

Figures 22-23. Paratype female (TTU-Z 51548) palp chelal of Parobisium yosemite, n. sp.
from Indian Cave, Yosemite National Park. 22, basal group of macrodenticles of fingers;
more flattened on moveable (ventral) finger of palp, upper denticles appear eroded with
lateral rows of minute grooves; 23, trichobothria of lateral side; internal position of venom
gland drawn in red; granulations of hand also visible.

COKENDOLPHER AND KREJCA-CaVERNICOLOUS PSEUDOSCORPION

17

Figures 24-25. Paratype female (TTU-Z 51548) Parobisium yosemite, n. sp. from Indian Cave,
Yosemite National Park. 24, upside-down view of palp hand (moveable finger should open
ventrally), setal positions marked as well as in another view in Fig. 23 (damage to base of chela
and basal portion of hand broken off); 25, lateral view of chelicera; seven setae on the hand and
a setae on moveable finger are highlighted in yellow, macrodenticles of the fingers as well as the
lyrifissure (= If) are also marked.

18

Occasional Papers, Museum of Texas Tech University

Figures 26-27. Paratype female Parobisiumyosemite, n. sp. (TTU-Z 51548) from Indian Cave,
Yosemite National Park. 26, lateral view of chela tips; 27, mesal side of chela; five long hirsute
filaments (arrow on right), plus three shorter hirsute filaments (arrow on left) in rallum.

COKENDOLPHER AND KREJCA-CaVERNICOLOUS PSEUDOSCORPION

19

Figures 28-29. Galea on moveable finger of chelicera of paratype female (TTU-Z 51548) Parobisium yosemite, n.
sp. from Indian Cave, Yosemite National Park. 28, lateral aspect with 4 numbered pore positions; depression above
1 does not appear to be a pore; 29, anterior close-up of pores. Pores at 0.6 pm diameter (measured at much higher
resolution). Numbers 1, 4 are certainly pores, but the openings are less clear of the depressions at 2 and 3, but likely
pores of the galea (spinneret).

Figure 30. Map of Yosemite Valley and locations of caves in Yosemite National Park; insert is location within California.
Parobisium yosemite, n. sp. is recorded from Indian Cave and Elf Village Cave. Map by K. McDermid.

20

Occasional Papers, Museum of Texas Tech University

Figures 31-32. Entrances to caves in Yosemite National Park which are known
to have Parobisium yosemite , n. sp. 31, Sarah Stock in front of the double
entrances to Indian Cave; 32, G. Robert Myers, III (see middle bottom of
picture) in the entrance to Elf Village Cave. Photographs by JKK.

COKENDOLPHER AND KREJCA-CaVERNICOLOUS PSEUDOSCORPION

21

Woody debris

East

Branch

INDIAN CAVE

Mapped length: 63.9 m (209.6 fl)
Mapped depth: -14.0 m (45.9 ft)

Plan view

burvty by brag block ana Joel Dr-
Oafting by Grog Stock

connection

Twilight

Keen

Woody debris

Seasonal pool

Entrance

Journal

Room

Profile view, view to the northwest

Entrance 0.0 m

hd

-6,3 m

Figure 33. Map of Indian Cave, Yosemite National Park, showing locations mentioned in text. Map by
G. Stock.

22

Occasional Papers, Museum of Texas Tech University

Figures 34-35. Habitat within Indian Cave, Yosemite National Park, which is the type locality for
Parobisiumyosemite, n. sp. 34, Krista McDermid in the Journal Room with granite boulder walls and
ceiling, and smaller granitic rocks on sand and sediment floor with woody debris; 35, Greg Stock in
the West Branch at the woody debris. Photographs by JKK and Steven Bumgardner, respectively.

COKENDOLPHER AND KREJCA-CAVERNICOLOUS PSEUDOSCORPION

23

(text continued from page 8)

linyphiid, uloborid, and theridiid spiders. The only item
eaten was a small theridiid spider (probably Theridion).
Pseudoscorpions held in captivity would slowly walk
around the dish with palps extended and held open.
Whenever they encountered a new potential prey in the
container, they would tap the tips of the palps around
the animal or appendage. As soon as the animal was
touched the pseudoscorpion would retract the palps
and change direction of movement. After this first
encounter, the pseudoscorpion would not react upon
touching the potential prey again and would often just
walk over the potential prey. In one case an ant ( Sole-
nopsis invicta ) reacted to the pseudoscorpion and bit
and held onto a leg tarsus. The sting had been removed
from the ant prior to introduction into the Petri dish.

The ant was dead, but still clinging to the leg when first
observed, so it is not certain if the pseudoscorpion tried
to eat the ant or visa-versa.

Ecological Status. —Extensive searching (>27
person-hours) of similar microhabitats nearby but
outside of the granite talus caves where the species
was found yielded other pseudoscorpion species but no
examples of Parobisium yosemite. We searched under
rocks, in packrat nests, under bark and in leaf litter
using hand collections and Berlese funnel extractions
(Zara 2009). While it is impossible to prove absence,
this habitat association, combined with the eye loss and
propensity for cave-adaptation of pseudoscorpions, lead
us to believe this species is troglobitic.

