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Cave-dwelling arthropods and vertebrates of north rim Grand Canyon, with notes on ecology and management


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Prior to this study, there was no information on arthropods, bats, and other vertebrates of caves in northwesternmost Arizona. Based on invertebrate and vertebrate inventory work conducted during 2005 and 2006, we provide future directions for conservation and management for caves on Grand Canyon–Parashant National Monument, northwestern Arizona. Baseline investigations to find and identify arthropods, bats, and other vertebrates were conducted at 7 of the largest known caves on the monument. We identified 52 morphospecies including 44 arthropods, 4 bats, and 4 other vertebrates. Of the cave-dwelling arthropods, we found 10 eisodophiles, 6 troglophiles, 8 questionable troglophiles, 7 trogloxenes, 8 accidentals, 3 taxa of unknown cave affiliations, and 2 mammalian parasites. We made several contributions to the entomological record including 2 new genera, 6 new species, 3 possible new species, one range extension, and one possible range extension. Also, we identified 5 bat roosts—1 hibernaculum, 2 night roosts, and 3 summer roosts of unconfirmed use. Observed arthropod richness per cave ranged from 1 to 14 morphospecies, and observed bat and other vertebrate (combined) richness was 1–3 morphospecies. We did not detect any cave-adapted arthropods during this investigation. For the caves sampled, we are uncertain whether the lack of cave-adapted taxa is due to (a) low nutrient input and high cryptoaridity associated with many southwestern cave systems or (b) lack of intensive sampling. Despite the lack of cave-adapted species, 5 of the 7 caves inventoried are considered of high management concern. Additional research at these caves will be required to obtain the data necessary to best manage and protect these systems.
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Information related to the biospeleology
of northern Arizona is limited. Prior to this
study, no investigations on cave use by ar -
thropods, bats, and other vertebrates in north-
western Arizona had been undertaken. Based
on our literature review, most of the work on
the southern Colorado Plateau (within north-
ern Arizona) has been baseline in nature, and
these studies were largely focused on Wupatki
National Monument and Grand Canyon Na -
tional Park. Given limited information and the
desire to provide a regional summary of cave
biology, we present a brief overview of the
taxa reported during previous work.
At Wupatki National Monument, Welbourn
(1976) conducted 1–2 site visits at 5 earth cracks
(volcano-tectonic fissures) and identified 19 ar -
thropods, including 5 troglophiles, 13 troglox-
enes, and 5 accidentals; no troglobites were
identified during his work (see below for defi-
nitions on cave specificity functional groups).
From Welbourn’s inventory (1976), Muchmore
Western North American Naturalist 74(1), © 2014, pp. 1–17
J. Judson Wynne1and Kyle D. Voyles2
ABSTRACT.—Prior to this study, there was no information on arthropods, bats, and other vertebrates of caves in north-
westernmost Arizona. Based on invertebrate and vertebrate inventory work conducted during 2005 and 2006, we pro-
vide future directions for conservation and management for caves on Grand Canyon–Parashant National Monument,
northwestern Arizona. Baseline investigations to find and identify arthropods, bats, and other vertebrates were con-
ducted at 7 of the largest known caves on the monument. We identified 52 morphospecies including 44 arthropods, 4
bats, and 4 other vertebrates. Of the cave-dwelling arthropods, we found 10 eisodophiles, 6 troglophiles, 8 questionable
troglophiles, 7 trogloxenes, 8 accidentals, 3 taxa of unknown cave affiliations, and 2 mammalian parasites. We made sev-
eral contributions to the entomological record including 7 new species (with 2 new genera), 3 possible new species, one range
extension, and one possible range extension. Also, we identified 5 bat roosts—1 hibernaculum, 2 night roosts, and 3
summer roosts of unconfirmed use. Observed arthropod richness per cave ranged from 1 to 14 morphospecies, and
observed bat and other vertebrate (combined) richness was 1–3 morphospecies. We did not detect any cave-adapted
arthropods during this investigation. For the caves sampled, we are uncertain whether the lack of cave-adapted taxa is
due to (a) low nutrient input and high cryptoaridity associated with many southwestern cave systems or (b) lack of inten-
sive sampling. Despite the lack of cave-adapted species, 5 of the 7 caves inventoried are considered of high manage-
ment concern. Additional research at these caves will be required to obtain the data necessary to best manage and pro-
tect these systems.
RESUMEN.—Anterior a este estudio, no había información sobre los artrópodos, murciélagos y otra fauna en las cav-
ernas al noroeste de Arizona. Basados en el inventario de invertebrados y vertebrados realizados durante 2005 y 2006,
nosotros proveemos las futuras direcciones para la investigación y gestión de cavernas del Grand Canyon–Parashant
National Monument, noroeste de Arizona. Investigaciones iniciales fueron realizados en artrópodos, murciélagos y fauna
en 7 de las más grandes cavernas conocidas. Identificamos 52 morfoespecies incluyendo 44 artrópodos, 4 murciélagos
y 4 fauna silvestre. De los artrópodos, hubo 10 eisodofilos, 6 troglofilos, 8 pseudo-troglofilos, 7 trogloxenos, 8 acciden-
tales, 3 desconocidas y 2 ectoparásitos. Realizamos varias contribuciones al registro entomológico incluyendo 7 nuevas
especies (con 2 nuevos géneros), 3 posibles nuevas especies, una expansión distribucional y una posible expansión dis-
tribucional. También, identificamos 5 refugios de murciélagos: 1 hibernáculo, 2 dormideros nocturnos y 3 dormideros
estivales de uso indeterminado. La riqueza observada de artrópodos oscilo entre 1–14 morfoespecies y la riqueza combi-
nada para murciélagos y fauna vario entre 1–3 morfoespecies. Durante este trabajo, no fueron encontrados artrópodos
troglomorficos. En las cavernas muestreadas, se desconoce si la falta de taxones adaptados a las cavernas es debido a
(i) el bajo aporte de nutrientes y la alta cripto-aridez asociada generalmente con los sistemas de cavernas del suroeste, o
(ii) el insuficiente muestreo. A pesar de ello, 5 de las 7 cavernas inventariadas son consideradas como de alto interés
de gestión. Investigaciones adicionales en estas cavernas serán necesarios para obtener los datos requeridos para una
mejor gestión y protección de estos sistemas.
1Department of Biological Sciences, Colorado Plateau Biodiversity Center, Colorado Plateau Research Station and Landscape Conservation Initiative,
Northern Arizona University, Box 5614, Building 56, Suite 150, Flagstaff, AZ 86011. URL:
2Saint George Field Office, Bureau of Land Management, 345 E. Riverside Dr., St. George, UT 84790.
(1981) described an endemic pseudoscorpion
(Archeolarca welbourni) from 2 earth cracks.
Bat research has been conducted on a nearly
decadal scale at Wupatki. Gustafson (1964) col -
lected one Townsend’s big-eared bat (Cory -
norhinus townsendii) during the summer near
the bottom of an earth crack. Welbourn (1976)
identified C. townsendii at 4 earth cracks (one
of the earth cracks was the same visited by
Gustafson [1964]). Day roost activity was docu -
mented at all features in September, and 2
earth cracks were identified as hibernacula.
Bain (1986) documented winter use of 7 earth
cracks (confirming continued use of the one
previously visited by Gustafson and 4 visited
by Welbourn [1976]) by C. townsendii. Con-
cerning other vertebrate species, Welbourn
(1976) identified the remains of 2 accidental
morphospecies in Wupatki earth cracks: Ro -
dentia (from 2 earth cracks) and Lagomorpha
(from one earth crack).
Wynne et al. (2007) compiled a review of
cave-dwelling arthropod research in Grand
Canyon National Park (GRCA) and synthe-
sized information from 15 caves representing
9 studies (conducted between 1975 and 2001).
At least 47 cave-dwelling arthropods, includ-
ing 3 troglobites, one stygobite (aquatic cave-
adapted animals), 7 trogloxenes, 16 troglophiles,
and 16 unidentified cavernicoles (presumed
troglophiles) were identified (Wynne et al.
2007). Additionally, bats known to use GRCA
caves include the Brazilian free-tailed bat (Ta -
darida brasiliensis) from 2 caves (Pape 1998,
Hill et al. 2001) and C. townsendii from 2
caves (Welbourn 1978). Other cave-dwelling
vertebrates known to occur in GRCA include
Neotoma sp. from 4 caves (Peck 1980, Boden-
hamer 1989, Drost and Blinn 1998, Hill et al.
2001), brush mouse (Peromyscus boylii) from
one cave (Drost and Blinn 1998), American
porcupine (Erethizon dorsatum) from one cave
(Hill et al. 2000) and ringtail (Bassariscus astu-
tus) from 4 caves (Peck 1980, Bodenhamer
1989, Pape 1998, Hill et al. 2000, 2001).
Peck (1982) conducted invertebrate surveys
and reported on the largely depauperate fauna
(n= 2 morphospecies) occurring in 4 lava tube
caves at Sunset Crater National Monument and
in the greater Flagstaff region.
Although much of the biospeoleological re -
search over the past several decades has been
baseline in nature, these studies combined with
our present work begin to produce a regional
picture of the natural history of caves on the
southern Colorado Plateau.
