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A study of the biology and ecology of Bat Cave, Grand Canyon National Park, was conducted during a series of four expeditions to the cave between 1994 and 2001. A total of 27 taxa, including 5 vertebrate and 22 macro-invertebrate species, were identified as elements of the ecology of the cave. Bat Cave is the type locality for Eschatomoxys pholeter Thomas and Pape (Coleoptera: Tenebrionidae) and an undescribed genus of tineid moth, both of which were discovered during this study. Bat Cave has the most species-rich macro-invertebrate ecology currently known in a cave in the park.
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Department of Entomology, University of Arizona, Tucson, Arizona 85721,
Abstract: A study of the biology and ecology of Bat Cave, Grand Canyon National
Park, was conducted during a series of four expeditions to the cave between 1994 and
2001. A total of 27 taxa, including 5 vertebrate and 22 macro-invertebrate species,
were identified as elements of the ecology of the cave. Bat Cave is the type locality
for Eschatomoxys pholeter Thomas and Pape (Coleoptera: Tenebrionidae) and an
undescribed genus of tineid moth, both of which were discovered during this study. Bat
Cave has the most species-rich macro-invertebrate ecology currently known in a cave in
the park.
This paper documents the results of a biological and
ecological analysis of Bat Cave on the Colorado River
within Grand Canyon National Park conducted during
four expeditions to the cave between 1994 and 2001. The
study focused on the macro-invertebrate elements present
in the cave and did not include any microbiological
sampling, identification, or analysis. Access to caves in
the park is strictly regulated and requires either an
approved research permit or a cave entry permit, granted
through the Grand Canyon Division of Science and
Resource Management.
Grand Canyon National Park has extensive cave
resources, but there has been little research on cave-
inhabiting invertebrates in the park. The earliest effort was
conducted by the Cave Research Foundation at Horseshoe
Mesa in 1977 and 1978 (Welbourn, 1978). The scope of that
project was not strictly biological, but did include a search
for invertebrates in eight caves contained in the Mooney
Falls member of the Redwall limestone. The effort produced
a total of fourteen invertebrate species, including a new
species of garypid pseudoscorpion (Archeolarca cavicola
Muchmore). The lack of species diversity was attributed to
the lack of available moisture in the caves. Peck (1980)
looked at three caves in the Grand Canyon, all of which, like
Bat Cave, were in the Muav limestone. All three of these
caves contain active streams that have their outfall in major
tributaries to the main gorge of the Colorado River below
the North Rim of Grand Canyon. Peck’s effort identified a
total of 15 invertebrate species from the 3 caves. One of those
caves, Roaring Springs Cave, was revisited by Drost and
Blinn in 1994 and 1995 (Drost and Blinn, 1997), and their
effort increased the invertebrate species recorded from that
cave from the original 10 found by Peck to 19, including the
first records of aquatic invertebrates from a Grand Canyon
Cave. A review of cave-invertebrate studies in the park,
which included 9 reports addressing 16 caves, was prepared
by Wynne et al. (2007). Their compilation resulted in a list
of approximately 37 species of cave macro-invertebrates
currently known from caves there. Wynne and others have
recently performed invertebrate surveys in caves in the
Grand Canyon–Parashant National Monument, and have
already encountered several undescribed cave-inhabiting
macro-invertebrate species, including three new genera.
These include a camel cricket (Rhaphidophoridae), a
barklouse (Psocoptera), and a macrosternodesmid millipede,
Pratherodesmus (Shear et al., 2009; Wynne and Drost, 2009).
Bat Cave is located in the north wall of the main gorge
of the Colorado River within Grand Canyon National
Park (Fig. 1, insert). The entrance is at an elevation of
580 meters, approximately 260 meters above the river. The
cave is formed along a major vertical fault in the Cambrian
Muav limestone and consists of just over one thousand
meters of surveyed passage, mostly along the single main
trunk passage. From the entrance, the cave trends a few
degrees east of north (Fig. 1). The cave is essentially
horizontal, with a total vertical relief of only 75 meters.
Bat Cave is named for the large maternity colony of
Mexican free-tailed bats (Tadarida brasiliensis I. Geoffroy)
that resides there. The first record of the cave is from the
1930s. Shortly thereafter, there were efforts to mine the
bat guano for fertilizer. These early efforts were mostly
unproductive. In 1958, the property was purchased by the
U.S. Guano Corporation. U.S. Guano invested 3.5 million
dollars in the setup of the mining operation, only to find
that the extent of the guano deposit was only 1 percent of
their original estimate. Difficulties with the haul system
across the canyon, and specifically the lack of a significant
guano deposit, forced the abandonment of the mining
operation in 1960 (Billingsley et al., 1997). The support
towers for the haul system’s cables are still present on the
top of the south rim of the canyon and below the cave on the
R.B. Pape – Biology and ecology of Bat Cave, Grand Canyon National Park, Arizona. Journal of Cave and Karst Studies, v. 76, no. 1,
p. 1–13. DOI: 10.4311/2012LSC0266
Journal of Cave and Karst Studies, April 2014 N1
north side of the river. A wooden plank-way, portions of
which are elevated, traverses approximately the first 145 me-
ters of the cave from an elevated entrance platform. Sections
of large flanged pipe approximately 30 cm in diameter and
other mining equipment remain in the cave. Most of the
animal species associated with the cave are arthropods tied
directly to the bat guano-supported food web.
The four expeditions to Bat Cave were conducted on
March 12, 1994; March 29 and 30, 1996; October 24, 1997;
and October 26, 2001. Access to the cave was made up the
river by boat from Lake Mead. Visits to the cave were
scheduled seasonally and temporally to avoid disturbance
Figure 1. Map of the biologically active front portion of Bat Cave, Grand Canyon National Park, showing regional location
(inset) and survey stations mentioned in the text.
2NJournal of Cave and Karst Studies, April 2014
to the resident bat population. Even so, at least some bats
were present on all visits to the cave. During the March
visits, no more than a few hundred bats were present at any
time, mostly around the roost at survey station D1 (Fig. 1).
All efforts were made to keep disturbance to the bats at a
minimum. This roost is at the east edge of the main trunk
passage, and the roost area can be skirted with minimal
disturbance to the bats.
The 5 visits involved approximately 34 hours of
observation and sampling of invertebrates in the cave.
The methods were low impact, consisting of a visual search
of passage floors, walls, crevices, ledges, and breakdown-
boulder accumulations. Additional habitat niches have
been created in Bat Cave by the presence of wooden
walkways and machinery remaining from the abandoned
mine operation. Particular attention was paid to floor
debris, under which many invertebrates commonly shelter.
Debris that was overturned was replaced after examination
to minimize impact to microhabitats. The principal benefits
of this type of documentation are minimal habitat
disturbance, low animal population impact, and ease of
sampling in the cave environment. Since obligate cave
invertebrate populations may be small and may be
adversely impacted by over-sampling, no pitfall or other
trapping devices were used. Pitfall traps should only be
utilized where the traps can be regularly monitored.
Initial sampling of specimens was kept to a minimum,
and no more than six individuals of each type were taken.
Both males and females were included in these samples for
species where their sex was readily evident. Additional
sampling for type-series specimens was later performed for
the tenebrionid beetle and the undescribed tineid moth.
Specimens were sampled with forceps or a camel hair brush
dampened with ethanol. Specimens were placed in vials of
ethanol for transport to the laboratory.
Two approximately 0.3 kg samples of fresh bat guano
were taken on March 29, 1996, for Berlese extraction. Each
of these samples was a composite of smaller sub-samples
taken in closely spaced areas within an approximately
0.5 m
area near the apex of the guano deposits. One
sample was taken at each of the two main roosts, near
survey stations D1 and B7. Specimens were sampled under
two park permits (9503-09-02 and GRCA-2001-SCI-0043).
