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Troglomorphic pseudoscorpions (Arachnida: Pseudoscorpiones) of northern Arizona, with the description of two new short-range endemic species

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This study reports on the pseudoscorpion fauna of the subterranean ecosystems of northern Arizona, U.S.A. Our work resulted in the descriptions of two new species, Hesperochernes bradybaughi sp. nov. and Tuberochernes cohni sp. nov. (Chernetidae) and the range expansion of one species, Larca cavicola (Muchmore 1981) (Larcidae). All of these species were cave-adapted and found within caves on Grand Canyon-Parashant National Monument in northwestern Arizona. Based upon this work, the genus Archeolarca Hoff and Clawson is newly synonymized with Larca Chamberlin, and the following species are transferred from Archeolarca to Larca, forming the new combinations L. aalbui (Muchmore 1984), L. cavicola (Muchmore 1981), L. guadalupensis (Muchmore 1981) and L. welbourni (Muchmore 1981). Despite intensive sampling on the monument, the two new species were detected in only one cave. This cave supports the greatest diversity of troglomorphic arthropod species on the monument—all of which are short-range endemics occurring in only one cave. Maintaining the management recommendations provided by Peck and Wynne (2013) for this cave should aid in the long-term persistence of these new pseudoscorpion species, as well as the other troglomorphic arthropods.
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Troglomorphic pseudoscorpions (Arachnida: Pseudoscorpiones) of northern Arizona, with the description
of two new short-range endemic species
Mark S. Harvey
1,2,3,4,5
and J. Judson Wynne
6
:
1
Department of Terrestrial Zoology, Western Australian Museum, Locked
Bag 49, Welshpool DC, Western Australia 6986, Australia;
2
Research Associate, Division of Invertebrate Zoology,
American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024-5192, U.S.A;
3
Research Associate, California Academy of Sciences, 55 Music Concourse Drive, San Francisco, California 94118,
U.S.A;
4
Adjunct, School of Animal Biology, University of Western Australia, Crawley, Western Australia 6009,
Australia;
5
Adjunct, School of Natural Sciences, Edith Cowan University, Joondalup, Western Australia 6027,
Australia;
6
Department of Biological Sciences, Colorado Plateau Biodiversity Center, Landscape Conservation
Initiative, Northern Arizona University, Box 5640, Flagstaff, Arizona 86011, U.S.A. E-mail:
mark.harvey@museum.wa.gov.au
Abstract. This study reports on the pseudoscorpion fauna of the subterranean ecosystems of northern Arizona, U.S.A.
Our work resulted in the descriptions of two new species, Hesperochernes bradybaughi sp. nov. and Tuberochernes cohni sp.
nov. (Chernetidae) and the range expansion of one species, Larca cavicola (Muchmore 1981) (Larcidae). All of these species
were cave-adapted and found within caves on Grand Canyon-Parashant National Monument in northwestern Arizona.
Based upon this work, the genus Archeolarca Hoff and Clawson is newly synonymized with Larca Chamberlin, and the
following species are transferred from Archeolarca to Larca, forming the new combinations L. aalbui (Muchmore 1984),
L. cavicola (Muchmore 1981), L. guadalupensis (Muchmore 1981) and L. welbourni (Muchmore 1981). Despite intensive
sampling on the monument, the two new species were detected in only one cave. This cave supports the greatest diversity of
troglomorphic arthropod species on the monument—all of which are short-range endemics occurring in only one cave.
Maintaining the management recommendations provided by Peck and Wynne (2013) for this cave should aid in the long-
term persistence of these new pseudoscorpion species, as well as the other troglomorphic arthropods.
Keywords: Nearctic, troglomorphy, troglobite, new synonymy, cave
urn:lsid:zoobank.org:pub:A15CB9DB-5B36-4A7C-8052-08E2EC1F4D34
The pseudoscorpion fauna of North American caves is
moderately well known, thanks largely to the efforts of J.C.
Chamberlin, C.C. Hoff, E.M. Benedict, D.R. Malcolm and W.B.
Muchmore who have characterized and described many different
North American troglobites and troglophiles. There are currently
144namedspeciesfoundincavehabitats across the United
States including six species in five families from Arizona:
Pseudogarypus hypogeus Muchmore 1981 (Pseudogarypidae),
Albiorix anophthalmus Muchmore 1999 (Ideoroncidae), Chit-
rellina chiricahuae Muchmore, 1996 (Syarinidae), Archeolarca
cavicola Muchmore 1981, A. welbourni Muchmore 1981
(Larcidae) and Tuberochernes ubicki Muchmore 1997 (Cherne-
tidae) (Muchmore 1996; Muchmore & Pape 1999; Harvey &
Muchmore 2013). Only A. anophthalmus and C. chiricahuae had
troglobitic modifications including the complete lack of eyes
and pallid body color (Muchmore 1996; Muchmore & Pape
1999; Harvey & Muchmore 2013), whereas the others are less
obviously modified with only the slightly attenuated append-
ages hinting at an obligate subterranean existence (Muchmore
1981, 1997).
Prior to this work, all of these cavernicolous species from
Arizona occurred south of the Colorado River with P.
hypogeus,A. cavicola and A. welbourni from northern Arizona
(Coconino County) and A. anophthalmus,C. chiricahuae and
T. ubicki from south-eastern Arizona (Pima, Cochise and
Santa Cruz Counties, respectively). During biological inven-
tories of caves on the Grand Canyon-Parashant National
Monument (hereafter referred to as Parashant) in northwest-
ern Arizona, one of us (J.J.W.) and colleagues found
representatives of three different pseudoscorpion species,
which are the subject of this study.
Over the past several years, Parashant caves have yielded
other significant and interesting arthropod species—many of
which are restricted to the cave environment. These include
two new genera (comprising two new species)—a book louse
(order Psocoptera, family Sphaeropsocidae: Troglosphaerop-
socus voylesi Mockford 2009 (Mockford 2009), and a cave
cricket (family Rhaphidophoridae: cf. Ceuthophilus n. gen. n.
sp., Cohn and Swanson, unpublished data). This work also
resulted in the identification of several cave-adapted and cave-
limited species including a leiodid beetle, Ptomaphagus
parashant Peck and Wynne 2013 (Peck & Wynne 2013), an
undescribed species of centipede (family Anopsobiidae;
Wynne, unpublished data), an undescribed Isopod species,
Brackenridgia n. sp. (S. Taiti, in litt.), and a recently described
cave limited millipede, Pratherodesmus voylesi Shear 2009
(Shear et al. 2009). Additionally, three new species of
trogloxenic beetles were reported from Parashant caves
including Eleodes wynnei Aalbu, Smith, and Triplehorn 2012
(Tenebrionidae; Aalbu et al. 2012), an undescribed species of
the carabid beetle genus Rhadine LeConte (Carabidae: the
perlevis species-group; T.C. Barr, in litt.), and an undescribed
carabid beetle species Pterostichus Stephens (Carabidae, K.
Will, in litt.).
METHODS
The junior author and colleagues sampled caves on Grand
Canyon-Parashant National Monument during 4–14 August
2014. The Journal of Arachnology 42:205–219
205
2005, 1–6 May 2007, 16–25 August 2007, 12–21 May 2008,
and 5–12 March 2009. They sampled all caves identified as
having deep zone like conditions (n510). Given the short
duration of study (between two to four site visits), and
potential seasonal effects, confidently identifying this zonal
environment was not possible. The cave deep zone is required
habitat for cave-adapted arthropods and is characterized by
complete darkness, stable temperature, a near-water saturated
atmosphere and limited to no airflow (as in Howarth 1980,
1982). Parashant is located in northwestern Arizona, encom-
passes approximately 4,451 km
2
, and is characterized by
rugged terrain containing deeply incised canyons, mesas, and
mountains. Vegetation zones include Mojave Desert contain-
ing creosote bush (Larrea tridentata) and Joshua trees (Yucca
brevifolia) at lower elevations, gradating through Great Basin
pinyon (Pinus edulis) and juniper (Juniperus spp.) woodlands
to Colorado Plateau grasslands and Ponderosa pine (Pinus
ponderosa) forest with aspen (Populus tremuloides) groves on
Mt. Trumbull (elevation 2,447 m). All of the caves referred to
in this paper were located within the Supai, Kaibab, or
Redwall limestone formations. Elevation for the caves that
were studied ranges from 736 to 1,590 m.