Here we report the first regional all-taxa
biological inventory of known caves in Grand
Canyon–Parashant National Monument, Ari-
zona. Our objectives were to conduct baseline
inventories by (1) sampling cave-dwelling ar -
thropods; (2) identifying bat use of caves either
directly or by examining previous evidence of
use; (3) documenting all other vertebrate ac -
tivity observed within each cave; and (4) iden-
tifying caves of high management concern for
further study.
Cave-Dependency Functional Groups
We divided Grand Canyon–Parashant cave-
dwelling taxa into 7 cave-dwelling organism
groups and one special case category. The fol-
lowing definitions were taken from Barr (1968),
Howarth (1983), and Wynne (2013): (1) trog -
lobites, obligate cave dwellers who can only
com plete their life cycle within the hypogean
environment; (2) troglophiles, spe cies that oc -
cur facultatively within caves and complete
their life cycles there, but also exist in simi -
lar dark and humid epigean microhabitats; (3)
trogloxenes, species that frequently use caves
for shelter but forage in the epigean realm; (4)
accidentals, morphospecies that oc cur in caves
but cannot survive within the hypogean envi-
ronment; (5) eisodophiles, spe cies that facul -
tatively use cave entrances and twilight zones
and may complete their life cycles there, but
also exist in similar par tially sheltered epi -
gean environments; (6) eisodoxenes, species
that fre quently use cave en trances and twi-
light zones for shelter but forage on the sur-
face; (7) unknown, species for which informa-
tion is lacking to place them within one of the
6 aforementioned groups; and, (8) parasites, a
special-case group, which includes parasitic ar -
thropods detected in caves due to the presence
of their host (e.g., bats or other mammals).
Study Site
Located in northwestern Arizona, Grand
Canyon–Parashant National Monument (here-
after shortened to Parashant) is jointly man-
aged by the National Park Service and the
Bureau of Land Management. Encompassing
1.1 million acres, Parashant is characterized
by rugged terrain containing deeply incised
canyons, mesas, and mountains. Two geologi-
cal provinces converge here: the Basin and
Range and the Colorado Plateau. Vegetation
zones include Mojave Desert at lower eleva-
tions, grading through grassland and juniper
shrubland to ponderosa pine forest on Mt.
Trumbull (elevation 2447 m).
Parashant caves were selected for inventory
if they contained deep zone–like conditions.
Cave deep zones are characterized by com-
plete darkness, stable temperature, water-satu -
rated atmosphere, and limited to no airflow
(see Howarth 1980, 1982). This zonal environ-
ment serves as habitat for cave-adapted arthro-
pods. Two of 7 caves met all the deep zone cri-
teria. However, caves meeting most of these
criteria (i.e., complete darkness, stable tem-
perature, limited airflow) were also included
in our study. Because troglomorphic animals
often represent rare and endemic taxa, caves
supporting these animals are considered of
high management and conservation value.
Therefore, we sampled caves with deep zones
and deep zone–like conditions.
To safeguard caves and their resources, we
referred to all of the caves in this study by
using an alphanumeric coding system devel-
oped by the Na tional Park Service (NPS) rather
than actual cave names. Parashant headquarters
in St. George, Utah, has the cipher table with
cave codes and names. A copy of this paper
with actual cave names is on file with both the
National Park Service in St. George and the
National Cave and Karst Research Institute,
Carlsbad, New Mexico.
We sampled arthropods, bats, and other ver -
tebrates in the following date ranges: 4–14
August 2005 and 16–26 September 2005. Win-
ter roost surveys for hibernating bats were
conducted during 4–10 March 2006.
Arthropod Sampling
Based on our site visits, we identified 6
caves that were believed to support deep zone–
like conditions, and we included one addi-
tional cave due to the presence of a summer
bat roost. Five caves (PARA 1801, PARA 2602,
PARA 2204, PARA 1001, and PARA 1401)
were sampled using a combined baited and
unbaited pitfall trapping approach and direct
intuitive searches. Two caves (PARA 0802 and
PARA 2202) were sampled using direct intui -
tive searches and opportunistic hand-collect-
ing throughout each cave’s length.
In caves sampled with pitfall traps, we de -
ployed traps for 4 days, with 2 traps placed
within each of the 3 primary zones—light, twi-
light, and dark. Because the location of the
twilight zone shifts temporally and season -
ally, we estimated the location of this photic
zone during summer. Pitfall trap construction
consisted of two 32-ounce stacked plastic
containers (13.5 cm height, 10.8 cm diameter
rim and 8.9 cm base). We used approximately
4.9 mL of peanut butter as bait and placed it
in the bottom of the exterior container. At the
bottom of the interior container, we made sev-
eral dozen holes so the bait could “breathe”
to attract insects. We buried containers to
the rim when possible, built rock ramps to the
trap rim in other cases, and covered all traps
with a caprock. Prior to removing traps at the
end of the sample period, we searched around
each trap to identify and capture individuals
attracted to the bait but not captured within
the trap (Poulson and Culver 1969).
No pitfall traps were placed in PARA 2202
due to safety issues associated with a return
visit. Although PARA 0802 lacked deep zone–
like conditions and we considered the pres-
ence of cave-adapted arthropods un likely, we
sampled the cave for arthropods because of
the presence of a bat roost (and thus guano).
We searched each cave in areas deemed
most likely to contain certain arthropods. These
areas included detritus deposits, areas with
condensed water and mud, guano deposits,
and active speleothems. We searched for ar -
thropods for at least 40 minutes (2 observers
×20 min each) in each zone (i.e., light, twi-
light, and dark). We also opportunistically col-
lected arthropods by traversing the length of
the cave and searching for and collecting ar -
thropods as encountered (2 observers per cave).
Arthropod Identification
We used a combination of existing keys and
worked with staff members at both the Colo -
rado Plateau Museum of Arthropod Biodiver-
sity and the Colorado Plateau Research Sta-
tion at Northern Arizona University (NAU) to
identify arthropods to the lowest taxonomic
level possible. For several taxonomic groups,
we sent specimens to experts, including Rolf
Aalbu (Coleoptera: Tenebrionidae), Thomas
Barr Jr. (Coleoptera: Rhadine), Earnest Ber -
nard (Col lembola), Theodore Cohn (Orthop -
tera: Rhamphidophoridae), Carl Dick (Dip tera:
Nycteri biidae), Mark Harvey (Pseudoscorpi-
ones: Cher netidae), Robert Johnson (Hyme -
noptera: For micidae), Edward Mockford (Pso -
coptera), Pierre Paquin (Arachnida), William
Shear (Opiliones), Jon Gelhaus (Diptera: Tipu -
lidae), and Chen Young (Diptera: Tipulidae).
Voucher specimens for most arthropod groups
identified through this work are deposited at
the Museum of Northern Arizona, Flagstaff.
Vertebrate Sampling
BATS.—During August and September
2005, we surveyed for bats by employing a
combination of techniques including (i) mist-
netting and harp-trapping at cave entrances
during evening emergence; (ii) hand-netting
within one cave; (iii) visually identifying bats
as encountered within each cave; and (iv) exam -
ining caves for evidence of bat use. For each
bat captured, we determined sex, age, weight,
and reproductive status (Kunz 1982). Cave
entrances were presumed to serve as night
roosts if we documented guano and/or insect
parts on the cave floor within 5 m of a cave’s
We set 2 mist nets and a harp trap within
the entrances of PARA 1801, PARA 0802, and
PARA 1401. Mist nets and harp traps were
deployed before sunset and removed 2 hours
after bats had started their evening emer-
gence. We used 6-m, 2-shelf Avinet mist nets
( and a G6 Cave Catcher harp
trap (Bat Management, www.batmanagement
.com; maximum catch area 3.35 m2). We hand-
netted bats within PARA 2602.
During late winter (4–10 March 2006), each
cave was visited once to search for and count
hibernating bats. When possible, we visually
identified all bats encountered to species. No
hibernating bats were handled during this work.
OTHER VERTEBRATES.—Within each cave,
we searched for and recorded the presence of
all other vertebrates. Sign of other vertebrates
included direct observation, scat, feathers, and
skeletal remains.
We detected 52 cavernicolous taxa, including
44 arthropods, 4 bats, and 4 other vertebrates
(see the Annotated List of Morphospe cies, page
8). Of these, arthropods comprised 10 eiso -
dophiles, 6 troglophiles, 8 questionable troglo -
philes, 7 trogloxenes, 8 accidentals, 3 unknowns
TABLE 1. Arthropods, bats and other vertebrates identified across all cave specificity functional groups, Grand Canyon–Parashant National Monument.
Eisodophiles Eisodoxenes Troglophiles Troglophiles? Trogloxenes Accidentals Unknown Parasites
Arthropods 10 6 8 7 8 3 2
Bats 4 — —
Other vertebrates 2 1 1
and 2 parasites; all bats were trog loxenes, and
other vertebrates included one troglophile, one
accidental, and 2 eisodoxenes (Table 1). Ob -
served arthropod richness ranged from one to 14
morphospecies per cave. The rank order of caves
by arthropod species richness was PARA 1801
(14), PARA 2602 (13), PARA 1001 (11), PARA
2204 (8), PARA 2202 (5), PARA 0802 (1), and
PARA 1401 (1). We detected bats in most caves
(71.4%); PARA 0802, PARA 1401, and PARA
2602 all contained at least 2 bat morpho species.