Relative-humidity and ambient air-temperature mea-
surements were recorded at several points along the main
trunk passage of the cave on March 29, 1996 using a digital
temperature/humidity meter (Hanna Instruments model HI
A total of 27 taxa, including vertebrates, were identified
from Bat Cave during this study. Table 1 is a taxonomic
summary of the species documented and lists the assigned
ecological group (trogloxene, troglophile, troglobite, or
accidental), guild (scavenger, fungivore, predator, parasite,
or phytophage), and relative abundance (abundant, com-
mon, uncommon, or rare) for each species. Ecological
groups used in this paper are defined as follows: A
trogloxene is an animal that enters caves to fill some
ecological need, such as obtaining food, water, shelter, etc.,
but that cannot survive without returning to the surface to
meet some of its life-cycle requirements. A troglophile is an
animal that is capable of completing its life cycle within
caves, but may also do so elsewhere. A troglobite is an
obligate cave animal, which cannot live outside of the cave
environment. The occurrence of parasites in the cave is in
association with other species of animals, but because of
their presence in the cave during part of their life cycle we
consider them to be trogloxenes. Accidentals are just what
the label implies, animals that wander or fall into the cave,
would not normally occur in such habitats, and obtain no
benefit from their presence in the cave.
Phylum: Chordata
Class: Aves
Order: Passeriformes
Family: Troglodytidae
Catherpes mexicanus Swainson
A single canyon wren was observed in the entrance
room of the cave on October 26, 2001. Canyon wrens are
common in cave entrance areas, where they forage, shelter,
and occasionally nest.
Class: Mammalia
Order: Chiroptera
Family: Vespertilionidae
Myotis californicus Audubon & Bachman
The presence of this insectivorous bat at Bat Cave is
known from acoustic monitoring that was conducted at the
entrance in 1997 (Charles Drost, pers. comm). It is
assumed that only small numbers of these bats use the
cave. Their contribution to the nutrient input to the cave is
greatly overshadowed by the population of the Mexican
free-tailed bat.
Myotis yumanensis H. Allen
The presence of this insectivorous bat at Bat Cave is
known from acoustic monitoring that was conducted at
the entrance in 1997 (Charles Drost, pers. comm). It is
assumed that only small numbers of these bats use the cave.
Their contribution to the nutrient input to the cave is greatly
overshadowed by the population of the Mexican free-tailed
bat. This species almost always occurs near a permanent
water source, and it is likely a relatively common animal in
caves and crevices along the Colorado River.
Family: Molossidae
Tadarida brasiliensis I. Geoffroy
The Mexican free-tailed bat maternity colony at Bat
Cave is the major source of nutrient input into the cave, with
its guano serving as the foundation of the cave’s invertebrate
Journal of Cave and Karst Studies, April 2014 N3
Table 1. Summary of fauna documented at Bat Cave, Grand Canyon National Park, Arizona.
Taxon Ecological Group Guild Abundance
Class Aves
Order Passeriformes
Catherpes mexicanus Trogloxene Predator Uncommon
Class Mammalia
Order Chiroptera
Myotis californicus Trogloxene Predator Uncommon?
Myotis yumanensis Trogloxene Predator Uncommon?
Tadarida brasiliensis Trogloxene Predator Abundant
Order Carnivora
Bassariscus astutus Trogloxene Predator Common
Class Arachnida
Order Sarcoptiformes
Glycyphagus sp. Troglophile Fungivore Uncommon
Cubaglyphus sp. Guanophile Scavenger Common
Sapracarus tuberculatus Troglophile Scavenger? Rare
Order: Mesostigmata
Chiroptonyssus robustipes Troglophile Parasite Abundant
Mitonyssoides stercoralis Troglophile Predator Common
Order: Trombidiformes
Pimeliaphilus sp. (Undes.?) Trogloxene Parasite Common
Order Araneae
(Kukulcania sp.?) Troglophile Predator Rare
Kibramoa suprenans Troglophile Predator Common
Loxosceles sp. Troglophile Predator Uncommon
Psilochorus sp. Troglophile Predator Uncommon
Selenops sp. Trogloxene Predator Uncommon
Order Pseudoscorpiones
Neoallochernes stercoreus Troglophile Predator Abundant
Class Hexapoda
Order Psocoptera
Psyllipsocus ramburii Troglophile Fungivore Common
Order Neuroptera
Eremoleon pallens Troglophile Predator Common
4NJournal of Cave and Karst Studies, April 2014
food web. During the period the field work was conducted,
the summer population of the bats was estimated at 285,000
(Charles Drost, pers. comm). The bats currently roost at two
major sites within the cave. The primary site is at survey
station B7, 265 meters from the entrance. The second,
smaller roost is located at survey station D1, 229 meters into
the cave. Overflow bats from these roosts probably
congregate in wider sections in the main corridor closer to
the entrance, between survey stations A11 and B2. Recent
guano deposits in the front sections of the cave are not
extensive. Currently, the bats seldom go deeper into the cave
than the passage constriction at survey station B9,
308 meters from the entrance. No recent guano deposition
was observed beyond survey station B9.
A large fossil guano deposit is present at the back of the
cave beginning at survey station B14. Over a period of time
the main cave passage was gradually cut off at a passage
constriction at survey station B9. The bedrock ceiling of
the cave passage at this point dips steeply downward.
Evidently the annual accumulations of guano deposited at
the roost centered at survey station B7 eventually increased
in depth until the slope of the deposit met with the ceiling
at survey station B9, isolating the back portion of the cave.
Based on the presence of old mining pipes at this point, the
back section of the cave was apparently reopened during
the last mining efforts. A sample of the top of the fossil bat
guano beyond station B9 was dated at 6.5 ka, and the
bottom of the fossil guano deposit was dated at 14.5 ka
(Wurster et al., 2008). The more recent date would
represent the time of last regular use of the back portion
of the cave by the large Tadarida colony.
Order: Carnivora
Family: Procyonidae
Bassariscus astutus Lichtenstein
Ringtail scats and tracks were observed throughout the
front half of the cave, back to the active bat roosts at
survey stations D1 and B7. A ringtail was observed on
two of the five visits into the cave. Both were nocturnal
sightings, which is the normal activity period for the
animals. One sighting was at 1929 hrs on March 29, 1996,
along the east wall of the cave adjacent to survey station
B2, where a single animal was observed headed deeper
into the cave. At the time there were a few hundred free-
tailed bats active at the roost at survey station D1. A
second sighting was at 1930 hrs on October 24, 1997,
between survey stations A5 and A6, where the main
passage narrows. Along the west wall of the cave at this
point there is a wooden walkway support structure
that extends toward the middle of the passage. We had
earlier noted that there were several dead Tadarida lying on
and around this small structure (Fig. 2). There were
abundant ringtail scats present along the west wall here.
The ringtail was observed perched on the east edge of
the support structure. This put it near the center of
the walkway area and well within the flight path of any
low-flying bats. There were still a few bats leaving the
cave at this time. The ringtail was not observed attempt-
ing to take bats out of the air, but it is probable that is
how it was capturing live bats. It is unknown why the
several dead bats on the support structure noted earlier
in the day had not been consumed by the ringtail. With
the abundant bats as a dependable food source from at
Taxon Ecological Group Guild Abundance
Order Coleoptera
Dermestes carnivorus Troglophile Scavenger Rare
Eschatomoxys pholeter Troglophile Scavenger Abundant
Smicronyx imbricatus Accidental Phytophage Uncommon
Order Siphonaptera
Sternopsylla distincta Trogloxene Parasite Common
Order Diptera
Fannia sp. Trogloxene? ? Rare?
Order Lepidoptera
New genus and species Troglophile Scavenger Common
Tinea (pallescentella?) Troglophile? Scavenger Rare
Order Hymenoptera
Sceliphron caementarium Trogloxene Predator Uncommon
Table 1. Continued.
Journal of Cave and Karst Studies, April 2014 N5
least April through October and the natural shelter
provided by the cave, it is possible that ringtails may also
den in the cave. Ringtails would probably use the front
entrance room for denning, since there is a jumble of rocks
which could provide a secluded area for that purpose.