Although we inventoried 10 Parashant caves, we provide
descriptions for only the three caves (PARA-1001, PARA-
2204 and PARA-3503) where pseudoscorpions were detected.
PARA-1001 Cave was the second most biologically diverse
cave on Parashant (Wynne, unpublished data), and supports
the largest known cricket roost in northern Arizona (Wynne &
Voyles 2014). A small solution cave within the Kaibab
limestone, it had a total surveyed length and depth of 76.2 m
and 10.4 m, respectively. This cave had a small south-facing
vertical entrance (135uaspect) at bottom center of a large
sinkhole. Vegetation was characterized as juniper scrublands
at 1,585 m elevation, and was located on the north side of the
lower Colorado River along the western extent of the Grand
Canyon. PARA-2204 Cave was the most biologically diverse
cave on the monument (Wynne, unpublished data). The
deepest extent of this cave contained active speleothem
formations and supported a near-saturated water atmosphere
year-round. Located within the Supai formation, this large
solution cave (total surveyed length 175 m) was comprised of
several sinuous phreatic passages. This cave has one horizon-
tal entrance (330uaspect) and was situated within a canyon
near the base of the canyon’s north-face. Located at 1,272 m
elevation, this cave occurred within the vegetation transition
zone of Mojave Desert scrub and juniper woodlands. PARA-
3503 Cave was a dry cave with no evidence of recent
speleothem activity, and supported a summer roost of bats,
Myotis sp. (Wynne, unpublished data). The cave had a large
horizontal entrance (135uaspect) situated upon a high bench
(1,102 m elevation on an exposed cliff face). This cave was
situated along the south-face of one of the largest canyons
draining into the Colorado River from the north. Occurring
within the Redwall formation, this large solution cave
contained 540 m of surveyed length with an estimated survey
depth of 14.2 m. Vegetation was characterized as Mojave
Desert scrub.
The work conducted in 2005 was part of a biological
baseline study [refer to Wynne & Voyles (2014) for a
description of sampling methods]. Later (between 2007 and
2009), these caves were systematically sampled to characterize
the cave-dwelling arthropod communities. Interval sampling
using baited pitfall traps, timed searches, and opportunistic
sampling techniques were used. To apply these techniques,
detailed maps for each cave were required. For interval
sampling, we established up to 10 sampling intervals (which
included a sampling station at either wall and one at cave
centerline for a total of #3 sampling stations per interval). We
used 10%of the total cave length to establish the sampling
interval (e.g., for a 1,000 m long cave, the sampling interval
was every 100 m).
At each sampling station, we deployed live capture baited
pitfall traps and conducted timed searches. For pitfall traps,
we used two 907 g stacked plastic containers (13.5 cm high,
10.8 cm diameter rim and 8.9 cm base). A teaspoon of peanut
butter was used as bait and placed in the bottom of the
exterior container. At the bottom of the interior container, we
made several dozen holes so the bait could ‘‘breathe’’ to
attract arthropods (e.g., Barber 1931). Attempts were made to
counter-sink each pitfall trap within the cave sediment or
rockfall. When this was not possible, we built ramps around
each trap using local materials (e.g., rocks, wooden debris,
etc.) so arthropods could access the trap and fall in (e.g.,
Ashmole et al. 1992). To discourage small mammals, we
placed small rocks around the edges of the trap and then
covered the mouth of the trap with a cap rock. Pitfall traps
were deployed for three to four days (a three day deployment
occurred once due to scheduling constraints). For timed
searches, we established a 1 m radius around each sampling
station (where the pitfall trap would be deployed) and
searched for arthropods within that ,3 m circle. A one to
three minute timed search (one minute if no arthropods were
observed, three minutes if arthropods were detected) was
conducted before pitfall trap deployment and prior to trap
removal.
Opportunistic collecting was executed by two to three trained
observers as they traversed the length of each cave. This
technique was applied as the observers were in transit between
sampling intervals while deploying and removing pitfall traps
and conducting timed searches. Opportunistic collecting was
not conducted while at sampling stations and was resumed only
when the observers were in transit once again. This technique
was used at least twice per cave (both during pitfall trap
deployment and retrieval trips). For example, a cave containing
10 sample station arrays, there were 27 individual ‘‘random
walks’’ per site visit (i.e., nine random walk samples times three
observers collecting along their between stations). Because we
conducted two site visits per cave, there would be a total of 54
samples. For one cave, PARA 1001 Cave, we had two observers
conduct the opportunistic collecting.
An alpha-numeric coding system developed by the National
Park Service (NPS) was used to safeguard the location of both
caves and their resources. We only provide generalized latitude
and longitude coordinates of the area to keep the precise
location of the cave confidential. Parashant National Monu-
ment headquarters in Saint George, Utah has the cipher table
with cave codes. A copy of this paper with actual cave names
is on file at both monument headquarters, National Park
Service and the National Cave and Karst Research Institute,
Carlsbad, New Mexico.
206 THE JOURNAL OF ARACHNOLOGY
Specimens representing three species collected by one of us
(J.J.W.) and colleagues form the basis of this study. All
specimens were collected and stored in 70%ethanol. The
holotypes of both new species and specimens of the known
species are deposited in the Museum of Northern Arizona,
Flagstaff, Arizona (MNA). Temporary slide mounts were
prepared by mounting them on microscope slides with 10 or
12 mm coverslips supported by small sections of 0.25, 0.35 or
0.50 mm diameter nylon fishing line in a drop of lactic acid at
room temperature for two or more days. After study the
specimens were rinsed in water and returned to 75%ethanol
with the dissected portions placed in 12 33 mm glass genitalia
microvials (BioQuip Products, Inc.). All specimens were
studied using a Leica DM2500 compound microscope and
illustrated with the aid of a drawing tube. Measurements were
taken at the highest possible magnification using an ocular
graticule. Terminology and mensuration mostly follow
Chamberlin (1931), with the exception of the nomenclature
of the pedipalps, legs and with some minor modifications to
the terminology of the trichobothria (Harvey 1992), cheliceral
setation (Harvey & Edward 2007), cheliceral rallum (Judson
2007) and faces of the appendages (Harvey et al. 2012).
TAXONOMY
Family Larcidae Harvey 1992
Larca Chamberlin 1930
Larca Chamberlin 1930:616.
Archeolarca Hoff and Clawson 1952:2–3. Syn. nov.
Type species.—Larca: Garypus latus Hansen 1884, by
original designation.
Archeolarca: Archeolarca rotunda Hoff and Clawson 1952,
by original designation.
Remarks.—The genus Larca was created by Chamberlin
(1930) for the type species L. lata (Hansen) from Europe and
L. granulata (Banks 1891) from eastern U.S.A. Since then,
other species have been added from Europe (Beier 1939a;
Gardini 1983; Henderickx & Vets 2002; Zaragoza 2005) and
North America (Hoff 1961; Benedict & Malcolm 1978;
Muchmore 1981). Archeolarca was described for the type
species A. rotunda which was collected from pack rat middens
and porcupine nests in Utah (Hoff & Clawson 1952). Since
then, four additional species have been described from other
parts of western North America, all from cave ecosystems
(Muchmore 1981, 1984), and A. rotunda has been found in
New Mexico and Oregon (Hoff 1956a; Benedict & Malcolm
1978). Archeolarca only differs from Larca in the possession of
four trichobothria on the movable chelal finger of adults,
whereas species of Larca have only two or three trichobothria
(e.g. Hoff 1961; Benedict & Malcolm 1978; Muchmore 1981;
Gardini 1983; Muchmore 1984, 1990; Henderickx & Vets
2002; Zaragoza 2005). Most adult specimens from the
Parashant have four trichobothria on the movable chelal
finger (Fig. 12), consistent with being a species of Archeolarca,
but one male has four on the right chela and three on the left
(Fig. 11) raising the issue of whether the genera should be
retained.