Numbers for other vertebrates ranged from
one to 2 morphospecies per cave; both PARA
1001 and PARA 2202 supported 2 vertebrate
Taxonomically, the most morphospecies-
diverse groups of arthropods were spiders
(Araneae; n= 14), beetles (Coleoptera: 6 differ-
ent families; n= 7), cave crickets (Or thoptera:
Rhaphidophoridae; n= 5), ants (Hy menoptera:
Fo r mi c i d ae ; n= 4), flies (Diptera: 4 different
families; n= 4), and book lice (Psocoptera: 3
different families; n= 3). Other groups of par-
ticular interest due to their ecological and evo-
lutionary relationships to cave environments
include harvestmen (Opiliones; n= 1), mites
(Acari; n= 2), and springtails (Collembola; n=
1). The remaining arthropods detected within
caves include true bugs (Hemiptera; Reduvii -
dae; n= 1), fleas (Si phonaptera; n= 1) and
bristletails (Thysanura; n= 1).
On a regional level, several taxonomic
groups were represented by 4 or more mor-
phospecies. Spiders were the most commonly
detected animal in all caves visited, represent-
ing ~30% of the arthropods detected. PARA
1801 and PARA 2202 contained the highest
diversity of spiders, 5 and 4, respectively.
Other notable groups include beetles (7 mor -
pho species from 5 caves), crickets (5 morpho -
species from 3 caves), Psocoptera (3 morpho -
species from 3 caves), and ants (4 morphospecies
from 3 caves). All ants that we detected repre-
sent accidental occurrences (R. Johnson, per-
sonal communication, 2006).
Across all arthropod groups, most morpho -
species were limited to a single observation
within a single cave. However, both Leio bunum
townsendii and Entomobrya zona were en -
countered across 3 caves.
When we considered total numbers of ob -
served individuals per light zone, we found
that the twilight zone contained the largest
number of individuals (~2523), followed by
the light (~543) and dark zones (~39; Table
2). PARA 1001 contained a large cricket roost
(estimated to contain >1000 individuals). Al -
though the crickets occurred throughout PARA
1001, most were detected within the light and
twilight zones. They were often found den-
ning in large groups along the walls and ceil-
ings and within several ceiling fissures. PARA
2202 and PARA 2204 supported large num-
bers of Leiobunum townsendii (estimated in the
hundreds) denning in packed clusters in large
overhead fissures within the twilight zones of
each cave. When we shined our light on these
clusters, the harvestmen would rock, sway,
and often aggregate into tighter clusters.
This work resulted in discovery of 7 new
species (with 2 new genera), 3 possible new spe -
cies, 1 range extension, and 1 possible range
extension. The 2 new genera (comprising 2 new
spe cies) are a book louse (Psocoptera: Sphae -
ropsocidae: Troglosphaeropsocus voylesi, Mock -
ford 2009) and a cave cricket (Rhaphidophori-
dae: cf Ceuthophilus n. gen. n. sp., Cohn and
Swanson, unpublished data). Additional new
spe cies included the following: a new pseudo -
scorpion (Chernetidae: Tuberochernes n. sp.,
det. M. Harvey), a cave cricket (Ceuthophilus
n. sp., det. T. Cohn); a carabid beetle (Rhadine
n. sp., Perlevis species group, det. T. Barr), and
2 tenebrionid beetles (Eschatomoxys pholeter
Thomas and Pape 2007 [Pape et al. 2007] and
Eleodes [Caverneleodes] wynnei Aalbu 2012
[Aalbu et al. 2012]). Images of select taxa are
provided in Fig. 1. Two additional Ceutho philus
sp. (n. sp.? a and n. sp.? b) from PARA 1801
may represent new species; the specimens are
“distinctive, and both may be undescribed” (T.
Cohn, personal communication, 2006).
TABLE 2. Total number of arthropods observed per
photic zone by cave.
Light Twilight Dark
PARA 1801 20 16 2
PARA 2202 ~1000 4
PARA 2602 14 6 13
PARA 2204 9 ~1000 13
PARA 0802 1
PARA 1401 1
PARA 1001 >500 >500 6
TOTAL 543a2523a39
aThe values were estimated for large populations of crickets (PARA 1001) and
harvestmen (PARA 2202 and PARA 2204) and were likely underestimated for
each cave.
Our efforts extended the range of the Spe -
leketor flocki (Psocoptera: Psyllipsocidae), which
was previously known from 2 other localities in
the southwest: Tucson Mountains, southern Ari-
zona, and southeastern Nevada. One accidental
ant species, Paratrechina hystrix?, may repre-
sent the first record of this species in Arizona
(R. Johnson, personal communication, 2006).
Specimens match the description of P. hystrix
in part; however, because no workers were
collected, this identification cannot be confirmed
(R. Johnson, personal communication, 2006).
Five of the 7 caves (71.4%) contained bats.
Of these, we identified 3 roosts of an unknown
function, 2 night roosts, and a hibernaculum.
PARA 1401 serves as both a summer roost and
We documented a fringed myotis (Myotis
thysanodes; Fig. 2C) colony (~30 individuals)
at PARA 1801 during the September 2005
survey. We trapped one nonreproductive fe -
male using a harp trap. We were unable to
determine the function of this roost based
upon the one individual captured. Given the
presence of a fresh guano pile, we ascertained
the general location of the roost, which was at
the back of the cave.
We confirmed 2 pallid bat (Antrozous pal-
lidus; Fig. 2B) roosts during August surveys. In
PARA 0802 and PARA 2602, we observed ~50
and ~100 individuals, respectively. We cap-
tured one postlactating female, 4 nonreproduc-
tive females (1 juvenile, 1 adult, and 2 undeter-
mined), and one nonscrotal male, us ing a harp
trap and mist nets at PARA 0802. These bats
were not marked, so there was the possibility of
double counting. In PARA 2602, we captured
one nonreproductive female and one nonscro-
tal male with handheld nets. We tentatively
identified a pallid bat (on the basis of its light
brown color and size) flying within PARA
Fig. 1. Five new species (including 2 new genera) discovered on Grand Canyon–Parashant National Monument, Arizona:
A, Rhadine n. sp., Perlevis species group, on cave floor of PARA 2204; B, high resolution postmortem image of Tub er och er nes
n. sp.; C, a breeding pair of Eleodes (Caverneleodes) wynnei (Aalbu et al. 2012) from PARA 1001; D, postmortem image of
Troglosphaeropsocus voylesi (courtesy of E. Mockford); E, a breeding pair of cf Ceuthophilus n. gen. n. sp. from PARA 1001.
Fig. 2. Three cave-roosting bat species confirmed on Grand Canyon–Parashant National Monument, Arizona: A, 3
hibernating Corynorhinus townsendii from PARA 1401; B, 3 Antrozous pallidus from PARA 0802; C, Myotis thysanodes
in hand during harp-trapping and mist-netting operations at the entrance of PARA 1801.
In August 2005, we counted approximately
10 Myotis sp. emerging from PARA 1401.
Numerous lepidopteran wings were docu-
mented within the entrance and twilight zones
of PARA 0802 and PARA 2602. Corynorhinus
townsendii is one of the more common trog -
loxenic bats in northern Arizona. This bat uses
cave entrances as night roosts, where it has
been documented removing the wings of moths
prior to consuming the abdomen (e.g., Lopez-
Gonzalez 2005). In West Virginia, Sample and
Whitmore (1993) have shown this species to
preferentially consume moths over other ar -
thropods. Because of this documented diet pref -
erence and the large accumulation of moth
wings, we suggest these caves are probably
used as night roosts by C. townsendii.
During our early March 2006 surveys, we
observed 9 C. townsendii (Fig. 2A) hibernating
within the deep zone of PARA 1401 at mid-cave.
Other Vertebrates
We identified woodrat (Neotoma spp.) mid-
dens in 4 caves, American porcupine use (Er -
ethizon dorsatum) at 2 caves, owl use of one
cave, and small carnivore use of one cave. We
documented Neotoma spp. middens in the en -
trance and twilight zone of 4 caves (PARA
2202, PARA 2602, PARA 2204, and PARA 0802).
Woodrats and their middens are commonly ob -
served along sheltered rock outcrops, crags, rock
shelters and cave entrances; thus, wood rats are
considered troglophiles. Because we did not trap
small mammals during this study, we were un -
able to provide species-level identifications.
Three woodrat species are known to occur
within Parashant. The Mexican woodrat (Neo -
toma mexicana) occurs within rocky outcrops
and slopes and is found in open woodland and
transition-zone vegetation (Cornely and Baker
1986). Stephen’s woodrat (Neotoma stephensi)
is often found in association with juniper trees
(Juniperus spp.; Jones and Hildreth 1989). Also,
the white-throated woodrat (Neotoma albigula)
may also occur at the mid-elevations of the
monument (Macêdo and Mares 1988). Thus,
woodrat activity may represent one or more of
these species at any given site.
We found significant use by American por-
cupine in both PARA 1001 and PARA 1401.