Discussion

Troglobites are not well documented from talus
caves, and this may be the first North American troglo-
bite described from a talus cave, and possibly the first
troglobite described from a granitic bedrock region in
the world. Terrestrial troglobites are most commonly
found in caves formed in calcareous bedrock or lava
tubes, however in a few cases researchers described
them from other dark, high humidity voids in the sub¬
surface. Researchers found troglobitic fauna in Europe
in thick layers of talus covering karstic bedrock, the
presumed source of the subterranean fauna (Chris¬
tian 1987). These voids are described as the “milieu
souterrain superficiel” (MSS) (Juberthie et al. 1980),
characterized by cracks and fissures in the surface of the
bedrock and adjacent talus. The discovery of troglo¬
bites in this habitat has lead to more extensive searching
in the MSS in Europe, and even extended the range for
some species beyond areas of calcareous bedrock and
into regions of schists (Crouau-Roy 1989). However,
there are also MSS species described from Europe from
non-karstic MSS (e.g., Traegaardhia distosolenidia
Zacharda [Zacharda et al. 2010]).

In Australia, troglobites are documented in
atypical calcareous bedrock areas including pisolite,
calcrete, and also in a (primarily volcanic) greenstone
belt (Anonymous 2007, 2008; Barranco and Harvey
2008). In his summary of obligate cave-dwelling fauna
of the U.S.A. and Canada, Peck (1998) noted that the

MSS and other similar aquatic habitats are not well
explored in North America. In the U.S.A., Slay and
Bitting (2007) found the same number of troglobites
in mines dug into karstic bedrock as in natural caves,
and there are many records of subterranean fauna from
other types of voids in karstic rocks such as boreholes
and wells.

To our knowledge there is only one other report of
a troglobite from a granite talus cave in North America,
the pseudoscorpion Tyrannochthonius troglodytes
Muchmore from central Texas. Muchmore (1986)
described this species as cave adapted, though the type
locality is from the “twilight zone,” or near-entrance
area of the cave with some light and temperature
fluctuation, of a granite cave in Llano County, Texas.
This granite area in central Texas, with several talus
caves, is relatively small and surrounded on all sides
by extensive limestone karst (Smith and Veni 1994).
The granite area in Yosemite Valley, by contrast, is quite
large, with only isolated calcareous bedrock (limestone
and marble) being located farther, at least 15 km away.
Consequently the likelihood is fairly high that Paro¬
bisium yosemite evolved in a granite talus setting, to
which it is limited, whereas it is less certain that this is
the case for Tyrannochthonius troglodytes.

It is interesting to consider the immigration and
evolution of this troglobite given that these specific

24

Occasional Papers, Museum of Texas Tech University

caves were likely formed within the past 1,000 years
(Wieczorek et al. 1999). Although it is possible that
the species evolved in these most recent caves, more
likely they evolved in caves that are now eroded
away or buried, and immigrated to the caves that we
can access nearer to the surface. In either scenario,
the area was glaciated as recently as 15,000-20,000
ybp, indicating this is either a recently-evolved spe¬
cies or one that survived the lastest glaciation of the
area, possibly taking refuge in talus slopes above or
adjacent to the glaciers. The valley was formed over
a million years ago by glacial activity, but none of the
more recent glaciers have been as large and therefore
talus forming slopes have been available as a habitat
for the past million years. Another possibility is that
the species invaded after glaciers retreated. Given
that Parobisium yosemite occurs at two nearby caves
in the same talus slope, it is likely that further collect¬
ing effort at other caves in this rockfall will yield new
localities, and the rockfall area itself (approximately
1.5 km 2 ) could be considered the total known range for
the species. Even with this liberal definition, however,

its range is relatively small and conservation priority
should be given to this species. In these very small
passages, substrate compaction (Zara 2009) and small
chemical spills (e.g., insect repellent, batteries) could
play a large role in habitat.

Research priorities for this narrow endemic spe¬
cies include learning more about the talus cave ecosys¬
tem and the biology of the species. For example, what
other species occur in the same environment? Are there
species associations? Do the cave limited species rely
primarily on organic debris carried into the cave via
rains or does some other trogloxene species play a role?
In the case of other cave systems, bats or cave crickets
are important nutrient carriers, therefore the manage¬
ment of those species is critical for the cave ecosystem.
The biology of P. yosemite is poorly known. General
information on life history, including but not limited
to seasonality, longevity, fecundity, population size,
temperature and humidity tolerances, prey, and preda¬
tors, is needed for management and understanding of
this interesting species.

Acknowledgments

The National Park Service provided funding for
this project and the Yosemite Fund contributed funding
to the field work that lead to the discovery of the spe¬
cies. Krista McDermid, George R. (Rob) Myers III,
Robert Sas, Greg Stock, and Sarah Stock assisted with
the field work. Greg Stock initiated and coordinated
the various projects, performing a multitude of tasks
ranging from applying for the initial funding to detail¬
ing field logistics - his vision about the potential value
in these talus caves and motivation to sample them
resulted in the discovery of the new pseudoscorpion.

We thank Krista McDermid and Greg Stock for the
maps and Steven Bumgardner for help with a photo¬
graph. The collections were made under National Park
Service permit # YOSE-2009-SCI-0121. We thank the
Texas Tech University Imaging Center and Department
of Biological Sciences for facilities and guidance to
all things related to the Hitachi S-4300SE/N (grant #
NSF MRI 04-511). Mark Judson and Steve Taylor are
thanked for their helpful comments on a draft of the
manuscript.

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Addresses of authors:

James C. Cokendolpher

Natural Science Research Laboratory
Museum of Texas Tech University
Lubbock, TX 79409 USA
james. cokendolpher@ttu. edu

Zara Environmental LLC. 2009. Continuing biological in¬
ventory of talus caves in Yosemite Valley. Report
prepared for Yosemite National Park, 31 August
2009, 42 pp.

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Jean K. Krejca

Zara Environmental LLC
1707 W.FM 1626
Manchaca, TX 78652 USA
jean@zaraenvironmental. com

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Production Editor: Lisa Bradley

ISSN 0149-175X

Museum of Texas Tech University, Lubbock, TX 79409-3191