The maintenance of garypoid genera based solely on
trichobothrial number has been abandoned for several other
groups including the garypid genera Anagarypus Chamberlin
1930 with seven trichobothria on the fixed finger and one or
two on the movable finger forming a pattern of 7/1–2
(Muchmore 1982), Eremogarypus Beier 1955, with a pattern
of 5–8/1–3 (e.g., Beier 1962; Beier 1973), Synsphyronus
Chamberlin 1930, with a pattern of 5–8/1–3 (e.g., Chamberlin
1943; Harvey 1987b, 2011) and Thaumastogarypus Beier 1947,
with a pattern of 7–8/3–4 (e.g. Beier 1947; Mahnert 1982), and
the geogarypid genus Geogarypus Chamberlin 1930 in which
adults normally have an 8/4 pattern, but G. bucculentus Beier
1955 and G. connatus Harvey 1987 have a 7/4 pattern (Harvey
1986, 1987a). Intra-specific variation in the number of
trichobothria of the movable chelal finger has been reported
in the genus Serianus Chamberlin 1930 (Garypinidae). Hoff
(1950) found that a small series of specimens of S. minutus
Hoff 1950 (now known as S. argentinae Muchmore 1981 due
to secondary homonymy of the original name) included adults
with the normal four trichobothria on the movable chelal
finger, as well as some with only two or three trichobothria.
Similarly, Mahnert (1988) found that the type series of
Paraserianus bolivianus Beier 1939 possessed three or four
trichobothria on the movable chelal finger. Given that the
main feature used to substantiate the genus Paraserianus by
Beier (1939b) was the presence of only three such trichobo-
thria (as opposed to four in Serianus), Mahnert (1988) placed
Paraserianus as a synonym of Serianus.
Comparison of specimens of many species of Larca and
Archeolarca by one of us (M.S.H.), including the type species
of both genera, has revealed no other significant differences
that could be considered to maintain distinct genera, and
Archeolarca is here regarded as a synonym of Larca, resulting
in the following new combinations: L. aalbui (Muchmore
1984), comb. nov.,L. cavicola (Muchmore 1981), comb. nov.,L.
guadalupensis (Muchmore 1981) comb. nov. and L. welbourni
(Muchmore 1981) comb. nov.
Larca cavicola (Muchmore) comb. nov.
(Figs. 1–14)
Archeolarca cavicola Muchmore 1981: 55–56, Figs. 11, 12.
Material examined.—U.S.A.: Arizona: Mohave County: 1
male, PARA-3503 Cave, Grand Canyon-Parashant Nation-
al Monument, ca. UTM 0247400 N, 4020000 E, Zone 12S,
baited pitfall trap 1A, 20 May 2008, J.J. Wynne (MNA); 1
female, same data except baited pitfall trap 1C, 6 March
2009, J.J Wynne (MNA); 1 tritonymph, 1 deutonymph,
same data except trap 2B, 10 March 2009, J.J. Wynne
(MNA); 1 tritonymph, same data except trap 7A (MNA); 1
tritonymph, same data except opportunistic collecting in a
possible deep zone (MNA); 1 male, PARA-2204 Cave,
Grand Canyon-Parashant National Monument, ca. UTM
025100 N, 4041000 E, Zone 12S, M, baited pitfall trap 2B,
17 May 2008, J.J. Wynne (MNA); 1 female, same data
except 20 May 2008 (MNA); 1 tritonymph, same data
except trap 1A (MNA); 1 male, same data except trap 1B
(MNA).
Diagnosis.Larca cavicola resembles the other species
previously included in the genus Archeolarca in possessing
four trichobothria on the movable chelal finger, but occasion-
ally this is reduced to three trichobothria. It differs from these
species by having reduced eyes, especially the posterior pair,
which are noticeably smaller than the anterior pair.
HARVEY & WYNNE—TROGLOMORPHIC PSEUDOSCORPIONS FROM ARIZONA 207
Description.—Adults: Color: carapace, pedipalps and coxae
deep red-brown, abdomen pale red-brown and legs pale
yellow-brown.
Chelicera: with 4 setae on hand, with sbs absent, and 1
subdistal seta on movable finger (Fig. 7); all setae acuminate;
seta bs slightly shorter than others; with 2 dorsal lyrifissures
and 1 ventral lyrifissure; galea of Land Kvery long with 3
terminal rami, rami of male smaller than on female; rallum of
4 blades, the most distal blade with several serrations on
leading edge, other blades smooth; serrula exterior with 14 (L),
16 (K) blades; lamina exterior present.
Pedipalp (Fig. 9): most surfaces of trochanter, femur,
patella and chelal hand lightly and sparsely granulate, chelal
fingers smooth; trochanter, femur, patella and chelal hand
with prominent, curved, slightly denticulate setae arranged
sparsely; patella with 3 small sub-basal lyrifissures; trochanter
1.83–1.99 (L), 1.90–1.93 (K), femur 4.74–5.94 (L), 4.57–4.95 (K),
patella 3.63–4.47 (L), 3.69–3.94 (K), chela (with pedicel) 4.47–
5.28 (L), 4.08–4.54 (K), chela (without pedicel) 4.22–5.02 (L),
3.85–4.26 (K), hand (with pedicel) 2.17–2.49 (L), 1.94–2.08 (K)
3longer than broad, movable finger (with pedicel) 0.96–1.01
(L), 0.99–1.00 (K)3longer than hand. Fixed chelal finger with
8 trichobothria, movable chelal finger with 4 trichobothria
(Fig. 12), although sb absent from left chela of one male
(Fig. 11): eb,esb,ib and ist situated subbasally, est, isb and it
submedially, et subdistally, and est opposite it;band sb
situated subbasally, and st and tsituated submedially, with st
situated very close to t; patch of microsetae not present on
external margin of fixed chelal finger near et. Venom
apparatus present in both chelal fingers, venom ducts fairly
short, terminating in nodus ramosus slightly distal to et in
fixed finger (Figs. 11, 12). Chelal teeth pointed, slightly
retrorse, becoming rounded basally; fixed finger with 32 (L,
K) teeth; movable finger with 32 (L), 33 (K) teeth; accessory
teeth absent.
Carapace (Figs. 3, 6): 0.77–0.86 (L), 0.74 (K)3longer than
broad; anterior margin straight; with 2 pairs of rounded
corneate eyes, tapetum present; with 31 (L), 32 (K) setae,
arranged with 4 (L,K) near anterior margin and 4 (L,K) near
posterior margin; with 1 deep, broad median furrow.
Coxal region: manducatory process rounded with 1 small
sub-oral seta, and 9 (L), 12 (K) additional setae; median
maxillary lyrifissure large, rounded and situated submedially;
posterior maxillary lyrifissure rounded. Coxae I to IV
becoming progressively wider. Chaetotaxy of coxae I–IV: L,
6: 6: 6: 14; K,6:7:9:16.
Legs: femora I and II longer than patellae; junction between
femora and patellae III and IV very angulate; femora III and
IV much smaller than patellae III and IV; femur +patella of
leg IV 5.92 (L), 5.27 (K)3longer than broad (Fig. 10);
metatarsi and tarsi not fused; tarsus IV without tactile seta;
subterminal tarsal setae arcuate and acuminate; claws simple;
arolium much longer than claws, not divided.
Abdomen: tergites II–X and sternites IV–VIII of male and
female with medial suture line fully dividing each sclerite,
sternite IX partially divided. Tergal chaetotaxy: L, 4: 6: 10: 10:
11: 12: 11: 10: 10: 6 (arranged T4T): 7: 2; K,6:5:7:9:10:11:
11: 13: 9: 6 (arranged T4T): 8: 2; tergites I–X uniseriate.