Woods (1973) indicates that porcupines den in
caves, hollow trees, and logs. However, we did
not find evidence of recent activity in either
cave. In PARA 1001, we confirmed use by
porcupine based on an extensive deposition of
scat and quills, extending from the middle por -
tion to the back of the cave. We found scat and
quills littering the entrance of PARA 1401, as
well as a fully articulated porcupine skeleton
in the cave. Evidence of owl roosting activity
(i.e., pellets) was documented in the twilight
zone of PARA 2202.
We found scat from a small carnivore (possi-
bly ringtail) in the dark zone of PARA 2202.
This could be one of 3 small carnivores known
to use caves in the Southwest, including ring-
tail (Bassariscus astutus; e.g., Peck 1980, Boden -
hamer 1989, Pape 1998, Hill et al. 2000, 2001,
Strong 2006), skunks (Mephitidae), or raccoon
(Procoyon lotor; e.g., Winkler and Adams 1972).
The following is an annotated list of morpho -
species detected from 7 caves on Grand Can -
yon–Parashant National Monument, Arizona.
Phylum Arthropoda
Class Arachnida
Order Araneae
Famil y Pho lcid ae
Undetermined genus and species.
Det. P. Paquin. Eisodophile.
One juvenile specimen was collected from
the twilight zone of PARA 1801. Because the
specimen was an immature, it could not be
identified beyond family level (P. Paquin, per-
sonal communication, 2006).
Physocyclus sp.
Det. P. Paquin. Eisodophile.
One male specimen was collected from the
light zone of PARA 2204.
Psilochorus sp.
Det. P. Paquin. Troglophile?
One female specimen was collected from
the dark zone of PARA 2602. Given that it was
collected from the dark zone, we suggest this
spider may be a troglophile.
Family Theridiidae
Achaearanea sp. Det. P. Paquin. Eisodophile.
One female specimen was removed from a
web within the light zone of PARA 1801. Peck
(1980) suggests this genus occurs in caves, as
well as in more mesic epigean environments
in Arizona, California, New Mexico, and Utah.
Family Linyphiidae
Undetermined genus and species.
Det. P. Paquin. Eisodophile.
Two juvenile specimens belonging to the
same genus and species were collected from
the light zone of PARA 1801. Because they
were juveniles, they could not be identified
beyond family level.
Agyneta sp.
Det. P. Paquin. Troglophile?
We collected one female specimen from the
dark zone of PARA 2204. Given that it was
collected from the dark zone, we suggest this
spider may be a troglophile.
Family Tetragnathidae
Metellina sp. Det. P. Paquin. Eisodophile.
Two juvenile specimens were collected from
the light zone of PARA 1001.
Family Tengellidae
Undetermined genus and species a.
Det. P. Paquin. Troglophile?
One juvenile specimen was collected from
the dark zone of PARA 2202. Given that it was
collected from the dark zone, we suggest this
spider may be a troglophile.
Undetermined genus and species b.
Det. P. Paquin. Troglophile?
We collected 2 juvenile specimens of a
genus and species different from the afore-
mentioned specimen from the dark zone of
PARA 2202. Given that these specimens were
collected from the dark zone, we suggest this
spider may be a troglophile.
Undetermined genus and species c.
Det. P. Paquin. Eisodophile.
One juvenile specimen was collected from
the light zone of PARA 1801.
Family Plectreuridae
Plectreurys sp.
Det. P. Paquin. Eisodophile.
We collected one specimen from a web
within the twilight zone of PARA 2202.
Family Uloboridae
Uloborus sp.
Det. P. Paquin. Eisodophile.
One female specimen was collected from
the light zone of PARA 1001.
Family Oecobiidae
Oecobius sp. Det. P. Paquin. Eisodophile.
One juvenile wall spider was collected from
the twilight zone of PARA 2602.
Order Opiliones
Family Sclerosomatidae
Leiobunum townsendii Weed, 1893.
Det. W. Shear. Trogloxene.
We collected 3 specimens in the light zone
of PARA 1801. In the twilight zones of both
PARA 2204 and PARA 2202, several hundred
individuals were observed within ceiling fis-
sures. These individuals were denning close
together in large masses. We suspect these
opillionids den in fissures during the day, and
exit to forage at night. This species was also
observed throughout the length of PARA 1001.
Order Pseudoscorpiones
Family Chernetidae
Tuberochernes n. sp.
Det. M. Harvey. Troglophile?
We collected 3 specimens from the dark
zone of PARA 1001. These specimens are with
a taxonomist and will ultimately be described.
Subclass Acari
Acari species a.
Specimens misplaced by taxonomist. Unknown.
Two specimens were collected from the
twilight zone of PARA 2602.
Acari species b.
Specimens misplaced by taxonomist. Unknown.
Two specimens were collected from the
light zone of PARA 1801.
Class Entognatha
Subclass Collembola
Order Entomobryomorpha
Family Entomobryidae
Entomobrya zona Christiansen and Bellinger,
1980. Det. E. Bernard. Troglophile.
Specimens from this family were collected
in the light (n= 3) and twilight (n= 10) zones
of PARA 1801; the light (n= 4), twilight (n=
1), and dark (n= 7) zones of PARA 2602; and
the light zone (n= 4) of PARA 2204. We ob -
served this collembolan in the light zones of 3
caves and the twilight zones of 2 caves. We sug-
gest this animal is a troglophile because (1) it
was found throughout the lengths of 2 caves; (2)
it is a largely nonvagile edaphic organism, and
(3) collembolans are routinely found in caves.
Class Insecta
Order Coleoptera
Family Anobiidae
Undetermined species.
Det. C. Drost. Accidental.
We collected 3 specimens from the light
zone of PARA 1801. Most members of the
family of death watch or spider beetles occur
in dry vegetation (Triplehorn and Johnson
2005). Members of this family are occasionally
noted in caves where they may occur in pack-
rat midden material.
Family Carabidae
Rhadine n. sp. (Perlevis species group).
Det. T. Barr. Troglophile.
A total of 6 specimens were collected, con-
sisting of 3 males and one female from PARA
2204 and one male and one female from PARA
2202. These specimens represent an unde-
scribed species (T. Barr, personal communica-
tion, 2007).
Family Bruchidae
Undetermined species.
Det. R. Delph. Accidental.
We collected 2 specimens from the light
zone of PARA 1801. This seed-boring beetle
was detected within the cave entrance but
is not expected to use caves unless seeds
are transported into the cave by aeolian
Family Leiodidae
Undetermined species.
Det. J. Wynne. Troglophile?
Twelve specimens were collected from 2
Parashant caves. The specimens consist of one
from the light zone of PARA 1001 and 10 from
PARA 2204 (4 from the light zone and 6 from
the twilight zone). This beetle species was ob -
served denning in large numbers within the
fissures in the twilight zone and in association
with Leiobunum townsendii. Most leiodid bee-
tles are habitat generalists and feed as both
adults and larvae on certain fungi or microor-
ganisms associated with decaying organic mat-
ter (e.g., Majka and Langor 2008). We suggest
these beetles may have a commensal relation-
ship with harvestmen (opilionid spiders) and
may feed on fungi growing on spiders’ feces.
There is a literature, rich with examples of
troglomorphic species, that confirms that leio-
dids use caves (e.g., Peck 1974, 1978). We sug-
gest this morphospecies may be a troglophile.
Family Mordellidae
Undetermined species.
Det. R. Delph. Accidental.
Fifteen specimens were collected including
3 specimens from PARA 2602 (2 from the dark
zone and one from the light zone) and 12 speci -
mens from PARA 2204 (2 from the dark zone
and 10 from the twilight zone). Mordellid lar-
vae feed on decaying wood and vegetation,
while adults feed on flowers (Triplehorn and
Johnson 2005). Because both of these caves
were dry and detritus deposition was low, we
suggest this morphospecies is accidental.
Family Tenebrionidae
Eschatomoxys pholeter Thomas and Pape
2007 (new species). Troglophile.
This species was newly discovered and de -
scribed through this research. This beetle was
collected from the twilight zone of PARA 2602.
Pape et al. (2007) considered this beetle a trog -
lophile. Representing the fourth documented
locality, this species was also collected in Ram-
part, Bat, and Christmas Tree Caves, Grand
Canyon National Park (Pape et al. 2007).
Eleodes (Caverneloedes)wynnei Aalbu, Smith
& Triplehorn 2012 (new species). Troglophile.
This species was newly discovered and de -
scribed through this research. Three speci-
mens were collected from the twilight zone of
PARA 1801 and the light zone of PARA 1001.
Also, we collected tenebrionid larvae within
PARA 1801. All Caverneleodes species are con -
sidered troglophiles (Aalbu et al. 2012).
Order Diptera
Family Nycteribiidae (formerly
known as Hippoboscidae)
Basilia antrozoi Townsend, 1893.
Det. C. Dick. Parasite.
One female specimen was removed and col -
lected from a captured Antrozous pallidus at
the entrance of PARA 0802.
Family Phoridae
Undetermined species.
Det. J. Wynne. Troglophile?
We collected 2 specimens from the dark
zone of PARA 1001. This cave contains cricket
guano and decaying vegetation. Adults of this
species may be feeding on the fungus growing
on these 2 nutrient sources.