Other rock-pile accumulations occur near survey station
Based on sightings and an abundance of their scats,
ringtails apparently regularly use caves in the southwestern
U.S. They use caves for shelter, as a source of water and
some foods, and probably den in caves, at least occasion-
ally. Ringtails are opportunistic in their feeding, taking
equally from animal and plant materials seasonally. Birds,
wood rats, bats, mice, and other small rodents are
commonly taken (Taylor, 1954; Murie, 1974; Poglayen-
Neuwall and Toweill, 1988). Ringtails probably regularly
take dying adult bats and young that fall onto cave floors
below roosts. Records for lizards and snakes from two-
thousand-year-old ringtail scats were documented in caves
in the western part of Grand Canyon by Mead and Van
Devender (1981). Arthropods make up a significant part of
the diet of ringtails (Toweill and Teer 1977; Poglayen-
Neuwall and Toweill, 1988). I found cave cricket (Ceutho-
philus nr. pinalensis) parts in a ringtail scat at Arkenstone
Cave in southern Arizona. Taylor (1954) also recorded
Ceuthophilus sp. from ringtail scats. A microscopic
examination of ringtail scats from Bat Cave revealed
abundant body parts of the Bat Cave tenebrionid beetle
(Eschatomoxys pholeter). The only other invertebrate of
adequate abundance in Bat Cave that might be used as
food by ringtails is the plectreurid spider (Kibramoa
suprenans Chamberlin). Ringtails are known to take
arachnids (Poglayen-Neuwall and Toweill, 1988), and it
is doubtful that they would be intimidated by spider-web
silk in pursuing this species.
Phylum: Arthropoda
Class: Arachnida
Order: Sarcoptiformes
Family: Glycyphagidae
Glycyphagus sp.
This dust mite is likely fungivorous and likely feeds on
molds associated with the bat guano.
Family: Rosensteiniidae
Cubaglyphus sp.
This animal is an undescribed species of Cubaglyphus
(Barry OConnor, pers. comm.).
Family: Suidasiidae
Sapracarus tuberculatus Fain & Philips
This is a species of very small mites that is known from
a variety of habitats, including caves (Barry OConnor,
pers. comm.). The feeding habits of the species are not
known (Barry OConnor, pers. comm.). Other mite records
from caves in Grand Canyon National Park are an anystid
mite from several of the Horseshoe Mesa caves (Welbourn,
1978) and a rhagidiid mite (Rhagidia cf. hilli) from Roaring
Springs Cave (Peck, 1980).
Order: Mesostigmata
Family: Macronyssidae
Chiroptonyssus robustipes Ewing
This mite is an obligate parasite of the Mexican free-
tailed bat. The bats groom them off their bodies and the
mites drop to the guano pile below. It is not known
whether any of the mites manage to return to the bats.
Many of them are likely consumed by the various
invertebrate predators in the guano pile. The mite
population in the guano piles was not at its peak during
our visits to the cave since the number of bats present was
generally low. The fresh guano accumulation was quite
limited when the Berlese samples were taken.
Mitonyssoides stercoralis Yunker, Lukoschus, and Giesen
This mite is the only species in its family (Macronyssidae)
that is not a blood-feeding parasite on a vertebrate host, but
is predatory on the rosensteiniid mites that occur in bat
guano. M. stercoralis is found in the guano of both molossid
and vespertilionid bats. The previously known range of the
species was from Indiana in the U.S. to southern Brazil
(Radovsky and Krantz, 2003). Presence of M. stercoralis in
Grand Canyon National Park is a significant range
extension for the species.
Order: Trombidiformes
Family: Pterygosomatidae
Pimeliaphilus sp.
Species of Pimeliaphilus are parasitic on lizards and a
variety of arthropods (Field et al., 1966; Anderson, 1968;
Ibrahim and Abdel-Rahman, 2011; Delfino et al., 2011). A
single deutonymph and larvae of an undetermined species
of Pimeliaphilus were sampled from Eschatomoxys pholeter
Figure 2. Dead Tadarida killed by ringtail near survey
stations A5 and A6; scale is 15 cm.
6NJournal of Cave and Karst Studies, April 2014
in the cave (Fig. 3). These mites are relatively common on
the beetles, and are typically parasitic at these stages. This
animal may be a new species of Pimeliaphilus (Barry
OConnor, pers. comm).
Order: Araneae
Family: Filistatidae?
Kukulcania sp.?
A single large, blackish spider was found in the apron of
its web on the north side of a large boulder in the middle of
the main trunk passage of the cave at survey station A5. The
spider eluded capture, but is believed to be Kukulcania sp.
Family: Plectreuridae
Kibramoa suprenans Chamberlin
The population of this spider at Bat Cave is pale in color
and may represent a troglophilic population (Darrell Ubick,
pers. comm). No troglophilic plectreurid spider population
has been previously recorded (Ribera and Juberthie, 1994).
This spider is the only large invertebrate predator in the cave-
invertebrate food web here. The primary food of the older,
larger spiders is the tenebrionid beetle Eschatomoxys pholeter.
Below their webs an accumulation of the beetle bodies is
often evident (Fig. 4). The spiders are abundant wherever
there is support for their webs. They build their webs along
the base of the cave walls, among rock piles, and in and under
the walkway and machinery left from the guano-mining
operation. Some recesses in the walls of the cave are
festooned with old webs containing their ecdysed skins, old
egg cases, and carcasses of E. pholeter (Fig. 5). A close-up
view of an individual abandoned web is shown in Fig. 6.
The juvenile spiders spin their webs across small
depressions on the cave walls or in recesses or depressions
in the guano deposits on the floor of the cave. The latter
generally occur at the edge of the active guano deposition
areas where they are proximal to smaller invertebrates
present on the surface of the guano. Interestingly, this
species has not been recorded in other caves in the park.
Family: Sicariidae
Loxosceles sp.
There are several troglophilic and a few troglobitic
species in this genus that occur in the tropics (Ribera and
Juberthie, 1994). This species exhibits no obvious morpho-
logical modifications that would indicate that it is adapted
for living in a cave environment, and it is probably an
epigean form. This may be the same species recorded from
Thunder Cave by Peck (1980) and from Roaring Springs
Cave by Drost and Blinn (1997).
This spider is not a prominent member of the biota
within the cave, but is probably relatively common outside
Figure 3. Undescribed Pimeliaphilus sp. mite on thorax of
Eschatomoxys pholeter.
Figure 4. Numerous carcasses (white arrows) of Eschato-
moxys pholeter in debris pile at the base of an old K.
suprenans web; scale is 15 cm.
Journal of Cave and Karst Studies, April 2014 N7
the cave in rocky areas along the cliff faces. The single
female found was located on the cave floor at the base of the
west wall of the passage, approximately 64 meters from the
cave entrance. A reduction of cave passage dimensions in
this area has an ameliorating affect on the cave microcli-
mate. Closer to the entrance, where passage dimensions are
considerably larger, hot, dry desert air from outside the cave
easily mixes with the cave atmosphere, resulting in
significant drying of the front part of the cave. The area of
this initial passage constriction is where invertebrates of the
guano food-web are first encountered. The brown spider has
encroached on the periphery of the active food web, where it
is not in competition with the dominant plectreurid spider.
When first observed, this spider had a muscid fly (Fannia
sp.) in its chelicerae.
Family: Pholcidae
Psilochorus sp.
This troglophile is a small relative of the common cellar
spider. Members of this family are often a common element
of invertebrate biota of caves in the western U.S.
Psilochorus has not previously been recorded from caves
in the park, but is likely to be present in habitats similar to
that present in Bat Cave. There is some depigmentation in
this species, but no observable reduction of the eyes. Only a
small population of these delicate spiders is found in the
cave, and they typically occur away from areas of intense
invertebrate activity at the fresh guano deposits. Their
more delicate webs and the situations where they build may
indicate that mostly small airborne prey are taken. No prey
debris was observed in their webs. Their diet may include
Figure 5. Several old webs of Kibramoa suprenans hanging from a wall in the cave; scale is 15 cm.
Figure 6. Single old web structure of Kibramoa suprenans
showing white egg-case enclosures at top and aggregation of
shed spider skins trapped in lower portion.
8NJournal of Cave and Karst Studies, April 2014
the guano moth and small dipterous species associated with
the fungus-covered guano and dead bats in the cave.
Family: Selenopidae
Selenops sp.