Sternal chaetotaxy: L, 19: (0) 19 [3 +3] (0): (0) 6 (0): 7: 9: 7: 8:
8: 6: 3: 2; K, 14: (0) 8 (0): (0) 4 (0): 6: 7: 6: 8: 9: 6: 4: 2; sternites
IV–X uniseriate; Land Ksternite II and III with all setae
situated near posterior margin. Spiracles with helix. Anal
plates (tergite XII and sternite XII) situated between tergite XI
and sternite XI, and surrounded by desclerotized region of
Figures 1–5.—Larca cavicola (Muchmore), male from PARA-2004 Cave: 1. Body, dorsal; 2. Body, ventral; 3. Carapace, dorsal; 4. Left chela,
lateral; 5. Anal region, posterior.
208 THE JOURNAL OF ARACHNOLOGY
Figures 6–14.—Larca cavicola (Muchmore), specimens from PARA-3503 Cave: 6. Carapace, dorsal, male; 7. Chelicera, dorsal, male; 8.
Rallum, lateral, male; 9. Right pedipalp, dorsal, male; 10. Left leg IV, male; 11. Left chela, lateral, male; 12. Left chela, lateral, female; 13. Left
chela, tritonymph; 14. Left chela, deutonymph. Scale lines 50.1 mm (Figs. 7, 8), 0.2 mm (Figs. 11–14), 0.5 mm (Figs. 6, 9, 10).
HARVEY & WYNNE—TROGLOMORPHIC PSEUDOSCORPIONS FROM ARIZONA 209
tergite XI and sternite XI; sternite XI with ca. 18 (L), 24 (K)
small lyrifissures. Pleural membrane finely wrinkled-plicate;
without any setae.
Genitalia: male: very similar to that described for L. laceyi
Muchmore, 1981 by Muchmore (1981). Female with 1 pair of
lateral cribriform plates and 2 pairs of median cribriform
plates; spermathecae absent.
Dimensions: male (PARA-3503 Cave) followed by other
males (where applicable): Body length 2.40 (2.14–2.42).
Pedipalps: trochanter 0.371/0.186 (0.351–0.387/0.192–0.207),
femur 1.021/0.172 (0.923–0.976/0.187–0.206), patella 0.859/
0.192 (0.768–0.832/0.200–0.229), chela (with pedicel) 1.220/
0.231 (1.173–1.286/0.262–0.272), chela (without pedicel) 1.160
(1.106–1.216), hand length 0.576 (0.569–0.622), movable finger
length 0.582 (0.547–0.595). Chelicera 0.200/0.115, movable
finger length 0.130. Carapace 0.605/0.784 (0.621–0.656/0.763–
0.772); anterior eye diameter 0.059, posterior eye diameter
0.043. Leg I: femur 0.382/0.090, patella 0.249/0.092, tibia 0.350/
0.067, metatarsus 0.252/0.042, tarsus 0.218/0.042. Leg IV: femur
+patella 0.740/0.125, tibia 0.605/0.079, metatarsus 0.285/0.055,
tarsus 0.270/0.048.
Female (PARA-3503 Cave) followed by other female (where
applicable): Body length 2.85 (2.72). Pedipalps: trochanter
0.422/0.219 (0.408/0.215), femur 1.108/0.224 (0.978/0.214),
patella 0.992/0.252 (0.822/0.223), chela (with pedicel) 1.394/
0.307 (1.304/0.320), chela (without pedicel) 1.309 (1.232), hand
length 0.640 (0.621), movable finger length 0.643 (0.616).
Chelicera 0.240/0.131, movable finger length 0.150. Carapace
0.708/0.960); anterior eye diameter 0.049, posterior eye
diameter 0.048. Leg I: femur 0.410/0.103, patella 0.289/0.117,
tibia 0.382/0.075, metatarsus 0.261/0.059, tarsus 0.237/0.048.
Leg IV: femur +patella 0.828/0.157, tibia 0.660/0.095,
metatarsus 0.300/0.067, tarsus 0.282/0.058.
Tritonymph: Color: carapace, pedipalps and coxae red-
brown, abdomen pale red-brown and legs pale yellow-brown.
Chelicera: with 4 setae on hand and 1 on movable finger;
galea long and slender with 3 terminal rami.
Pedipalp: trochanter 1.97, femur 5.05, patella 3.90, chela
(with pedicel) 4.58, chela (without pedicel) 4.32, hand (without
pedicel) 2.17 3longer than broad, and movable finger 1.02 3
longer than hand (without pedicel). Fixed chelal finger with 7
trichobothria, movable chelal finger with 3 trichobothria
(Fig. 13): eb,esb,ist and ib situated basally; est and it
medially; et distally, isb absent; bsubbasally, st and t
submedially, sb absent. Fixed chelal finger with 26 teeth;
movable finger with 22 teeth.
Carapace: 0.85 3longer than broad; with 2 pairs of small
rounded corneate eyes; with 4 setae near anterior margin and 3
near posterior margin; with deep median furrow.
Legs: much as in adults.
Abdomen: tergal chaetotaxy: 4: 4: 6: 7: 8: 7: 8: 6: 6: 6
(arranged T4T): 7: 2. Sternal chaetotaxy: 2: (0) 7 (0): (0) 3 (0):
4: 4: 4: 5: 6: 4: 2: 2.
Dimensions (mm) (PARA-3503 Cave): Body length 1.75.
Pedipalps: trochanter 0.314/0.159, femur 0.768/0.152, patella
0.643/0.165, chela (with pedicel) 1.040/0.227, chela (without
pedicel) 0.981, hand length 0.493, movable finger length 0.501.
Carapace 0.544/0.640.
Deutonymph: Color: carapace, pedipalps and coxae pale
red-brown, abdomen and legs pale yellow-brown.
Chelicera: with 4 setae on hand and 1 on movable finger;
galea long and slender with 3 terminal rami.
Pedipalp: trochanter 2.11, femur 5.16, patella 3.50, chela
(with pedicel) 4.19, chela (without pedicel) 3.94, hand (without
pedicel) 2.02 3longer than broad, and movable finger 0.97 3
longer than hand (without pedicel). Fixed chelal finger with 6
trichobothria, movable chelal finger with 2 trichobothria
(Fig. 14): eb,ist and ib situated basally; est and it medially; et
distally; it subdistally, esb and isb absent; bsubbasally, t
submedially, sb and st absent. Fixed chelal finger with 24
teeth; movable finger with 21 teeth.
Carapace: 0.82 3longer than broad; with 2 pairs of small
rounded corneate eyes; with 4 setae near anterior margin and 4
near posterior margin; with deep median furrow.
Legs: much as in adults.
Abdomen: tergal chaetotaxy: 4: 4: 4: 6: 6: 6: 6: 6: 6: 6
(arranged T4T): 4: 2. Sternal chaetotaxy: 0: (0) 2 (0): (0) 2 (0):
3: 2: 4: 4: 4: 4: 4: 2.
Dimensions (mm) (PARA-3503 Cave): Body length 1.49.
Pedipalps: trochanter 0.278/0.132, femur 0.629/0.122, patella
0.514/0.147, chela (with pedicel) 0.850/0.203, chela (without
pedicel) 0.800, hand length 0.410, movable finger length 0.397.
Carapace 0.490/0.600.