Family Sciaridae
Undetermined species.
Det. J. Wynne. Troglophile?
We collected one specimen from the dark
zone of PARA 1001. This cave contains cricket
guano and decaying vegetation. Larvae of this
morphospecies are likely feeding on the fungus
growing on these 2 nutrient sources. Addition-
ally, larvae of some of these sciarid species are
known to be predaceous (Cole and Schlinger
1969, Triplehorn and Johnson 2005). No larvae
were de tected within PARA 1001.
Family Tipulidae
Tipula kaibabensis Alexander 1946.
Det. J. Gelhaus and C. Young. Trogloxene.
One specimen was photographed in the twi-
light zone of PARA 1001, and one was collected
from PARA 0802. Tipulids use dark damp places
for dens during the day (J. Gelhaus, personal
communication, 2013). Peck (1981) and Reeves
et al. (2000) considered Tipulids detected in
caves in the SE United States to be trogloxenes.
Order Hemiptera
Family Reduviidae
Triat oma cf rubida.
Det. J. Wynne. Accidental.
We collected one specimen from the light
zone of PARA 1001. Given that this species is
sanguinivorous and that we didn’t observe any
vertebrates within this cave, we suggest the
occurrence is accidental.
Order Hymenoptera
Family Formicidae
Note: All ant morphospecies were detected within cave
entrances and in only association with baited pitfall traps.
We consider all of these ants to be accidental.
Solenopsis xyloni McCook, 1879.
Det. R. Johnson. Accidental.
Five specimens were collected from the en -
trance of PARA 2602.
Pheidole vistana Forel, 1914.
Det. R. Johnson. Accidental.
We collected 3 specimens from the en -
trance of PARA 2602.
Paratrechina cf hystrix.
Det. R. Johnson. Accidental.
We collected 2 specimens from the entrance
of PARA 1801. R. Johnson (personal commu-
nication, 2006) suggests it may be P. hystrix. If
so, this record is the first for Arizona (R. John-
son, personal communication, 2006). Paratre -
china hystrix is a northern species known to
occur in Nevada and Utah. R. Johnson (per-
sonal communication, 2006) suggests the spec-
imens match the description in part; however,
because we did not collect any workers, a reli-
able identification is not possible.
Pheidole sp.
Det. R. Johnson. Accidental.
One specimen was collected from the en -
trance of PARA 2204. R. Johnson indicated
the specimen was a single minor worker. Mi -
nors are often difficult to identify to species,
and the specimen was not examined further
(R. Johnson, personal communication, 2006).
Order Orthoptera
Family Rhaphidophoridae
cf Ceuthophilus n. gen. n. sp.
Det. T. Cohn and A. Swanson.
Trogl oxe ne.
Nine specimens (8 females and one male)
were collected from the light and twilight zone
of PARA 1001. T. Cohn (personal communica-
tion, 2006) indicated that this new genus has
clasping cerci with unique short spines on
their base, a subgenital plate that matches no
other specimens in his collection, and a dis-
tinctive last tergite.
Ceuthophilus n. sp.
Det. T. Cohn.
Trogl oxe ne.
One male was collected from the twilight
zone of PARA 2204. T. Cohn (personal com-
munication, 2006) suggests that the structures
on this specimen are unique and that the speci -
men represents an undescribed species.
Ceuthophilus n. sp. a?
Det. T. Cohn. Undescribed?
Trogl oxe ne.
We collected one female from the entrance
of PARA 1801. Although we will require adult
males for confirmation, T. Cohn (personal com-
munication, 2006) indicated that the fe male has
distinctive characters and may be undescribed.
Ceuthophilus n. sp. b?
Det. T. Cohn. Undescribed?
Trogl oxe ne.
Another female was collected from the en -
trance of PARA 1801. T. Cohn (personal com-
munication, 2006) indicated that the female
has an “extraordinarily elongate and curved ovi -
positor” and is distinctive and may be unde-
scribed. Male specimens will be required to
confirm this.
Undetermined Ceuthophilus sp.
Det. T. Cohn.
Trogl oxe ne.
There was at least one undetermined im -
mature female Ceuthophilus species collected
in PARA 1401. Because the specimen was an
immature, it was not possible to identify it
beyond genus. However, it may also represent
a new species (T. Cohn, personal communica-
tion, 2006).
Order Psocoptera
Family Sphaeropsocidae
Troglosphaeropsocus voylesi Mockford 2009
(new genus and species).
Det. E. Mockford. Unknown.
This animal represents both a new genus
and species discovered through this research.
We collected one specimen from the twilight
zone of PARA 2602 (located in the Mojave
Desert). The cave is dry and dusty with little
to no aeolian-deposited detritus; however, it is
used as a summer roost by Antrozous pallidus.
Until we have additional information on this
psocid’s occurrence within caves, we consider
its cave affiliation to be unknown.
Family Psyllipsocidae
Psyllipsocus ramburii Sélys-Longchamps,
1872. Det. E. Mockford. Troglophile.
This species was trapped in the light, twi-
light, and dark zones of PARA 1801 and the
dark zone of PARA 2602. Because psocids are
routinely found living in caves, we suggest it is
a troglophile.
Family Prionoglarididae
Speleketor flocki Gurney, 1943.
Det. E. Mockford. Troglophile.
We collected a nymph of this species from
the dark zone of PARA 2602. E. Mockford
(personal communication, 2006) indicated that
although the specimen was a nymph, its head
markings are unmistakable. This specimen
rep resents the third locality for this species in
the western United States. It has been con-
firmed from a cave in the Tucson Mountains
and Gypsum Cave, southeastern Nevada.
This psocid routinely lives in caves, and be -
cause a nymph was found in the dark zone of
this cave, we suggest it is probably a troglophile.
Order Siphonaptera
Undetermined family, genus, and species.
Det. J. Wynne. Parasite.
One specimen was collected from the dark
zone of PARA 2602. This cave supports a pos-
sible maternity roost for Antrozous pallidus,
which is likely the host of this siphonapteran
and explains its occurrence.
Subclass Apterygota
Order Thysanura
Family Lepidotrichidae
Undetermined species.
Det. J. Wynne. Eisodophile.
We trapped one silverfish within the en -
trance of PARA 2602.
Phylum Chordata
Class Mammalia
Order Chiroptera
Family Vespertilionidae
Myotis sp.
Det. J. Wynne. Trogloxene.
Approximately 3 Myotis sp. were observed
during exit counts at PARA 1401. According to
Arizona Game and Fish Department’s (AGFD)
Heritage Data Management System, this ob -
servation is likely either M. thyasanodes or M.
yumanensis. Only M. yumanensis is known to
roost both in caves and human-made struc-
tures (AGFD 2003a, 2003b). Williams et al.
(2006) suggest this species is commonly found
near small- to moderately sized bodies of wa -
ter, and of the 4 habitat types investigated, this
species occurs in riparian woodland more than
in all other habitats combined. Though there
are 2 water tanks within 3 miles of PARA
1401, these tanks are intermittent water sources.
We suggest the best candidate water source is
probably Imlay Reservoir, approximately 8.85
km from the cave.
Myotis thysanodes Miller 1897. Fringed
myotis. Det. J. Wynne. Trogloxene.
One nonreproductive female was captured
in a harp trap at the entrance of PARA 1801.
We estimated roost size between 20 and 30
individuals. We suggest PARA 1801 may be a
maternity roost. This bat roosts in caves, mines,
and buildings (O’Farrell and Studier 1980).
Antrozous pallidus Allen, 1862. Pallid bat.
Det. J. Wynne. Trogloxene.
Pallid bats were identified in PARA 2602,
PARA 0802, and possibly PARA 2202. Two
individuals (one adult female and one non-
scrotal male) were captured and identified in a
handheld net in PARA 2602. During our sur-
vey in August 2005, we observed ~100 indi-
viduals roosting in the boulder breakdown at
the entrance of PARA 2602. At PARA 0802,
we used a combined mist-netting / harp-trap-
ping approach to capture 7 individuals (2 post-
lactating adult females, one nonreproductive
juvenile female, 2 nonreproductive undeter-
mined females, one nonscrotal juvenile male,
and one nonreproductive adult female). This
roost contained ~50 individuals and was lo -
cated in the rock fissures in the ceiling within
the cave’s light zone. In PARA 2202, we ob -
served a bat flying in the twilight zone whose
pelage was consistent with A. pallidus. Al -
though we did not confirm it was a pallid bat,
the observation likely represented this spe -
cies. While pallid bats are found roosting in
caves, a majority of data suggest they roost
primarily in rock crevices and outcrops (Her-
manson and O’Shea 1983).
Corynorhinus townsendii Cooper 1837.
Townsend’s big-eared bat.
Det. J. Wynne. Trogloxene.
Nine individuals were observed hibernat-
ing in the twilight zone of PARA 1401. This
was the only documented hibernaculum on
the Grand Canyon–Parashant National Monu-
ment. Lepidopteran wings were also observed
within the entrance of PARA 2602 and PARA
0802, suggesting night use by this species of
bat. Corynorhinus townsendii is a cave-roost-
ing bat, but it also roosts in mines and build-
ings (Kunz and Martin 1982).