The presence of Selenops sp. in the cave was limited to a
couple of shed skins found on the walls of the first room of the
cave within the first six meters from the entrance. These spiders
are foragers that typically occur in rocky habitats, spending
most of their time in rock piles, crevices, and caves. Their
presence here is not a surprise. They apparently do not go
deeply enough into the cave to capitalize on prey supported by
the bat guano deposit, where Kibramoa suprenans is the
dominant predator.
Order: Pseudoscorpiones
Family: Chernetidae
Neoallochernes stercoreus Turk
This is the first record of this pseudoscorpion associated
with a bat colony in Arizona (William Muchmore, pers.
comm). This species is apparently the common pseudoscor-
pion associated with active Mexican free-tailed bat guano
deposits, and is found in many caves in Texas where free-
tailed colonies reside (Muchmore, 1992). A mean density of
135 individuals/dm
was recorded for this species at Fern
Cave, Val Verde Co., Texas (Mitchell and Reddell, 1971).
The estimated density of this species in Bat Cave at the time
of sampling is approximately one fourth that value.
However, the sampling was not done during the summer
peak of the bat population and active guano deposition.
Interestingly, the species has not been found at Carlsbad
Caverns National Park, which has a free-tailed colony of
approximately 700,000 bats. The pseudoscorpion species
present there is Dinocheirus astutus Hoff (Muchmore, 1992).
Likely prey of N. stercoreus in Bat Cave includes mites, the
psocid, and collembola, if they are present. No collembola
were observed on or sampled from the guano.
The only other pseudoscorpion known from a cave in
Grand Canyon National Park is Archeolarca. cavicola from
Cave of the Domes on Horseshoe Mesa (Welbourn, 1978).
A. cavicola shows slight morphological modifications for
its subterranean habit (Muchmore, 1981). Other species in
that genus have been reported from caves at Wupatki
National Monument, Arizona, and Guadalupe Mountains
National Park, Texas (Muchmore, 1981).
Pseudoscorpions are a common element of cave and
guano deposit ecosystems. I have observed them in many
caves in the southwestern U.S., often in caves that are quite
dry and where food resources seem to be minimal. Overall,
the group appears to be more tolerant of arid conditions
than other arthropod predators. Additional studies in
caves in the park should reveal additional pseudoscorpion
Class: Hexapoda
Order: Psocoptera
Family: Psyllipsocidae
Psyllipsocus ramburii Selys-Longchamps
This barklouse seems to be the common psocid found
in caves in the southwestern U.S., and indeed, has a
worldwide distribution (Badonnel and Lienhard 1994).
Psocids occurring in cave environments are probably
detritivores, feeding on microflora or other organic sources
(Mockford, 1993). The guano deposits at Bat Cave likely
provide a whole suite of potential nutrient sources for
this species. P. ramburii is associated with cave cricket
(Ceuthophilus nr. pinalensis) guano at Arkenstone Cave in
southern Arizona. There it is the prey of the cave-adapted
pseudoscorpion Albiorix anophthalmus Muchmore (Mock-
ford, 1993; Muchmore and Pape, 1999). It is likely to be a
prey species for Neoallochernes stercoreus in Bat Cave.
Psocids (Psyllipsocidae) were recorded from several of the
Horseshoe Mesa caves (Welbourn, 1978), and they are
likely the same species. P. ramburii was also recorded from
a single specimen at Tapeats Cave (Peck, 1980).
Order: Neuroptera
Family: Myrmeleontidae
Eremoleon pallens Banks
The larval pits of this antlion species are common in the
dry, silty soils of the entrance room of the cave. Adults were
seen flying in the evenings in the main passage in the vicinity
of survey station A7, and small groupings of their wings
were found clustered on the floor of the passage where
Tadarida had captured the animals in flight and hung up on
the ceiling to feed on the insects. This is a new county record
for E. pallens in Arizona (Mohave County), and the only
record of the species from a cave. The type species for E.
pallens was from a mine shaft on Picacho Peak in southern
Arizona. At least one species of Eremoleon (E. longior
Banks) has previously been recorded from caves (Adams,
Order: Coleoptera
Family: Dermestidae
Dermestes carnivorus Fabricius
Dermestid beetles are common faunal elements on bat-
guano deposits in the New World and Asia (Mitchell and
Reddell, 1971; Decu et al., 1998; Gnaspini and Trajano,
2000). Where they occur in association with bat guano,
they are typically the dominant decomposer, and they may
be present in enormous numbers. No dermestid species is
established in the cave, and colonization attempts by
dermestids, which likely occur on occasion, are apparently
repelled by predatory or scavenging elements in the guano
community, possibly including Eschatomoxys pholeter,
which could feed on eggs and/or young larvae of
dermestids. Only a single D. carnivorus was found in the
cave, on October 26, 2001. The animal was probably
attracted to the odor of the guano deposit, which is
detectable even by humans at great distance. The beetle
was found near the front part of the cave (near survey
station A5).
Journal of Cave and Karst Studies, April 2014 N9
Family: Tenebrionidae
Eschatomoxys pholeter Thomas and Pape
This beetle species was described using specimens taken
at Bat Cave during this study. The species also occurs in
other caves in western Grand Canyon National Park and
the Grand Canyon–Parashant National Monument (Pape
et al., 2007). The species is probably not strictly a cave
species, and likely also occurs in dry situations among
cliffs, where it is probably associated with fecal materials
in rodent nests. This is similar to the habit of E. tanneri
Sorenson and Stones, which has been recorded in
southern Utah (Sorenson and Stones, 1959) and in
Marble Canyon, Arizona (Pape et al., 2007). Peck
(1980) recorded a single specimen of Eschatomoxys sp.
from Thunder Cave, also in the Grand Canyon. The
location of this record is about midway between the
known distributions of E. pholeter and E. tanneri.The
Thunder Cave record probably represents one of these
two species, but since the disposition of the specimen is
unknown it could not be examined. The specimen was
taken off a wall of a dry upper passage above some (bat?)
guano (Peck 1980).
Peck (1980) recognized that Eschatomoxys spp. show
morphological adaptations to a subterranean habitat. The
head and pronotum are narrowed, and there is a definite
attenuation of the legs and antennae. He suggested that
this morphology represents adaptation for scavenging in
animal nests and burrows rather than being associated with
caves. This seems reasonable, considering most records of
the genus have not been from caves, except for E. pholeter,
which has so far been recorded only from caves. The
species probably colonizes caves opportunistically from
adjacent shelter and crevice habitats where they are
probably associated with rodents.
The greatest density of the beetles in the cave is on the
active bat-guano deposits. Otherwise they are thinly
distributed throughout the front portion of the cave,
except in the entrance room, which may be too dry. They
scavenge over the surface of the guano and probably feed
on a variety of foods, including dead bats. They were
observed feeding on the dead bats presumed to have been
caught by a ringtail near the front part of the main
passage of the cave (Pape et al., 2007: figure 9). There
were no larvae of E. pholeter found in the guano deposits,
and they were not present in the two guano samples
removed from the cave. The larvae may be present only
during times of the year when bat activity is at its peak.
The beetle is the primary prey of the spider Kibramoa
suprenans at Bat Cave.
E. pholeter is the only invertebrate species that was
observed beyond the passage constriction at survey station
B9. Only a couple of individuals were present there, and it
is probable these were vagrants from the nearby active
guano deposit. A search of the fossil guano deposit for
invertebrate activity gave negative results, and the deposit
is likely depleted of useable nutrients.
Family: Curculionidae
Smicronyx imbricatus Casey
This weevil has been recorded from the Gulf coastal
plain and ‘‘central lowlands’’ (north-central plains) in
Texas to the Sierra Nevada and coast-range sections of the
Pacific mountain system, and it is widely distributed in the
basin and range province (Anderson, 1962). I assume the
weevils emerge in the cave from seeds contained in scats
deposited by ringtails. S. imbricatus was found in moderate
abundance at the peak of the guano deposit at survey
station D1. The natural proclivity of many weevils is to
climb up vegetation and other objects. The only thing
available for them to climb in the cave is a mountain of
guano, so they congregate at the top along with other
invertebrates. The presence of this species is coincidentally
associated with ringtail use of the cave.