Remarks.Larca cavicola was described from a single
female collected in Cave of the Domes, Grand Canyon
National Park, Coconino County, Arizona (Muchmore
1981). The new specimens were taken from two different
caves within the Parashant, PARA-3503 Cave and PARA-
2204 Cave, expanding the known range of this species some
160 km west of the type locality. Specimens from both cave
localities have shorter and slightly thinner pedipalpal segments
than the female holotype. In addition, the PARA-3503 Cave
specimens have slightly longer and thinner pedipalps than
those from PARA-2204 Cave. There do not appear to be any
other morphological features that would warrant the recog-
nition of more than one species amongst these specimens
which are all here attributed to L. cavicola. As noted by
Muchmore (1981), this species shows some obvious troglo-
morphic features consistent with an obligate subterranean
existence including long, slender pedipalps and legs, reduced
posterior eyes, and fewer setae on the carapace. Given the
findings of both Muchmore (1981) and the present study, we
consider this species to be troglobitic. A useful measure of
troglomorphic adaptation in larcid pseudoscorpions was
proposed by Gardini (1983), who found that the ratio
pedipalpal femur length/carapace length was lower in epigean
species of Larca than in cavernicolous species. This pattern
was also observed in two new Spanish species of Larca
(Zaragoza 2005). A similar condition is found in the species
formerly described in Archeolarca. The epigean L. rotunda has
a low ratio of 1.20 (male), 1.36 (female) (Hoff & Clawson
1952), whereas the cavernicolous species generally have higher
ratios: L. aalbui 1.57 (male), L. cavicola 1.44 (female), L.
guadalupensis 1.34 (female) and L. welbourni 1.47 (female)
(Muchmore 1981, 1984). The ratios of the new specimens of L.
cavicola recorded here [1.69 (male), 1.56 (female)] are higher
than the female holotype, but we ascribe this to individual
variation.
Two of the three post-embryonic nymphal stages (deuto-
nymph and tritonymph) are present in the samples, and they
210 THE JOURNAL OF ARACHNOLOGY
exhibit the same trichobothrial pattern as illustrated for L.
aalbui (under the name Archeolarca aalbui) by Harvey (1992).
Family Chernetidae Menge 1855
Subfamily Chernetinae Menge 1855
Hesperochernes Chamberlin 1924
Hesperochernes Chamberlin 1924:89–90.
Type species.Hesperochernes laurae Chamberlin 1924, by
original designation.
Remarks.—The genus Hesperochernes currently comprises
19 North American species, ranging as far south as the
Dominican Republic and Mexico (e.g., Ellingsen 1910;
Chamberlin 1924; Beier 1933, 1976) and as far north as
Canada (Hoff 1945), and a single Japanese species (Sato 1983).
Muchmore (1974) provided details on how to separate
Hesperochernes from the morphologically similar genera
Chernes Menge 1855 and Dinocheirus Chamberlin 1929, but
admitted that the composition of the genus was not fully
resolved due to uncertainties in the morphology of several
species. Hesperochernes is currently diagnosed by the follow-
ing combination of characters: rallum composed of 4 blades;
tarsus III and IV without conspicuous tactile seta; setae of
pedipalps and tergites not large and leaf-like; female
spermathecae with long paired ducts terminating in rounded
sacs; and cheliceral setae bs and sbs usually dentate or
denticulate. Of these characters, Muchmore (1974) was only
able to nominate the spermathecal morphology and the
denticulate bs and sbs as features that distinguish it from
Chernes. It appears, however, that some species currently
assigned to Hesperochernes have an acuminate bs, including
H. canadensis,H. holsingeri,H. molestus,H. montanus,H.
occidentalis and H. riograndensis (Chamberlin 1935; Hoff
1945; Hoff & Clawson 1952; Hoff 1956b; Hoff & Bolsterli
1956; Muchmore 1994). Moreover, the new species described
below clearly demonstrates the labile nature of this feature,
with the male having a strongly denticulate bs on both
chelicerae, but the two females having an acuminate bs.It
would seem that this feature should be used with considerable
caution, and that the nature of the spermathecae is the only
feature that can be reliably used to separate Hesperochernes
from Chernes.
Although Muchmore (1974) was able to confirm the generic
placement of several species from the U.S.A. and Canada
[H. laurae, H. mimulus Chamberlin 1952, H. mirabilis (Banks
1895), H. molestus Hoff 1956, H. occidentalis (Hoff and
Bolsterli 1956). H. riograndensis Hoff and Clawson 1952, H.
tamiae Beier 1930, and H. utahensis Hoff and Clawson 1952],
he was not able to ascertain whether others were correctly
placed [H. canadensis Hoff 1945, H. montanus Chamberlin
1935, H. pallipes (Banks 1893), H. paludis (Moles 1914), H.
thomomysi Hoff 1948, and H. unicolor (Banks 1908)]. The
same can be said of the Central American and Asian species
currently included in Hesperochernes,H. globosus (Ellingsen
1910), H. tumidus Beier 1933 and H. inusitatus Hoff 1946 from
Mexico, H. vespertilionis Beier 1976 from Dominican Repub-
lic, and H. shinjoensis Sato 1983 from Japan, as the morphology
of the spermathecae has not yet been ascertained (Ellingsen
1910; Beier 1933; Hoff 1946a; Beier 1976; Sato 1983).
Species of Hesperochernes are frequently collected in caves
or are associated with other animals. The cave-dwelling species
include three eyeless species that have long slender pedipalps
consistent with strong troglomorphisms, H. holsingeri,H.
mirabilis and H. occidentalis, as well as the new eyeless species
described below that has long legs but has robust pedipalps.
The species associated with rodents include H. mimulus,H.
molestus,H. riograndensis,H. tamiae,H. thomomysi and H.
utahensis (Beier 1930; Hoff 1945, 1946b; Chamberlin 1952;
Hoff & Clawson 1952; Hoff 1956b), while H. vespertilionis was
collected within a bat roost (Beier 1976). Hesperochernes
laurae and H. unicolor were found within both wasp’s and
ant’s nests (Banks 1908; Chamberlin 1924; Hoff 1947),
respectively, H. montanus was found in a bird’s nest
(Chamberlin 1935), and H. tumidus was collected ‘‘lying on
the ground in pods of Inga sp.’’ (translated from the original
German) (Beier 1933). The poorly described and most likely
misplaced H. paludis was taken from both rotten poplar tree
logs on the ground and live standing poplar trees (Moles
1914), and the only species recorded from outside of North
America, H. shinjoensis from northern Japan, was collected
from under tree bark (Sato 1983). The other species lack any
habitat data.
Hesperochernes bradybaughi sp. nov.
urn:lsid:zoobank.org:act:5419D319-EF22-4722-926F-1F8EC
080400B
Figs. 15–26
Material examined.Types. U.S.A.: Arizona: Mohave
County: holotype male, PARA-1001 Cave, Grand Canyon-
Parashant National Monument, ca. UTM 0264500 N, 4060700
E, Zone 12S, baited pitfall trap 3B, 20 August 2007, J.J. Wynne
(MNA); 1 female, same data as holotype except baited pitfall
trap 5A (MNA); 1 female, same data as holotype except
opportunistic, mid cave, 13 August 2005 (NMA).
Etymology.—This species is named for Jeff Bradybaugh,
former superintendent of Grand Canyon-Parashant National
Monument and an advocate for cave research, conservation
and management both on Parashant and within the National
Park Service.
Diagnosis.Hesperochernes bradybaughi most closely re-
sembles three other species of the genus that are also
completely eyeless and have long slender legs [e.g. femur +
patella IV 5.19 (male), 5.37–5.56 (female) 3longer than
broad], H. mirabilis,H. holsingeri and H. riograndensis.
Hesperochernes bradybaughi lacks the slender pedipalps
characteristic of H. mirabilis and H. holsingeri, and the male
chela of H. bradybaughi is markedly swollen, especially on the
dorsal face (Fig. 21), unlike the male of H. riograndensis which
is not swollen. It is also substantially larger than H.
riograndensis, e.g., chela (without pedicel) of H. riograndensis
is 0.956 (male), 0.970 (female) mm, whereas H. bradybaughi is
1.434 (male), 1.502–1.510 (female) mm.
Description.—Adults: Color: pedipalps and carapace dark
red-brown, legs light red-brown, tergites yellow-brown,
sternites pale yellow-brown.