Order Rodentia
Family Cricetidae
Neotoma sp. Packrat or woodrat.
Det. J. Wynne and K. Voyles. Troglophile.
Midden activity was documented in the en -
trances and into the light zones of PARA 2202,
PARA 2602, PARA 2204, and PARA 0802.
Woodrats use cave entrances rock outcrops,
rock fissures, and other suitable features for
establishing dens.
Family Erythizontidae
Erethizon dorsatum Cuvier 1822. North
American porcupine.
Det. J. Wynne. Eisodoxene.
Both PARA 1001 and PARA 1401 have been
used extensively by porcupine. In PARA 1001,
guano deposition of 2–10 cm was ob served
throughout the cave; however, there were no
signs of recent porcupine use. A fully articu-
lated porcupine skeleton was photo graphed
amid a deep deposition of guano in PARA
1401. Though the North American porcupine
dens in caves, it also will den in other features
offering similar microhabitats (e.g., hol low logs
and trees). Strong (2006) indicates that this
species is commonly documented in caves in
the Chihuahuan Desert, New Mexico.
Order Lagomorpha
Family Leporidae
Lepus californicus Gray 1837. Black-tailed
jackrabbit. Det. J. Wynne. Accidental.
A black-tailed jackrabbit skeleton was found
approximately 5 m from the entrance of PARA
1001. This animal likely fell into the cave,
became trapped, and eventually died. The car-
cass provided a pulsed food resource for scav-
enging cavernicoles.
Class Aves
Order Strigiformes
Undetermined family, genus and species.
Det. K. Voyles and J. Wynne. Eisodoxene.
We documented owl pellets within the twi-
light zone of PARA 2202. Owls routinely roost
in cave entrances and other suitable habitats
both during the day, and also during the night
between hunting outings.
This study represents the first regional all-
taxa biological inventory of caves in the south-
western United States. Our study resulted in
discovery of at least 7 new species (with 2 new
genera) and 3 potential new species (which
will be described in future publications), as
well as 2 range extensions and one possible
range extension of arthropods. Additionally,
we identified 5 bat roosts and cave use by sev-
eral other vertebrates. Though our study has
contributed significantly to the natural history
of the region, cave biological research within
Parashant remains incomplete.
Two of the most morphospecies-rich caves,
PARA 1801 (14 morphospecies) and PARA
2602 (13 morphospecies), supported a largely
epigean arthropod community. Both caves con -
tained roosting bats, which provided nutrients
via guano. PARA 1001 (11 morphospecies) con -
tained water condensation on the ceiling and
walls and supported the largest known cricket
den in Arizona (likely on the order of thou-
sands of individuals). Such a large number of
crickets generate a significant nutrient load in
the form of cricket eggs, nymphs, and guano.
The ecological importance of cave crickets
has been widely documented (e.g., Barr 1967,
Howarth 1983, Taylor 2003, Culver 2005, Poul -
son 2005), and we suggest that the presence
of crickets was why this cave supported a
higher arthopod diversity. Additionally, given
the significant nutrient loading provided by
crickets and the presence of a cave deep zone,
PARA 1001 may also support cave-adapted
No troglomorphic taxa were identified dur-
ing this survey. Detections of cave-adapted
taxa are reportedly low for northwestern Ari-
zona. However, there are cave-adapted taxa
known regionally. For example, 3 troglobites
and one stygobite are known from Grand Can -
yon National Park (Wynne et al. 2007), and a
cave-limited millipede (Shear et al. 2009) and
a cave-adapted copepod have been collected
from the BLM–Arizona Strip lands adjacent to
Parashant (J. Wynne unpublished data).
Concerning more regional patterns, we
identified at least 18 arthropods with strong
cave affinities (e.g., trogloxenes or troglophiles)
and 21 accidentals or eisodophiles from 7
caves. Given the arid conditions of the desert
Southwest, we suggest that few ground-dwell -
ing arthropods in the Southwest are genetically
predisposed to colonizing the often more mesic
cave environment. Barr (1968) suggests that
most trog lobites were preadapted to the cave
environment in that they previously inhabited
similar mesic habitats such as leaf litter, moss,
or deep soils. By extension, given that these
more mesic habitats are nonexistent in the
Mo jave Desert and juniper shrublands within
the study area, a contemporary preadapted
pool of cave colonists seems to be lacking, and
thus may be reflected by the low number of
ground-dwelling cavernicoles observed.
Similarly, Peck (1978, 1980) suggests that
the low numbers of cave-adapted taxa in
southwestern U.S. caves may be due to the
low nutrient input and high aridity associated
with southwestern cave systems. At Wupatki
National Monument, Welbourn (1976) indicated
that moisture was the most limiting abiotic
factor in the earth cracks he sampled. While
not completely excluding the likeli hood that
troglomorphic arthropod diversity will be low
in northern Arizona, Wynne et al. (2007) sug-
gested that the low numbers of cave-adapted
taxa in Grand Canyon caves may reflect lim-
ited sampling effort and perhaps inappropriate
techniques for detecting cave-adapted taxa.
While troglomorphic taxa in the South -
west may be depauperate in comparison to
cave-obligate communities in the mesic cen-
tral and eastern portions of the United States,
we maintain that sampling effort and ineffec-
tive sampling techniques may still explain the
lack of troglomorphic taxa detected in Para -
shant caves.
For future work, we recommend a multi-
year systematic study design, including an in -
creased number of sampling stations per cave
for timed searches and baited pitfall trapping
(see Wynne 2013), as well as direct intuitive
searches and bait stations (provisioned with
chicken liver, mushrooms, blue cheese, and
sweet potato) within deep cave zones (e.g.,
Howarth et al. 2007). This approach would (a)
provide us with a more thorough inventory
of cave-dwelling arthropods, (b) provide for
inferential statistical comparisons across sites,
and (c) in crease the likelihood of detecting
cave-adapted organisms. Secondly, we suggest
that sample size (i.e., the number of caves
inventoried) may also be an issue. We provide
the results from only 7 caves within Parashant.
In the adjacent BLM–Arizona Strip lands and
Grand Canyon National Park lands, there are
approxi mately 250 and >400 known caves,
respectively. As we expand our efforts into
other management units in Arizona and obtain
a larger sample of study sites, we hypothesize
that more troglomorphic taxa will be discov-
ered. Only through increased sampling ef fort
and use of systematic techniques will we be
able to make ecological comparisons to other
regions in the southwestern United States and
infer whether the American Southwest truly
supports low troglomorphic diversity.
Management Implications
Through this research, we tentatively iden-
tified one cave as a high management priority
because of the presence of cave-dwelling
arthropods. PARA 1001 supports the only
known type localities for the new cricket
genus, cf Ceuthophilus n. gen. n. sp. Though
this genus may occur elsewhere on and off
Grand Canyon–Parashant National Monument,
this has not been confirmed. To increase our
knowledge of the current population and to
better define its range, we recommend (1)
conducting a multiyear census of the popula-
tion during the most productive time of year—
between the late monsoonal season and the
early post-monsoonal season (mid-August to
mid-September); (2) sampling additional caves
within Parashant to search for this animal; and
(3) conducting surface surveys in the region to
estimate the actual geographic distribution of
this new cricket. Additionally, we know of only
one other cave in Arizona that supports such a
large cricket population. Because cave crickets
contribute important nutrients via guano, eggs,
nymphs, and cricket carcasses and are a prey
source for predaceous arthropods (e.g., Barr
1967, How arth 1983, Taylor 2003, Culver 2005,
Poulson 2005), the cricket population is likely
an im portant component of this ecosystem.
Given the population size within PARA 1001,
we suggest that the presence of crickets likely
has an important bottom-up effect on ecologi-
cal structure and species richness and also sug -
gest that the actual number of species will be
higher than the observed richness presented
in this study.
We recommend that all caves containing
both summer roosts and hibernacula (PARA
0802, PARA 1401, PARA 1801, and PARA 2602)
be closed to recreational use until these sites
can be thoroughly studied and their functions
determined. There is little argument that hu -
man disturbance to bat roosts is detrimental
(e.g., Humphrey 1969, Mohr 1972, McCracken
1988, 1989, Harnish 1992, Brown et al. 1993,
Boyles and Brack 2009). Additionally, there
is a growing threat of white-nose syndrome
(WNS), a disease caused by Pseudogymnoas-
cus destructans (refer to Minnis and Lindner
2013), which will likely spread through the
western United States. This psychrophilic fun-
gus has resulted in the “most precipitous de -
cline of North American wildlife in the past
century” (BCI 2010). Furthermore, WNS has
resulted in the mortality of over 7 million bats
and has been detected in 23 states and 5
Canadian provinces (Wynne 2014).
Little is known concerning the habitat char -
acteristics of bat hibernacula in the Ameri can
Southwest. In Arizona alone, we lack baseline
information regarding the locations of cave-
roosting hibernating bats (A. McIntire, Ari-
zona Game and Fish, personal communication,
2010). Thus, resource managers can best pre-
pare for the westward advance of WNS by
intesifying their efforts to establish population
estimates of nonmigratory cave-roosting bats
and characterize habitat of cave roosts in the
western United States.