Order: Siphonaptera
Family: Ischnopsyllidae
Sternopsylla distincta Rothschild
This flea is known from throughout the southern portion
of the United States, where it is associated primarily with the
Mexican free-tailed bat and the western bonneted bat
(Eumops perotis Schinz) ( Hubbard, 1968). It is found in
large numbers on the active guano piles at Bat Cave when the
bats are present. The bats groom the fleas off their bodies,
and the fleas drop to the guano pile. There is no way for the
fleas to return to the host once they are removed, and some
may become prey of predators on the guano piles. There are
currently no other records of fleas from caves in the park, but
associations with birds, rodents, bats, and other cave-
frequenting mammals makes their presence likely.
Order: Diptera
Family: Muscidae
Fannia sp.
A single individual of this muscid fly was found in the
cave. It had been captured by the single female Loxosceles
spider observed in the cave. Its association with the ecology
of the cave, if any, is not known.
Order: Lepidoptera
Family: Tineidae
Tineid moth (new genus and species)
This animal is a new genus and species (Don Davis,
pers. comm.). This small, weakly flying moth is common in
the cave. They are readily noticed because their wing scales
reflect a silvery color in the light from headlamps. The
larvae of the moths probably derive nourishment directly
from the fresh guano (Vandel, 1965) and would then be
considered guanobionts. Alternatively, they may be fungi-
vores. They may be preyed upon by spiders present in the
cave (Psilochorus sp., Kibramoa suprenans, and Loxosceles
sp.). The moths were apparently attracted to the dead
Tadarida found near the front of the cave. This association
is not understood. The only other record of a tineid moth
10 NJournal of Cave and Karst Studies, April 2014
from a cave in the park is from one of the caves on
Horseshoe Mesa (Welboun, 1978).
Tinea (pallescentella?)
A single individual of this animal was found in the cave.
The moth is larger and distinctly different from the
undescribed species listed above. The specimen was
damaged and was not positively identifiable to species,
but may be T. pallescentella. Its association with the
ecology of the cave is not known.
Order: Hymenoptera
Family: Sphecidae
Sceliphron caementarium Drury
The presence of this sphecid wasp was evident from the
abundant multi-celled mud nests observed throughout the
entrance area. Nests were present on the walls of the cave
and on the wooden walkway supports, pulleys, and cables
left from the guano mining operation. One of the nests
contained sixteen cells. None of the nests were in use
during our visits to the cave. S. caementarium provisions its
nests with spiders. Due to the dry, sheltered environment
inside the cave entrance, the old nest cells may have
accumulated over a very long time.
A series of temperature and relative humidity measure-
ments were taken in the cave on March 29, 1996, beginning
at the cave entrance (at 1617 hrs), and progressing deeper
into the cave to the bat-occupied roost areas. Temperature
and humidity at the cave entrance (drip line) were 17 uC
and 15.4 percent, respectively. Both of these climatic
parameters increased toward the rear of the cave at the
time of our visit. The lower values at the cave entrance were
the result of the large size of the entrance and its proximity
to, and air exchange with, the arid desert environment
outside of the cave. The microclimate of the cave at the bat
roosts is greatly influenced by the presence of the bats,
particularly when they are in residence in large numbers.
Their presence results in a localized increase in both
relative humidity and ambient air temperature. At the first
bat roost (229 meters from the entrance; Fig. 1, survey
station D1), where a few hundred bats were present, the
values were 26 uC and 89.0 percent at fifteen centimeters
above the guano deposit. Deeper in the cave, along the
main trunk passage, the air temperature drops slightly at
the high point in the cave at the main bat roost (265 meters
from the entrance; Fig. 1, between survey stations B7 and
B9). Here the air temperature was 25 uC and the relative
humidity 100.0 percent. The slightly lower air temperature
at this location may have been due to a larger volume of air
that was less affected by the small number of bats present
at the time the measurements were taken. Additionally, air
movement along the main trunk passage may have caused
some minor mixing with cooler air from the front part of
the cave.
The food web in the cave is supported primarily by the
annual deposition of guano introduced by the Mexican
free-tailed bat colony. While some bats are present
throughout much of the year, the greatest numbers are in
residence from April through September. The bats leave
the cave after sunset each evening to feed on insects and
return to roost in the cave and digest their food. Their
urine and fecal pellets drop to the floor and build into large
deposits over many years. The annual guano layer supports
most of the arthropod activity, with layers from previous years,
depleted of nutrients, showing little or no arthropod activity.
Adult bats that die of natural causes fall to the guano
pile and provide additional nutrients. During the summer,
juvenile bats that lose their grasp on the ceiling and are
unable to fly also drop to the guano deposit and are
consumed by arthropods and, probably, ringtails. Parasites
that are groomed from the bats drop to the guano pile,
where they become food sources for other microfauna.
Ringtail scats provide a small amount of additional
allochthonous nutrient input to the cave. The lack of a
rhaphidophorine cave cricket at Bat Cave may be due to
inadequate vegetation in proximity to the cave.
The diagram of the cave food web, as it is currently
understood, is presented in Fig. 7. Most associations
shown are assumed and are based on typical life habits
of the taxa present. However, several of the associations
were documented during the study, including the Mexican
free-tailed bats feeding on the antlion (E. pallens), the
ringtail feeding on both bats (T. brasiliensis) and the
tenebrionid beetle E. pholeter, the plectreurid spider (K.
suprenans) also feeding on E. pholeter, and the brown
spider Loxosceles sp. feeding on the muscid fly (Fannia sp.).
The annual deposit of Tadarida guano is the only
nutrient input of any consequence entering Bat Cave. The
entrance to the cave is horizontal, and the ceiling at the
entrance overhangs the debris slope from the cave,
precluding allochthonous organic debris washing into the
cave and contributing nutrients. Only very small quantities
of water penetrate the thick bedrock overburden at the
main fault along which the cave is developed. The only
water observed in the cave was several small drip points
concentrated in an area along the west wall 230 meters
from the entrance and at the top of the large guano deposit
265 meters from the entrance and very small quantities
of condensate water found in the fossil guano deposit
approximately 360 meters from the entrance. No arthro-
pods were found at any of these water sources.
Most of the invertebrates present in Bat Cave are
associated directly with the active guano deposit. Previous
annual guano layers beneath the freshly accumulating layer
are drier and appear mostly devoid of arthropod activity.
Journal of Cave and Karst Studies, April 2014 N11
However, the drier condition of the lower layers may
provide suitable habitat for eggs, larvae, or pupae of some
species inhabiting the guano. The drier guano may inhibit
potentially harmful fungi associated with the higher
moisture content of the fresh guano. No arthropod activity
was present in the nutrient-depleted fossil guano deposit at
the rear of the cave.
Except for the presence of the large bat colony and the
associated guano deposit, Bat Cave would appear to follow
the pattern described by Welbourn (1978) for the caves at
Horseshoe Mesa and many other caves in Grand Canyon
National Park. Because of the generally arid climate in the
region during the last ten to twelve thousand years (Van
Devender, 1990), many caves, especially those with
multiple entrances, have dried to a considerable degree.
Cave invertebrates that may have been present in more
mesic, wetter times were likely extirpated during this
extended drying period. While Bat Cave is nearly devoid
of free water sources, the humidity in the cave increases
from the front to the rear of the cave. This is due to the
cave having a single entrance, which minimizes air
exchange with the arid exterior environment, and moisture
contributed by the bats through their respiration and
A total of 27 taxa, including five vertebrates and 22
macro-invertebrate species, were identified as elements of
the ecology of Bat Cave. Two of the macro-invertebrates,
Eschatomoxys pholeter (Coleoptera: Tenebrionidae) and
an undescribed genus of tineid moth are species first
discovered during this study. Additional study would likely
reveal additional macro-invertebrate species in the cave.
Bat Cave has the most species rich macro-invertebrate
ecology currently known in a cave in the park.