Chelicera: with 5 setae on hand and 1 subdistal seta on
movable finger (Fig. 23); setae ls and is acuminate, es and bs
dentate, sbs denticulate in female, acuminate in male; with 2
dorsal lyrifissures and 1 ventral lyrifissure; galea of Land K
with 6 rami; rallum of 4 blades, the 2 distal blades with several
HARVEY & WYNNE—TROGLOMORPHIC PSEUDOSCORPIONS FROM ARIZONA 211
serrations on leading edge, other blades smooth; serrula
exterior with 18 (L), 17 (K) blades; lamina exterior present.
Pedipalp (Fig. 24): surfaces of trochanter, femur, patella
and chelal hand coarsely granulate, chela fingers mostly
smooth; patella with 5 small sub-basal lyrifissures; trochanter
1.84 (L), 1.86–1.88 (K), femur 3.17 (L), 2.95–3.09 (K), patella
2.62 (L), 2.54–2.66 (K), chela (with pedicel) 3.07 (L), 3.23–3.34
(K), chela (without pedicel) 2.83 (L), 2.98–3.09 (K), hand 1.49
(L), 1.34–1.64 (K)3longer than broad, movable finger 0.93
(L), 0.86–0.96 (K)3longer than hand. Fixed chelal finger with
8 trichobothria, movable chelal finger with 4 trichobothria
(Figs. 21, 22): eb and esb situated basally, ib and ist
subbasally, est and isb submedially, et and it subdistally, isb
situated midway between ist and it, and et slightly distal to it;t
situated subdistally, st situated closer to tthan to sb. Venom
apparatus only present in movable chelal finger, venom ducts
long, terminating in nodus ramosus distal to st (Figs. 21, 22).
Fixed finger with 2 large sensillae on retrolateral face, and 2 on
prolateral face; movable chelal finger with sensilla slightly
proximal to sb in male and slightly distal to sb in female, with
2 receptors. Chela of male without mound. Chelal teeth
pointed and slightly retrorse, basal teeth more rounded; fixed
finger with 44 (L), 48 (K) teeth, plus 11 (L), 9 (K) retrolateral
and 10 (L), 7 (K) prolateral accessory teeth; movable finger
with 46 (L), 50 (K) teeth, plus 9 (L,K) retrolateral and 6 (L), 4 (K)
prolateral accessory teeth.
Carapace (Fig. 17): coarsely granulate, 1.15 (L), 0.98–1.10
(K)3longer than broad; without eyes or eyespots; with 100
(L), 83 (K) setae, arranged with 61 (L), 42 (K) (including 6 near
anterior margin) in anterior zone, 25 (L), 34 (K) in median
zone, and 14 (L), 17 (K) in posterior zone; with 2 deep furrows,
posterior furrow situated slightly closer to posterior carapace
margin than to anterior furrow.
Coxal region: maxillae granulate; manducatory process
somewhat acute, with 2 apical acuminate setae, 1 small sub-
oral seta and 37 (L), 32 (K) additional setae; median maxillary
lyrifissure rounded and situated submedially; posterior max-
illary lyrifissure rounded. Leg coxae smooth; chaetotaxy of
coxae I–IV: L, 18: 19: 23: ca. 60; K, 18: 21: 25: ca. 65.
Legs: very slender; junction between femora and patellae I
and II strongly oblique to long axis; junction between femora
and patellae III and IV very angulate; femora III and IV much
smaller than patellae III and IV; femur +patella of leg IV 5.19
(L), 5.37–5.56 (K)3longer than broad; all tarsi with slit
sensillum on raised mound; male leg I not modified; tarsi III
and IV without tactile seta, but with paired subdistal setae;
subterminal tarsal setae arcuate and acute; claws simple;
arolium about same length as claws, not divided.
Abdomen: tergites I–X and sternites IV–X of male and
female with median suture line fully dividing each segment.
Tergal chaetotaxy: L, 11: 12: 11: 18: 19: 18: 20: 18: 17: 18: 13: 2;
K, 12: 13: 13: 17: 17: 18: 19: 19: 21: 16: 14: 2; uniseriate, except
Figures 15–20.—Hesperochernes bradybaughi, sp. nov.: 15. Body, dorsal, male holotype; 16. Body, ventral, male holotype; 17. Carapace,
dorsal, male holotype; 18. Body, dorsal, female paratype; 19. Body, ventral, female paratype; 20. Left chela, lateral, male holotype.
212 THE JOURNAL OF ARACHNOLOGY
for medial and lateral discal seta on tergites IV–IX; setae
thickened and strongly dentate. Sternal chaetotaxy: L, 30: (3) 22
[2 +2] (3): (1) 8 (1): 19: 21: 21: 20: 20: 16: 8 (arranged T6T): 2; K,
ca. 40: (3) 10 (3): (1) 5 (1): 14: 21: 20: 20: 18: 16: 11 (arranged
T9T): 2; uniseriate, except for lateral discal seta on sternites
VII–X; setae of anterior sternites acicular, becoming progres-
sively more denticulate on posterior sternites. Spiracles with
helix. Anal plates (tergite XII and sternite XII) situated between
tergite XI and sternite XI, anal setae not denticulate. Pleural
membrane wrinkled and somewhat stellate; without any setae.
Genitalia: male of the chernetid type. Female (Fig. 26): with
a pair of long thin-walled spermathecae terminating in
rounded sacs.
Dimensions: Male holotype: Body length 3.11. Pedipalps:
trochanter 0.518/0.282, femur 974/0.307, patella 0.824/0.314,
chela (with pedicel) 1.552/0.506, chela (without pedicel) 1.434,
hand length 0.756, movable finger length 0.704. Chelicera
0.322/0.165, movable finger length 0.252. Carapace 0.956/
0.830. Leg I: femur 0.268/0.161, patella 0.495/0.136, tibia
0.503/0.102, tarsus 0.495/0.079. Leg IV: femur +patella 0.883/
0.170, tibia 0.778/0.107, tarsus 0.557/0.085.
Female (paratype lodged in MNA) followed by other
female (where applicable): Body length 2.82 (4.21). Pedipalps:
trochanter 0.552/0.294 (0.566/0.304), femur 1.034/0.335 (1.042/
0.353), patella 0.904/0.340 (0.942/0.371), chela (with pedicel)
1.624/0.486 (1.635/0.506), chela (without pedicel) 1.502
Figures 21–26.—Hesperochernes bradybaughi, sp. nov.: 21. Left chela, lateral, male holotype; 22. Left chela, lateral, female paratype; 23.
Chelicera, dorsal, female paratype; 24. Right pedipalp, dorsal, male holotype; 25. Left leg IV, male holotype; 26. Spermathecae, female paratype.
Scale lines 50.1 mm (Fig. 23), 0.2 mm (Fig. 26), 0.5 mm (Figs. 21, 22, 24, 25).
HARVEY & WYNNE—TROGLOMORPHIC PSEUDOSCORPIONS FROM ARIZONA 213
(1.510), hand length 0.797 (0.698), movable finger length 0.768
(0.816). Chelicera 0.327/0.152, movable finger length 0.244.
Carapace 1.040/0.944 (1.000/1.021). Leg I: femur 0.300/0.182,
patella 0.540/0.146, tibia 0.56/0.108, tarsus 0.520/0.079. Leg
IV: femur +patella 1.010/0.188 (1.000/0.180), tibia 0.830/
0.121, tarsus 0.580/0.084.
Remarks.—As stated in the diagnosis, H. bradybaughi
appears to be most similar to H. riograndensis but differs in
being substantially larger and with a markedly swollen male
chela, especially on the dorsal face. The only known location
of H. riograndensis is located 670 km ESE of Parashant, and
the microhabitat of both species differs with H. bradybaughi
being found in a cave and H. riograndensis collected from the
nest of a kangaroo rat (Heteromyidae: Dipodomys) (Hoff &
Clawson 1952). Given the lack of eyes and eyespots, we
consider H. bradybaughi to be a troglobite.
Tuberochernes Muchmore
Tuberochernes Muchmore 1997:206–207.