To improve our knowledge regarding bat
distributions in northwestern Arizona, we rec-
ommend the following: (1) additional surveys
of summer roosts (at PARA 0802, PARA 1801,
and PARA 2602) during the middle of the
nursery period (mid- to late June) to deter-
mine whether these roosts are actually mater-
nity/nursery sites; (2) establishment of annual
or biennial winter bat censuses of the hiber-
naculum cave (PARA 1401); and (3) expanded
searches to identify additional cave roosts,
with follow-up inventories, roost monitoring,
and habitat characterization as appropriate.
We extend special thanks to Darla Sidles,
Mark Sogge, Karen Yanskey, and Carol Cham-
bers for their logistic support and adminis -
trative guidance. Ben Solvesky provided field
support and assisted with bat identifications.
The image of Troglosphaeropsocus voylesi was
provided by Edward Mockford, University of
Illinois, Normal. Eric Alejandro Pinto gra-
ciously translated our abstract to Spanish. Jeff
Bradybaugh, Charles Drost, Jennifer Fox, Stew -
art Peck, and 2 anonymous reviewers pro-
vided valuable suggestions leading to the im -
provement of this manuscript. This work was
funded by the National Park Service, Grand
Canyon–Parashant National Monument, ARCS–
Phoenix Chapter, and the American Museum
of Natural History, Theodore Roosevelt Memo-
rial Fund.
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Received 22 March 2013
Accepted 11 October 2013
... In the sub-Saharan environment at Djelfa (Algeria), Souttou et al. (2015) using pitfall trap sampling inventoried 45 families of Arthropoda, of which Formicidae dominated with 86.10% followed by an undetermined dipteran family. In Arizona, USA, Wynne and Voyles (2014) sampled approximately 29 Arthropoda families. Moreover, these workers mentioned the existence of seven families of Araneae and six families of Coleoptera. ...
... Moreover, these workers mentioned the existence of seven families of Araneae and six families of Coleoptera. Our observations are similar to the results of Souttou et al. (2015) and those of Wynne and Voyles (2014). According to Pizarro-Araya et al. (2012), in the Atacama desert (Chile), a richness of 30 families was found of which the Mummuciidae was the most sampled with 22.5% followed by the Tenebrionidae (19.4%) and the Gryllidae (18.8%). ...
An inventory of arthropods was carried out at locations in the desert area of Touggourt, southeast Algeria. Samples were collected from two diverse habitats, a palm grove (agricultural habitat) and dunes (natural habitat). Using the Barber pitfall trap, 1100 specimens, divided into four classes, 15 orders, 44 families and 99 species were obtained. In the palm grove, 660 arthropods were trapped, belonging to four classes and 12 orders. Of these four classes, Insecta dominated followed by Crustacea, Arachnida and Entognata. Insecta accounted for 59.49% of the total capture and was dominated by two orders: Hymenoptera (41.81%) and Amphipoda (34.55%). In the Hymenoptera, Cataglyphis sp. was the most abundant (38.2%), followed by Pheidole pallidula (2.3%). In the dunes, 440 individuals were trapped. Insecta was the most abundant (90.69%), and Crustacea and Arachnida were scarce. Of the dominance by insects, Hymenoptera was most abundant (68.15%), and within that order, Cataglyphis bombycina (35.5%) was the most abundant followed by Monomorium subopacum (8.9%). In the palm grove, 42 species were recorded, compared to 57 in the dunes. The Shannon–Weaver index and equitability varied in both stations. In the palm grove, the diversity was 2.6, and the equitability was 0.5. By contrast in the dunes, the diversity was equal to 4 and the Equitability equal to 0.7. The differences in vegetation between the two sites reflect the differences in species diversity.
... As dispersal is assumed to be restricted in these organisms, the degradation of subterranean habitats is believed to represent a threat for the conservation of such short-range endemics. Rare troglobites have therefore been the primary targets of cave conservation efforts worldwide Wynne & Voyles, 2013;Culver & Pipan, 2014;Ferreira, Oliveira & Silva, 2015). ...
... For instance, the cave's deep interior has been compared to a desert since it is largely deprived of trophic resources (White & Culver, 2012;Pipan & Culver, 2013). Troglobites thus rely on external material that is washed into the cave or brought in by mobile species (Poulson & White, 1969;Taylor, Krejca & Denight, 2005;White & Culver, 2012;Wynne & Voyles, 2013). Although our findings match those of a recent analysis of 844 iron caves from the Carajás region, which also found higher species richness in caves containing guano (Jaffé et al., 2016), they reveal that this trophic resource not only supports higher species richness but also a higher functional diversity (assessed via phylogenetic diversity). ...
Full-text available
The degradation of subterranean habitats is believed to represent a serious threat for the conservation of obligate subterranean dwellers (troglobites), many of which are short-range endemics. However, while the factors influencing cave biodiversity remain largely unknown, the influence of the surrounding landscape and patterns of subterranean connectivity of terrestrial troglobitic communities have never been systematically assessed. Using spatial statistics to analyze the most comprehensive speleological database yet available for tropical caves, we first assess the influence of iron cave characteristics and the surrounding landscape on troglobitic communities from the Eastern Amazon. We then determine the spatial pattern of troglobitic community composition, species richness, phylogenetic diversity, and the occurrence of frequent troglobitic species, and finally quantify how different landscape features influence the connectivity between caves. Our results reveal the key importance of habitat amount, guano, water, lithology, geomorphology, and elevation in shaping iron cave troglobitic communities. While mining within 250 m from the caves influenced species composition, increasing agricultural land cover within 50 m from the caves reduced species richness and phylogenetic diversity. Troglobitic species composition, species richness, phylogenetic diversity, and the occurrence of frequent troglobites showed spatial autocorrelation for up to 40 km. Finally, our results suggest that the conservation of cave clusters should be prioritized, as geographic distance was the main factor determining connectivity between troglobitic communities. Overall, our work sheds important light onto one of the most overlooked terrestrial ecosystems, and highlights the need to shift conservation efforts from individual caves to subterranean habitats as a whole.
... Only a limited number of studies have been undertaken to document cave ecosystems in Grand Canyon. While nearby Grand Canyon -Parashant National Monument has received much more focus and is now recognized to have diverse cave ecosystems (Wynne & Voyles 2014). In Grand Canyon, previous work has hinted at the unique nature of the cave-adapted ecosystems (Drost & Blinni 1997, Wang & Holsinger 2001, Pape 2014. ...
... Regarding prey species, we encountered a sizable number of flies (Diptera) within entrances through twilight zones of most caves. While cave crickets have been identified as providing significant nutrient loading in the form of cricket eggs, nymphs, guano and adult cricket carcasses (e.g., Barr 1967;Howarth 1983;Taylor 2003;Culver 2005;Poulson 2005;Wynne & Voyles 2014), flies may serve a similar ecological role in Sierra de las Nieves caves. Large numbers of cave-dwelling flies have been documented within cave entrances in Vancouver Island, Canada (Shaw & Davis 1999) and Meghalaya, India (Disney 2009). ...
Technical Report
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This work represents the first large scale cave biological inventory of caves in Sierra de las Nieves Natural Park, Andalucía, Spain. We sampled seven caves (three low and four high elevation caves) from 22 June through 01 July 2017. We have preliminarily identified at least 42 morphospecies and 13 coarse-level taxonomic groups (i.e., Order or higher) of cave-dwelling arthropods including the relict springtail species, Onychiurus gevorum Arbea 2012. Bats were detected in two of three low elevation caves; a bat roost of unknown type consisting of approximately 100 bats was observed in one cave, and one bat (Myotis sp.) was found torporing in another cave. The common toad (Bufo bufo (Linnaeus, 1758)) was identified in two low elevation caves. We also provide recommendations for additional research to aid in the future management of these resources.
... Through better characterizing this community, researchers and managers may elucidate the pseudoscorpions' role in the community, as well as better define the distribution of these two species within Maomaotou Cave based upon the distributions of suitable habitat and potential prey species. As our sampling was temporally constrained (2 people at~4 hours) during one site visit and employed a single technique, we suggest applying multiple techniques with multiple site visits conducted during the appropriate time of year to garner a more comprehensive picture of the cave arthropod community (Wynne & Voyles 2014;Wynne et al. 2018). If this is not possible, we minimally suggest the deep zone of Maomaotou Cave be intensively and systematically sampled using baits (Howarth et al. 2007), leaf-litter traps (Slaney & Weinstein 1996), and direct intuitive searches using a similar sampling protocol proposed by and Wynne et al. (2018). ...
Two new troglomorphic pseudoscorpion species, Bisetocreagris maomaotou sp. nov. (Family Neobisiidae) and Tyrannochthonius chixingi sp. nov. (Family Chthoniidae) are described from one cave in the tower karst of northern Guangxi Province, China. This cave is located at close proximity to a village and an adjacent urban area. As with many caves in the South China Karst, this feature occurs at an elevation slightly above agriculture and rural activities; thus, we suggest it may be partially buffered from human activities in the lowland areas. We discuss the likelihood of narrow range endemism and provide research and conservation recommendations to guide future management of these two species.