I thank Robert A. Winfree, Senior Scientist at Grand
Canyon National Park at the time this study was
conducted, for his encouragement and cooperation with
this project. Special thanks are due to Ray Keeler for
handling all the logistics that made the expeditions to the
cave possible. Ray also reviewed the manuscript and
provided the plan map of the cave for Figure 1.
Sincere appreciation is extended to the following specialists
who performed identifications: Barry M. OConnor, Univer-
sity of Michigan (Acari); Darrell Ubick, California Academy
of Sciences (Araneae); William Muchmore, University of
Rochester (Pseudoscorpiones); Don Davis, National Museum
Figure 7. Bat Cave food web diagram. The associations are identified as follows: Solid-filled block arrows indicate direct
consumption. Unfilled block arrows indicate deposition. Dashed-line block arrows indicate likely consumption associations. A
narrow solid line is an intraspecies connection and a narrow dashed arrow is an assumed association. Documented taxa are
shown as solid-bordered boxes and suspected taxa are shown as dash-bordered boxes.
12 NJournal of Cave and Karst Studies, April 2014
of Natural History, Smithsonian Institution (Lepidoptera);
and Charles W. O’Brien, Florida A & M Univ. (Coleoptera:
Curculionidae). I thank Carl Olson, University of Arizona
Department of Entomology, for reviewing the manuscript. I
also thank the three anonymous reviewers whose incorporat-
ed suggestions improved this manuscript. I thank Charles
Drost, USGS-CPRS, for providing the records of the two
species of Myotis from Bat Cave. I thank my wife Esty Pape
for assistance with graphics and proofreading of the
manuscript. This study was partially funded with the aid of
a grant from the National Speleological Society.
Adams, P.A., 1956, New ant-lions from the Southwestern United States
(Neuroptera: Myrmeleontidae): Psyche, v. 63, p. 82–108. doi:10.1155/
Anderson, D.M., 1962, The weevil genus Smicronyx in America north of
Mexico (Coleoptera: Curculionidae): Proceedings of the United States
National Museum, no. 3456, v. 113, p. 185–372.
Anderson, R.G., 1968, Ecological observations on three species of
Pimeliaphilus parasites of Triatominae in the United States (Acarina:
Pterygosomidae) (Hemiptera: Reduviidae): Journal of Medical
Entomology, v. 5, no. 4, p. 459–464.
Badonnel, A., and Lienhard, C., 1994, Psocoptera, in Juberthie, C., and
Decu, V., eds., Encyclopaedia Biospeologica, Vol. 1, Moulis –
Bucarest, Socie´te´ de Biospe´ologie, p. 301–305.
Billingsley, G.H., Spamer, E.E., and Menkes, D., 1997, Quest for the
Pillar of Gold – The Mines and Miners of the Grand Canyon: Grand
Canyon, Arizona, Grand Canyon Association Monograph 10, 112 p.
Decu, V., Juberthie, C., and Nitzu, E., 1998, Coleoptera (Varia), in
Juberthie, C., and Decu, V., eds., Encyclopaedia Biospeologica, Vol. 2,
Moulis - Bucharest, Socie´te´ de Biospe´ologie, p. 1164–1173.
Delfino, M.M.S., Ribeiro, S.C., Furtado, I.P., Anjos, L.A., and Almeida,
W.O., 2011, Pterygosomatidae and Trombiculidae mites infesting
Tropidurus hispidus (Spix, 1825) (Tropiduridae) lizards in northeastern
Brazil: Brazilian Journal of Biology, v. 71, no. 2, p. 549–555.
Drost, C.A., and Blinn, D.W., 1997, Invertebrate community of Roaring
Springs Cave, Grand Canyon National Park, Arizona: The South-
western Naturalist, v. 42, no. 4, p. 497–500.
Field, G., Savage, L.B., and Duplessis, R.J., 1966, Note on the cockroach
mite Pimeliaphilus eunliffei (Acarinae: Pterygosomidae) infesting
Oriental, German, and American cockroaches: Journal of Economic
Entomology, v. 59, no. 6, 1532 p.
Gnaspini, P., and Trajano, E., 2000, Guano communities in tropical caves,
in Wilkens, H., Culver, D.C., and Humphreys, W.F., eds., Subterra-
nean Ecosystems: Amsterdam, Elsevier, Ecosystems of the World 30,
p. 251–268.
Hubbard, C.A., 1968, Fleas of Western North America: Their Relation to
the Public Health: New York, Hafner Publishing Co, 533 p.
Ibrahim, M.M., and Abdel-Rahman, M.A., 2011, Natural infestation of
Pimeliaphilus joshuae on scorpion species from Egypt: Experimental and
Applied Acarology, v. 55, p. 77–84. doi:10.1007/s10493-011-9452-6.
Mead, J.I., and Van Devender, T.R., 1981, Late Holocene diet of
Bassariscus astutus in the Grand Canyon, Arizona: Journal of
Mammalogy, v. 62, no. 2, p. 439–442.
Mitchell, R.W., and Reddell, J.R., 1971, The invertebrate fauna of Texas
caves, in Lundelius, E.L., and Slaughter, B.H., eds., Natural History
of Texas Caves: Dallas, Texas, Gulf Natural History, p. 35–90.
Mockford, E.L., 1993, North American Psocoptera: Gainesville, Florida,
Sandhill Crane Press, Flora and Fauna Handbook 10, 455 p.
Muchmore, W.B., 1981, Cavernicolous species of Larca, Archeolarca, and
Pseudogarypus with notes on the genera, (Pseudoscorpionida,
Garypidae and Pseudogarypidae): Journal of Arachnology, v. 9,
no. 1, p. 47–60.
Muchmore, W.B., 1992, Cavernicolous pseudoscorpions from Texas and
New Mexico (Arachnida: Pseudoscorpionida), in Reddell, J.R., ed.,
Studies on the Cave and Endogean Fauna of North American II:
Austin, Texas Memorial Museum, Speleological Monograph 3,
p. 127–153.
Muchmore, W.B., and Pape, R.B., 1999, Description of an eyeless,
cavernicolous Albiorix (Pseudoscorpionida: Ideoroncidae) in Arizona,
with observations on its biology and ecology: The Southwestern
Naturalist, v. 44, no. 2, p. 138–147.
Murie, O.J., 1974, Field Guide to Animal Tracks, second ed.: Boston,
Houghton Mifflin, 375 p.
Pape, R.B., Thomas, D.B., and Aalbu, R.L., 2007, A revision of the genus
Eschatomoxys Blaisdell (Tenebrionidae: Pimeliinae: Edrotini) with
notes on the biology: The Coleopterist’s Bulletin, v. 61, no. 4,
p. 519–540. doi:10.1649/0010-065X(2007)61[519:AROTGE]2.0.CO;2.
Peck, S.B., 1980, Climatic change and the evolution of cave invertebrates
in the Grand Canyon, Arizona: NSS Bulletin, v. 42, no. 3, p. 53–60.
Poglayen-Neuwall, I., and Toweill, D.E., 1988, Bassariscus astutus:
American Society of Mammalogists, Mammalian Species, no. 327, 8 p.
Radovsky, F.J., and Krantz, G.W., 2003, Generic and specific synonymy
of Mitonyssoides stercoralis Yunker, Lukoschus, and Giesen, 1990
with Coprolactistus whitakeri Radovsky and Krantz, 1998 (Acari:
Mesostigmata: Macronyssidae): Journal of Medical Entomology,
v. 40, no. 4, p. 593–594. doi:10.1603/0022-2585-40.4.593.
Ribera, C., and Juberthie, C., 1994, Araneae, in Juberthie, C., and Decu,
V., eds., Encyclopaedia Biospeologica, Vol. 1, Moulis - Bucharest,
Societe de Biospeologie, p. 197–214.
Shear, W.A., Taylor, S.J., Wynne, J.J., and Krejca, J.K., 2009, Cave millipeds
of the United States VIII. New genera and species of polydesmidan
millipeds from caves in the southwestern United States (Diplopoda,
Polydesmida, Macrosternodesmidae): Zootaxa, no. 2151, p. 47–65.
Sorenson, E.B., and Stones, R.C., 1959, Description of a new tenebrionid
(Coleoptera) from Glen Canyon, Utah: The Great Basin Naturalist,
v. 19, no. 2 and 3, p. 63–66.