Type species.Tuberochernes aalbui Muchmore 1997, by
original designation.
Diagnosis.Tuberochernes differs from all other chernetid
genera by the combined presence of a distinct medium-sized
mound on the prolateral face of the pedipalpal chela of males,
and four blades in the cheliceral rallum.
Remarks.—The genus Tuberochernes was described by
Muchmore (1997) for two species of cave-dwelling pseudo-
scorpions from southwestern U.S.A., T. aalbui and T. ubicki,
but the discovery of a third species, also from a cave in
southwestern U.S.A., does not necessitate an alteration of the
original description apart from the nature of the tactile seta of
leg IV. Muchmore (1997) observed that the tactile seta of leg
IV was ‘‘short, distally located’’ and ‘‘variably acuminate or
finely denticulate’’. Close examination of the posterior tarsi of
the new species described below does not reveal a tactile seta
of this nature, and we suggest this feature appears to be
variable within the genus.
The most obvious feature that distinguishes Tuberochernes
is the presence of a medium-sized mound on the prolateral
margin of the chelal hand in males (Muchmore 1997). In this
respect, it resembles several other chernetid genera, including
males of Mirochernes Beier 1930 and Bituberochernes Much-
more 1974, and both males and females of Interchernes
Muchmore 1980 and Petterchernes Heurtault 1986, which
were distinguished from Tuberochernes by Muchmore (1997).
Bituberochernes further differs from Tuberochernes by a
mound being also present on the pedipalpal patella. The
function of the mound has not been ascertained, but the
mound of T. cohni has 5 small pores, which may be responsible
for discharging fluids, possibly during sexual interactions with
females.
Tuberochernes cohni sp. nov.
urn:lsid:zoobank.org:act:12896B35-DD1C-4E0B-B66F-F9B
30170D476
Figs. 27–37
Material examined.—Type: U.S.A.: Arizona: Mohave
County: holotype male, PARA-1001 Cave, Grand Canyon-
Parashant National Monument, ca. UTM 0264500 N,
4060700 E, Zone 12S, the deeper extent of the twilight zone
(near the dark zone), opportunistic collecting, 13 August 2005,
J.J. Wynne (MNA).
Etymology.—This species is named for the late Dr.
Theodore ‘‘Ted’’ Cohn. Cohn was an Orthopterist and the
leading authority who identified the new genus of rhaphido-
phorid cricket known from PARA-1001 Cave. Dr. Cohn
passed away in November 2013 at age 82. He was a passionate
educator and entomologist.
Diagnosis.Tuberochernes cohni differs from the other two
species of the genus, T. aalbui and T. ubicki, by the more
anteriorly positioned mound on the pedipalpal chela.
Description.—Adult male: Color: pedipalps and carapace
dark red-brown, legs light red-brown, tergites yellow-brown,
sternites pale yellow-brown.
Chelicera: with 6 setae on hand and 1 subdistal seta on
movable finger (Fig. 32); setae es,sbs and bs dentate, ls and is
acuminate; with 2 dorsal lyrifissures and 1 ventral lyrifissure;
galea broken; rallum of 4 blades, the most distal blade with
several serrations on leading edge, other blades smooth;
serrula exterior with 17 blades; lamina exterior present.
Pedipalp (Fig. 33): surfaces of trochanter, femur, patella
and chelal hand coarsely granulate, chela fingers mostly
smooth; patella with 5 small sub-basal lyrifissures; trochanter
1.73, femur 2.83, patella 2.88, chela (with pedicel) 3.39, chela
(without pedicel) 3.11, hand 1.40 3longer than broad,
movable finger 1.23 3longer than hand. Fixed chelal finger
with 8 trichobothria, movable chelal finger with 4 trichobo-
thria (Fig. 31): eb and esb situated basally, ib and ist
subbasally, est and isb submedially, et and it subdistally, isb
situated midway between ist and it, and et slightly distal to it;t
situated subdistally, st situated much closer to tthan to sb.
Venom apparatus only present in movable chelal finger,
venom ducts long, terminating in nodus ramosus midway at
level of st (Fig. 31). Fixed finger with 3 sensillae on retrolateral
face, and 1 on prolateral face; movable chelal finger with
sensilla slightly distal to sb, with 2 receptors. Chela with
prominent, medium-sized mound on prolateral face (Figs. 30,
34), with 5 small pores. Chelal teeth pointed and slightly
retrorse, basal teeth more rounded; fixed finger with 37 teeth,
plus 7 retrolateral and 3 prolateral accessory teeth; movable
finger with 42 teeth, plus 4 retrolateral and 0 prolateral
accessory teeth.
Carapace (Fig. 29): coarsely granulate, 1.19 3longer than
broad; without eyes or eyespots; with 96 setae, arranged with
54 (including 6 near anterior margin) in anterior zone, 28 in
median zone, and 14 in posterior zone; with 2 deep furrows,
posterior furrow situated closer to posterior carapace margin
than to anterior furrow.
Coxal region: maxillae granulate; manducatory process
somewhat acute, with 2 apical acuminate setae, 1 small sub-
oral seta and 25 additional setae; median maxillary lyrifissure
rounded and situated submedially; posterior maxillary lyr-
ifissure rounded. Leg coxae smooth; chaetotaxy of coxae I–IV:
13: 12: 14: 34.
Legs (Figs. 35–37): junction between femora and patellae I
and II strongly oblique to long axis; junction between femora
and patellae III and IV very angulate; femora III and IV much
smaller than patellae III and IV; femur +patella of leg IV 4.03
3longer than broad; all tarsi with slit sensillum on raised
214 THE JOURNAL OF ARACHNOLOGY
mound; leg I modified with tibia thickened, tarsus slightly
curved and ventral margins of patella and tibia with coarse
granulation; tarsi III and IV without tactile seta, but with
paired subdistal setae; subterminal tarsal setae arcuate and
acute; claws simple; arolium about same length as claws, not
divided.
Abdomen: tergites II–X and sternites V–X of with median
suture line fully dividing each segment. Tergal chaetotaxy: 15:
20: 20: 20: 22: 22: 21: 21: 22: 17: 10: 2; uniseriate, except for
medial and lateral discal seta on tergites IV–IX; setae
thickened and strongly dentate. Sternal chaetotaxy: 51: (0) 8
[2 +2] (0): (1) 8 (1): 12: 16: 17: 18: 17: 14: 8 (arranged T6T): 2;
uniseriate, except for lateral discal seta on sternites IV–XI;
setae of anterior sternites acicular, becoming progressively
more denticulate on posterior sternites. Spiracles with helix.
Anal plates (tergite XII and sternite XII) situated between
tergite XI and sternite XI, anal setae denticulate. Pleural
membrane longitudinally striate; without any setae.
Genitalia: of the chernetid type.
Dimensions: male holotype: Body length 3.38. Pedipalps:
trochanter 0.576/0.332, femur 0.944/0.334, patella 0.910/0.316,
chela (with pedicel) 1.390/0.410, chela (without pedicel) 1.276,
hand length 0.573, movable finger length 0.704. Chelicera
0.333/0.134, movable finger length 0.240. Carapace 1.009/
0.848. Leg I: femur 0.305/0.249, patella 0.560/0.253, tibia
0.621/0.174, tarsus 0.442/0.089. Leg IV: femur +patella 0.859/
0.213, tibia 0.692/0.134, tarsus 0.533/0.954.
Remarks.Tuberochernes cohni possesses some very slight
modifications consistent with troglomorphic adaptations of
which the most prominent is the complete lack of eyes
(Fig. 29) and the slightly elongated leg segments. Thus, this
animal is considered a troglobite. It appears to bear a closer
resemblance to T. ubicki from a cave in the Santa Rita
Mountains, Arizona (610 km), than to T. aalbui from a cave in
the Inyo National Forest, California (415 km), due to the
similarly expanded tibia I in males of the two Arizona species.