... It is the only known locality for a troglobitic fungus beetle, Ptomaphagus parashant (Peck & Wynne 2013), two cave--adapted pseudoscorpions endemic to this cave (Harvey and Wynne 2014), and the only known locality for a troglomorphic centipede (family Anopsobiidae, cf Buethobius n. sp.; the first cave--adapted centipede known to Arizona). By studying PARA--1001 Cave, we not only have the ability to compare the climate of a non--hibernacula cave to a hibernacula cave, but we also have the ability to characterize the habitat and microclimate of one of the most biological significant caves on the monument (Peck & Wynne 2013;Wynne and Voyles 2014;Harvey and Wynne 2014). ...
Technical Report
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Addressing a knowledge gap concerning the winter ecology of bats on Grand Canyon-‐Parashant National Monument in preparation for the western advance of white-‐nose syndrome (WNS), this paper provides a summary of a three‐year study to estimate population trends of two known cave-roosting bat hibernacula (PARA-0901 and PARA-1401 Caves). Beginning in 2011, we sampled all caves (total 11) likely to support hibernating bats on both Parashant and adjacent BLM lands. Through this effort, colleagues and I identified two hibernacula and three torpor roosts. All but one torpor roost was located on Parashant. The two hibernacula caves became the focus of work in subsequent years (2012 and 2013). Total numbers of hibernating bats ranged from 44 to 51 in PARA­‐0901 Cave, and four to 17 in PARA-­1401 Cave. Most of the bats detected were Corynorhinus townsendii with Myotis sp. infrequently detected in both caves. No visible signs of white­‐nose syndrome (WNS) were observed during the three-year period on either hibernating bats visually examined or in post-­field examination of photographs. Analysis of six sediment samples (with 1 control on surface) from PARA-­0901 Cave tested negative for Pseudogymnoascus destructans (the fungus that causes WNS). In PARA-0901 Cave, the largest hibernaculum, we deployed 41 data loggers and in PARA-­1001 Cave, a non-­hibernaculum cave, we deployed 42 to collect rock surface temperature, ambient temperature, relative humidity and barometric pressure data for two years. For both PARA-0901 and PARA-­1001, we collected 3D cartography data, 3D geospatial data of all microclimatic instrument locations, and 3D geospatial data of all observed hibernating bats. These data will be used to develop models to characterize how microhabitats are selected for hibernation. I will use these models to (a) parameterize habitat requirements of bat hibernacula for at least one cave and (b) simulate climate change effects on this cave to predict whether this roost will become unsuitable for bats at some point in the future. PARA-­1401 Cave was gated in 2009; as a result, the roost is now protected. Presently, PARA-­0901 Cave lacks any safeguards. This cave is the largest known hibernacula on the monument (and in northern Arizona, in general) and is located within one mile of a frequently used cattle tank and corral and is within 300 feet of a single-­track road. To best protect this roost, we recommend this cave be closed to recreational use and the lower chamber ultimately gated. Recommendations are also provided for the establishment of a Western states comprehensive sampling and monitoring strategy of hibernacula for early detection of WNS.
Full-text available
Subterranean habitats may be considered limiting for animal colonization, especially for ants, due to permanent darkness and mainly because of oligotrophic conditions. While not as deep as limestone caves, iron ore caves and other subterranean habitats may be more available for colonization because of their shallower depth. We use the richness and composition of ants to assess how differences in habitat structure affect the biodiversity and ecosystem function between cavities and surrounding epigean landscapes. We predicted that the distribution of ants would be different because of the variation in habitat structure and cavity conditions may act as a filter for colonization by ants. A high diversity of ants was found in the 20 sampled cavities (26 species), and most of them were grouped in the generalist trophic guilds. The distribution of ants occurred independently of the type of cavity to which they are associated (caves, impacted caves and mines). Significant differences were observed in ant richness between epigean and cavities habitats, with lower average richness in cavities. The physical attributes of the cavities did not influence richness, mainly because cavity use by ants can usually be explained by their opportunistic habits and generalist lifestyle. Ants can participate directly in the cavities assemblage, playing roles in species composition and trophic functionality, due to the lower use restriction.
Full-text available
Este trabajo representa el primer inventario, a gran escala, de la biología de las cuevas del Parque Natural de la Sierra de las Nieves, Andalucía, España. Se han muestreado siete cavidades, de las cuales tres se localizan a cota relativamente baja, a una altura media de unos 1000 m.s.n.m., mientras las otras cuatro se localizan a una cota relativamente alta, con una altura media de 1600 m.s.n.m. Se han identificado, de modo preliminar, al menos 40 morfoespecies y 13 grupos taxonómicos a escala general (esto es, categorías taxonómicas de nivel orden o superior) de artrópodos que viven en cuevas, incluyendo la especie relicta de colémbolo Onychiurus gevorum Arbea 2012. Los murciélagos se detectaron en dos de las tres cuevas de cota baja; una colonia de murciélagos, posiblemente Rhinolophus ferrumequinum (Schreber, 1774), consistente en aproximadamente 100 individuos que se vio en una de las cuevas; y un murciélago (Myotis sp.) que se encontró aletargado en otra cavidad. El sapo común (Bufo bufo (Linnaeus, 1758)) se ha encontrado en dos de las cuevas de cota baja. Se proponen recomendaciones para desarrollar una investigación complementaria que ayude a la gestión futura de estos recursos biológicos.
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Ever-increasing human pressures on cave biodiversity have amplified the need for systematic, repeatable, and intensive surveys of cave-dwelling arthropods to formulate evidence-based management decisions. We examined 110 papers (from 1967 to 2018) to: (i) understand how cave-dwelling invertebrates have been sampled; (ii) provide a summary of techniques most commonly applied and appropriateness of these techniques, and; (iii) make recommendations for sampling design improvement. Of the studies reviewed, over half (56) were biological inventories, 43 ecologically focused, seven were techniques papers, and four were conservation studies. Nearly one-half (48) of the papers applied systematic techniques. Few papers (24) provided enough information to repeat the study; of these, only 11 studies included cave maps. Most studies (56) used two or more techniques for sampling cave-dwelling invertebrates. Ten studies conducted ≥10 site visits per cave. The use of quantitative techniques was applied in 43 of the studies assessed. More than one-third (42) included some level of discussion on management. Future studies should employ a systematic study design, describe their methods in sufficient detail as to be repeatable, and apply multiple techniques and site visits. This level of effort and detail is required to obtain the most complete inventories, facilitate monitoring of sensitive cave arthropod populations, and make informed decisions regarding the management of cave habitats. We also identified naming inconsistencies of sampling techniques and provide recommendations towards standardization.
Technical Report
Full-text available
I provide a summary of field activities conducted at three caves on 02 February 2013 (0700-1900hr), Grand Canyon-Parashant National Monument, Arizona, USA.
Full-text available
Subgenus Caverneleodes of the genus Eleodes is diagnosed and revised. Six new species from the United States: California (E. microps); Utah and Northern Arizona (E. wynnei), Central Arizona (E. wheeleri), Southern New Mexico (E. quadalupensis), and Mexico (E. thomasi and E. grutus) are described. The biogeography of the subgenus is discussed. Diagnoses and a key are provided to known species of Caverneleodes. Relationships with other Eleodes are discussed. Cave associated Amphidorini are surveyed.
Full-text available
Local species diversity of terrestrial arthropods was determined from a combination of trapping and census in an area of variable passage type in Flint Ridge Cave System in Mammoth Cave National Park, Kentucky. We measured evaporative rate, substrate moisture, substrate organic content, predictability and stability of food and microclimate, substrate diversity, and intensity of flooding. We found significant correlations of species diversity with substrate diversity, substrate organic content, and intensity of flooding.
Of the 15 species found, all are terrestrial and 5 are probably now limited to stream caves in the canyon as troglobites or disjunct populations of troglophiles. These 5 species probably descended from forest litter-inhabiting ancestors living near the caves during past glacial-pluvial climates. This 'life zone' lowering occurred most recently from 24 000 to 14 000 yr ago. When the forest retreated upwards at the beginning of the present interglacial (about 8000 yr ago), some of the litter invertebrates which had entered the caves were locally isolated in them when adjacent epigean populations went extinct. -from Author
Twelve species of Eleodes Eschscholtz from the United States and Mexico are described as new: Eleodes aalbui (California), E. spiculiferus (Texas). Mexico: E. bidens (Durango), E. brucei (Durango), E. corrugans (Michoacán), E. mirabilis (Nuevo León), E. muricatulus (San Luis Potosí), E. platypennis (Jalisco), E. reddelli (Nuevo León), E. samalayucae (Chihuahua), E. scyropterus (Hidalgo), and E. watrousi (Durango).
Ptomaphagus lincolnensis and Ptomaphagus manzano, small-eyed, flightless, high elevation forest litter species from New Mexican mountains, are discribed as new. Their relation to cave species is possible. High elevation spruce-fir forest of New Mexico harbor a suite of terrestrial litter arthropods "preadapted" for cave colonization. That cave faunas in the southwest (west of the Edwards Plateau of Texas) are comparatively lacking in troglobites (obligate cavernicoles) must thus be due to environmental dryness, and not to a lack of ancestral species with cave colonizing potential and ability.