Taylor, W.P., 1954, Food habits and notes on life history of the ring-tailed
cat in Texas: Journal of Mammalogy, v. 35, no. 1, p. 55–63.
Toweill, D.E., and Teer, J.G., 1977, Food habits of ringtails in the
Edwards Plateau Region of Texas: Journal of Mammalogy, v. 58,
no. 4, p. 660–663.
Vandel, A., 1965, Biospeleology – The Biology of Cavernicolous Animals:
London, Pergamon Press, 524 p.
Van Devender, T.R., 1990, Late Quaternary vegetation and climate of the
Sonoran Desert, United States and Mexico, in Betancourt, J.L., Van
Devender, T.R., and Martin, P.S., eds., Packrat Middens – The Last
40,000 years of Biotic Change: Tucson, Arizona, University of
Arizona Press, p. 134–165.
Welbourn, C.W., 1978, Preliminary report on the cave fauna: in Cave
Resources of Horseshoe Mesa (Grand Canyon National Park):
Yellow Springs, Ohio, Cave Research Foundation, p. 36–42.
Wurster, C.M., Patterson, W.P., McFarlane, D.A., Wassenaar, L.I.,
Hobson, K.A., Beavan Athfield, N., and Bird, M.I., 2008, Stable
carbon and hydrogen isotopes from bat guano in the Grand Canyon,
USA, reveal Younger Dryas and 8.2 ka events: Geology, v. 36, no. 9,
p. 683–686. doi:10.1130/G24938A.1.
Wynne, J.J., and Drost, C., 2009, Southwestern caves reveal new forms of
life: U.S. Geological Survey Fact Sheet 2009-3024: http://pubs.usgs.
gov/fs/2009/3024/fs2009-3024.pdf (accessed May 9, 2012).
Wynne, J.J., Drost, C.A., Cobb, N.S., and Rihs, J.R., 2007, Cave-
dwelling invertebrate fauna of Grand Canyon National Park,
Arizona: in Proceedings, 8
Biennial Conference of Research on
the Colorado Plateau: Tucson, University of Arizona Press,
p. 235–246.
Journal of Cave and Karst Studies, April 2014 N13
... Records of troglophilic (cave-dwelling) invertebrates from various cave systems globally indicate that Tineidae are widely present, particularly in tropical and subtropical regions of the Americas as well as the Balkan states and Australia (Barr and Reddell, 1967;Hamilton-Smith, 1967;Peck, 1975Peck, , 1974Robinson, 1980;Trajano, 2000;Humphreys and Eberhard, 2001;Polyak, 2004, 1996;László, 2004;Wynne and Pleytez, 2005;Polak et al., 2012;Byun et al., 2014;Eberhard et al., 2014;Pape, 2014;Silva and Ferreira, 2015;Turbanov et al., 2016;Jakšić, 2017). The Tineidae is a cosmopolitan lepidopteran family (Slootmaekers, 2013), and so it is highly likely that tineids are present in cave systems globally, but records are lacking. ...
Full-text available
Bats and moths provide a textbook example of predator-prey evolutionary arms races, demonstrating adaptations, and counter adaptations on both sides. The evolutionary responses of moths to the biosonar-led hunting strategies of insectivorous bats include convergently evolved hearing structures tuned to detect bat echolocation frequencies. These allow many moths to detect hunting bats and manoeuvre to safety, or in the case of some taxa, respond by emitting sounds which startle bats, jam their biosonar, and/or warn them of distastefulness. Until now, research has focused on the larger macrolepidoptera, but the recent discovery of wingbeat-powered anti-bat sounds in a genus of deaf microlepidoptera (Yponomeuta), suggests that the speciose but understudied microlepidoptera possess further and more widespread anti-bat defences. Here we demonstrate that wingbeat-powered ultrasound production, likely providing an anti-bat function, appears to indeed be spread widely in the microlepidoptera; showing that acoustically active structures (aeroelastic tymbals, ATs) have evolved in at least three, and likely four different regions of the wing. Two of these tymbals are found in multiple microlepidopteran superfamilies, and remarkably, three were found in a single subfamily. We document and characterise sound production from four microlepidopteran taxa previously considered silent. Our findings demonstrate that the microlepidoptera contribute their own unwritten chapters to the textbook bat-moth coevolutionary arms race.
... 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. The most recent undertaking of an invertebrate inventory took place in 2017 and surveyed three stream caves (Zara Environmental 2017). ...
... Therefore, considering (i) that dermestids rarely colonise underground remains and hence their occurrence would likely be due to have been transported into the den together with vertebrate remains, and (ii) that it is highly unlikely that dermestids would colonise completely skeletonised remains unless dried tissues (as required for feeding) are still available, it can be safely assumed that the observed dermestid modifications on bone are contemporaneous with the cave hyena deposition. It must also be mentioned that several studies also recorded the presence of dermestids feeding of bats carcasses and guano within caves occasionally burrowing older underlying bone remains (Pape, 2014;Wrobel and Biggs, 2018). This scenario could suggest a different explanation behind the insects' colonisation of CM and, most important, imply surface modifications on bones occurred anytime following deposition. ...
Cava Muracci (Cisterna di Latina, central Italy) is a complex of seven karst caves located in the Pontine Plain region. One of them, denominate Area 3, displayed a well-preserved Late Pleistocene deposit dating from 35 ka BP and older containing abundant mammal remains and cave hyena coprolites. This work presents the recent discovery from this deposit of fossil bones exposing dermestid pupal chambers and the opportunity these offered to support and improve, with high-resolution inferences, a previous environmental reconstruction based on faunal assemblage and pollen from coprolites. Moreover, ichnological analyses also shed light on some of the post-depositional agents which affected remains within the cave. This paper encourages the valuable support of palaeoichnology, a little exploited and undervalued discipline, to archaeological and palaeoenvironmental studies.
... Only the best cliff climbers (mountain goats, packrats, ringtails) and fliers (bats) gain entrance to some of these caves. With all the caves in the GC, studies of extant and ancient bats and their guano (dung) deposits are abundant (e.g., Wurster et al. 2008;Pape 2014), with a number of mummified remains beginning to be studied in detail (e.g., Mead and Mikesic 2001; see Double Bopper Cave below). A number of medium to large mammals (some extinct) are reported from various caves throughout the GC region, both within the river corridor and above the Tonto Platform mid-canyon, including Shasta ground sloth (Nothrotheriops shastensis), camel (Camelops sp.)., Harrington's mountain goat (Oreamnos harringtoni), bighorn (Ovis canadensis), bison (Bison sp.), and horse (Equus sp.) . ...
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Full citation: Mead, J.I., J.S. Tweet, V.L. Santucci, B. Tobin, C.L. Chambers, S.C. Thomas, and M.C. Carpenter, 2020. Pleistocene/Holocene Cave Fossils from Grand Canyon National Park: Ice Age (Pleistocene) Flora, Fauna, Environments, And Climate Of The Grand Canyon, Arizona. in Santucci, V.L. and J.S. Tweet, (editors), Grand Canyon National Park: Centennial paleontological resource inventory (non-sensitive version). Natural Resource Report NPS/GRCA/NRR—2020/2103. National Park Service, Fort Collins, Colorado, p. 403-463.
... Current literature on the behaviors of meso-mammals in caves generally focuses on single caves, seasons, or consists of observations made secondary to a primary research questions (Elder and Gunier 1981;Pape 2014). Because of this, we can only speculate as to why mammal cave use in central Texas occurs or what constitutes typical or atypical use. ...
... This perhaps suggests that bodies were placed within the cave fairly regularly, providing sustenance to maintain the colony. Alternatively, several cave ecology studies from the New World and Asia have reported dermestids in association with bats, feeding primarily on their carcasses and on guano (Gnaspini & Trajano, 2000;Mitchell & Reddell, 1971;Mizutani, McFarlane, & Kabaya, 1992;Pape, 2014). Thus, evidence of infestation within Je'reftheel does not necessarily imply the presence of dermestids at the time of deposition of fleshed bodies. ...