DISCUSSION
Our review of the pseudoscorpions detected within the caves
of Grand Canyon-Parashant National Monument has re-
vealed a modest fauna of three species: Larca cavicola (family
Larcidae), Hesperochernes bradybaughi and Tuberochernes
cohni (both in the family Chernetidae). All show modifications
consistent with obligate existence in cave environments, but
none show the classic signs of extreme troglomorphism found
in many cave-adapted pseudoscorpions (e.g. Heurtault 1994;
Harvey et al. 2000). Both species of Chernetidae lack eyes and
have long slender legs, which appear to be troglomorphic
modifications due to their subterranean existence, although
their pedipalps do not appear to be modified compared to
epigean species of the genus. Other subterranean species of
Hesperochernes with thin legs and no eyespots—H. holsingeri
from Indiana, H. mirabilis from Alabama, Georgia, Indiana,
Kentucky, Ohio, Tennessee and Virginia, and H. occidentalis
from Arkansas, Missouri, Oklahoma and Texas—appear to be
more highly modified as they have elongate pedipalps. Both
new species described from the Parashant may represent short-
range endemic species as defined by Harvey (2002) and Harvey
Figures 27–30.—Tuberochernes cohni, sp. nov., male holotype: 27. Body, dorsal; 28. Body, ventral; 29. Carapace, dorsal; 30. Right
chela, dorsal.
HARVEY & WYNNE—TROGLOMORPHIC PSEUDOSCORPIONS FROM ARIZONA 215
Figures 31–37.—Tuberochernes cohni, sp. nov., male holotype: 31. Left chela, lateral; 32. Chelicera, dorsal; 33. Right pedipalp, dorsal; 34. Left
chela, detail of mound, ventral; 35. Left tarsus IV; 36. Left I; 37. Left leg IV. Scale lines 50.2 mm (Figs. 32, 34, 35), 0.5 mm (Figs. 31, 33, 36, 37).
216 THE JOURNAL OF ARACHNOLOGY
et al. (2011) due to their highly restricted distributions.
Although the junior author and colleagues sampled all known
caves on Parashant, they detected these new species in only
one cave (PARA-1001 Cave).
Larca cavicola appears to be less cave-adapted than the
others, as it retains eyes. However, the pedipalps are
noticeably thinner than epigean species of the genus,
suggesting moderate morphological modifications to the cave
environment. Larca cavicola was found in PARA-3503 and
PARA-2204 Caves and has been found in Cave of the Domes,
a small cave situated within Grand Canyon National Park,
Coconino County (Muchmore 1981). Although this cave is
also located in the Grand Canyon region, it lies on the south
side of the Colorado River some 160 km from the Parashant
caves, and we suggest these populations are genetically
isolated from each other.
The only known locality of Hesperochernes bradybaughi and
Tuberochernes cohni is PARA-1001. This is the second most
biologically diverse cave, and the most biologically significant
cave on the monument. It supports the largest known cricket
roost in Arizona, which represents an undescribed genus of
rhaphidophorid cave cricket, cf Ceuthophilus n. gen. n. sp.,
Cohn & Swanson, unpublished data; (Wynne & Voyles 2014).
Its population contributes significantly to the nutrient loading
via cricket guano, cricket eggs and nymphs, as well as deceased
individuals at various life stages. In other regions, the
ecological importance of crickets on cave ecosystems is well
documented (e.g., Barr 1967; Howarth 1983; Taylor 2003;
Culver 2005; Poulson 2005). Given the size of the roost, we
suggest that cf Ceuthophilus n. gen. n. sp. represents a keystone
species with the presence of this animal supporting at least
four cave-adapted species including a short-range endemic and
troglomorphic leiodid beetle, Ptomaphagus parashant (Peck &
Wynne 2013), an undescribed species of troglomorphic
centipede (family Anopsobiidae; Wynne, unpublished data),
and the two pseudoscorpion species described here. To date,
P. parashant, the anopsobiid centipede, and the two new
pseudoscorpion species have been detected only in PARA-
1001 Cave. Two other caves on the monument, with similar
deep zone like conditions, were sampled using the same
systematic sampling design are within a 9.7 km radius of
PARA-1001; neither of these new pseudoscorpions species
were detected at these caves.
Management Implications.—We recommend the same man-
agement strategies proposed by Peck & Wynne (2013) be
maintained for PARA-1001 Cave. This cave should not be
gated given its south-facing entrance and entrance structure,
and it should remain closed to recreational use. PARA-1001 is
considered the second most biologically diverse cave on the
monument and supports the greatest diversity of troglo-
morphic arthropod species. Presently, all of these animals
(including the two new pseudoscorpion species described here)
are known to occur only within PARA-1001 Cave. Maintain-
ing the management strategies suggested by Peck & Wynne
(2013) should aid in the long-term persistence of these
presumed short-range endemic arthropods.
ACKNOWLEDGMENTS
Special thanks to Jennifer Fox, Eathan McIntyre, Ray
Klein and Rosie Pepito of Grand Canyon-Parashant National
Monument, Danielle Nelson and Matt Johnson with the
Colorado Plateau Research Station, and Neil Cobb of the
Colorado Plateau Museum of Arthropod Biodiversity for
administrative and logistical support. Tama and John Cassidy,
Michael Gowan, John Kalman, Ty Spatta and Kyle Voyles
assisted with fieldwork. The San Bernardino Cave Search and
Rescue Team, Jon Jasper and Kyle Voyles, remained on
emergency stand-by during field operations. Dave Decker and
Kyle Voyles provided descriptions regarding the geological
and structural characteristics of the study caves. Dale Pate and
two anonymous reviewers provided suggestions leading to the
improvement of this manuscript. The Explorers Club recog-
nized two of these research trips as flag expeditions. Fieldwork
was funded through a Colorado Plateau CESU cooperative
agreement between the National Park Service and Northern
Arizona University.
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HARVEY & WYNNE—TROGLOMORPHIC PSEUDOSCORPIONS FROM ARIZONA 219
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... Balgachernes differs from most of these genera in the placement of trichobothrium est which is much closer to esb than to et in Balgachernes ( Figure 14), but approximately midway between them, or only slightly closer to esb, in Atherochernes (Beier 1954), Ceriochernes (Beier 1937), Cocinachernes (Hentschel and Muchmore 1989), Gigantochernes (Beier 1932;Vitali-di Castri 1972), Hesperochernes (e.g. Chamberlin 1924;Harvey and Wynne 2014;Muchmore 1974), Illinichernes (Benedict and Malcolm 1982;Hoff 1949), Marachernes bellus Harvey, 1992 (but not M. perup Harvey, 1992or M. simulans Harvey, 1992 (Harvey 1992a), Maxchernes birabeni Feio, 1960and M. plaumanni Beier, 1974(but not M. iporangae Mahnert and Andrade, 1998 (Beier 1974;Feio 1960;Mahnert and Andrade 1998), Mirochernes (Hoff 1949), Neochelanops (Beier 1964a;Mahnert 1984), Nesochernes (Beier 1932), Paraustrochernes (Beier 1966a), Satrapanus (Harvey and Volschenk 2007), Semeiochernes (Beier 1932) and Troglochernes (Harvey and Volschenk 2007), or est closer to et than to esb in Eumecochernes (Beier 1932). It differs from Marachernes by the lack of a dentate prolateral mound on the male movable chelal finger (Harvey 1992a), and from Maxchernes by having isb situated closer to ist than to it, whereas isb is closer to it in Maxchernes (Beier 1974;Feio 1960;Mahnert and Andrade 1998). ...
... While the Guangxi pseudoscorpions currently known only from their type locality may also occur in other regional caves, it is improbable they are wide-ranging species. In general, troglomorphic species are well documented as being either single endemics or having small distributional ranges (Christman et al. 2005;Deharveng et al. 2008;Latella & Chen 2008;Tian 2011;Borges et al. 2012;Harvey & Wynne 2014). Wynne (unpublished data) found that of the 99 known, cave-adapted arthropods for Guangxi, over half (or 58 species) were identified from one cave. ...
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