Human bones from the Maya mortuary cave of Je’reftheel in west‐central Belize show evidence of taphonomic modifications attributed to insects, with termites and dermestid beetles being the most likely culprits. This study represents the first detailed exploration of the effects of osteophageous insects on bones from the Maya area, and thus expands on recent efforts by other researchers working in the region to document taphonomic processes and distinguish them from intentional mortuary treatments.
... Many of these caves have multiple miles of passage surveyed and have a variety of resources present. Through this documentation, significant resources have been found throughout the caves, from archeology and paleontology (Henderek et al. 2015) to biology (Pape 2014) to hydrology (Jones et al. 2017). Previous research has shown that these types of resources are often rare, delicate, and vulnerable to human disturbance (Despain and Fryer 2002). ...
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Protected areas are tasked with mitigating impacts to a wide range of invaluable resources. These resources are often subject to a variety of potential natural and anthropogenic impacts that require monitoring efforts and management actions to minimize the degradation of these resources. However, due to insufficient funding and staff, managers often have to prioritize efforts, leaving some resources at higher risk to impact. Attempts to address this issue have resulted in numerous qualitative and semi-quantitative frameworks for prioritization based on resource vulnerability. Here, we add to those methods by modifying an internationally standardized vulnerability framework, quantify both resource vulnerability, susceptibility to human disturbance, and fragility, susceptibility to natural disturbance. This modified framework quantifies impacts through a six-step process: identifying the resource and management objectives, identifying exposure and sensitivity indicators, define scoring criteria for each indicator, collect and compile data, calculate indices, and prioritize sites for mitigations. We applied this methodology to two resource types in Grand Canyon National Park (GRCA): caves and fossil sites. Three hundred sixty-five cave sites and 127 fossil sites in GRCA were used for this analysis. The majority of cave and fossil sites scored moderate to low vulnerability (0–6 out of 10 points) and moderate to low fragility for fossils. The percentage of sites that fell in the high-priority range was 5.5% for fossils and 21.9% for caves. These results are consistent with the known state of these resources and the results present a tool for managers to utilize to prioritize monitoring and management needs.
... Around a quarter of global bat species are under threat largely as a consequence of habitat destruction and modification (Kunz and Racey, 1998). The alteration of cave and karst ecosystems represent major drivers of extinction for diverse cave-dependent species (McCracken, 2011;Medellin et al., 2017), which in turn support a widearray of macroinvertebrate species dependent on the organic nutrients from bat guano, respiration, and urination (Pape, 2014;Iskali and Zhang, 2015). Thus, developing standardised and comparable methodologies to develop priorities for management and conservation are crucial to effectively protecting cave biodiversity. ...
The identification of important habitats for wildlife is essential in order to plan and promote strategies for longterm effective conservation. Caves and subterranean habitats are frequently overlooked habitats with diverse communities, which are frequently endemic to a region, karst outcrop or even a single cave. These cave species include a wide range of taxa adapted to cave environments. Within cave systems, bats are key providers of energy for other cave-dependent species. However, identifying caves for conservation prioritisation requires an understanding of cave-dwelling species diversity, patterns of endemism, and conservation status, in addition to a standard mechanism to evaluate risk. In this paper, we present the ‘Bat Cave Vulnerability Index’ (BCVI) as a standard index for evaluating bat caves for conservation prioritisation by determining Biotic Potential (BP) and Biotic Vulnerability (BV) of caves. The Biotic Potential is represented by various species diversity and rarity measurements. The Biotic Vulnerability is represented by the cave geophysical characteristics and human-induced disturbance present. Pilot testing in the southern Philippines has demonstrated that the index is an effective and practicable method to identify bat caves for conservation prioritisation. The biotic potential variables assess the presence of endemic, rare, and threatened bat species and assays the priority level based on an equation. Relative risk and vulnerability were assayed using landscape vulnerability variables, which showed anthropogenic activities were important factors in conservation prioritisation. The application and mechanism of the index potentially provides a valuable, rapid and simple assessment tool in cave conservation with special relevance to bat diversity and vulnerability. Furthermore, the multiple and holistic criteria of the BCVI, and the accessible information for both biotic and landscape features can be adapted to prioritise caves in a wider scale in the tropics, and in other regions with diverse cave ecosystems.
... Though less common in Camp Bullis, ringtails are also known to use caves for feeding, as a water source, and for denning (Clark 1951, Wynne 2013, Pape 2014. Ninebanded armadillo cave use was particularly interesting because they were exclusively photographed at Up the Creek Cave. ...
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Knowledge of meso-mammal cave use is essential for natural resource managers, particularly in the management of endangered cave invertebrates. Scat left by meso-mammals represents significant nutrient inputs into the oligotrophic cave environment which can disrupt the invertebrate species composition and ecology. Since little is known about what constitutes typical timing or frequency of meso-mammal visitation, current management practices are largely speculative. Central Texas caves were historically associated with raccoons Procyon lotor, but with the loss of large predators and encroachment of woody vegetation, the now naturalized North American porcupine Erethizon dorsatum has become an established part of the local ecosystems whose effect on cave biology remains unknown. Our objective with this study was to quantify meso-mammal cave use according to seasons, time of day, weather conditions, as well ecological and physical cave characteristics. We monitored 30 caves by placing trail cameras at cave entrances for one year on Joint Base San Antonio-Camp Bullis military base just north of San Antonio, Texas. North American porcupines, raccoons, and Virginia opossums Didelphis virginiana were the three most commonly photographed meso-mammals (>87%). All meso-mammal groups showed significantly different cave use according to season, weather, and cave characteristics. Data suggested most meso-mammals were using caves for denning while raccoons and Virginia opossums also were feeding on resident invertebrate and rodent populations. In particular, Virginia opossum and raccoon, both potential predators of endangered species, showed greater use of caves containing endangered species. The results from our study represent an initial step in understanding meso-mammal cave use in central Texas.
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(Citation: Bossard, R.L. 2017. Population Model for the Bat Flea Sternopsylla distincta (Siphonaptera: Ischnopsyllidae) in Temperate Caves of the Brazilian Free-Tailed Bat Tadarida brasiliensis (Mammalia: Chiroptera). The Journal of the Utah Academy of Sciences, Arts, & Letters 94: 71-78.) The population dynamics of the bat flea Sternopsylla distincta are not quantitatively known. These fleas parasitize Brazilian free-tailed bats (Tadarida brasiliensis) in temperate caves. Using laboratory and field data, I model seasonal population changes. Bat flea populations are predicted to boom when bats arrive in the spring, and immature flea stages quickly reach equilibrium densities after a few days. From June to September, all stages are present, with the most abundant being eggs at 90 per m2 of guano. Larvae equilibrate at 20 per m2, pupae 1 per m2, and adults comprise only 10% of the summer's flea population. When bats depart for southern roosts in the fall, bat flea populations decline, but pupae survive the winter. These results imply that S. distincta overwinters both as pupae (in northern caves) and as adults (in southern roosts). Fleas are important in many caves as climbing ability and phoresy allow fleas to attain bats on ceilings, creating a "flea–feces loop" of cycling biomass. Fleas reach bats by crawling up cave walls, or when adult female bats retrieve fallen, flea-infested pups, or through uncommon earwig phoresy. The model provides baseline predictions enabling comparison with field data and insight into food limitation that appears to regulate bat-flea populations in temperate caves.
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The Chihuahuan Desert is a sensitive area for paleoclimatic studies because of its moisture-deficient climate and freezing winter temperature. Midden sequences from Maravillas Canyon and Rio Grande Valley and isolated records from six other sites provide a vegetation record for the past 40 000 yr in the Big Bend of Texas. Proxy records are provided by middens of the climatic history of the Chihuahuan Desert for the last 45 000 yr, and four areas yielded both Middle and Late Wisconsinan middens which reflect a latitudinal gradient with the magnitude of biogeographic change decreasing to the south. -S.J.Yates
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
A new species of pseudoscorpion, Albiorix anophthalmus, is described from Arkenstone Cave, Pima Co., Arizona. It is highly modified for life in the cave, being larger and more slender than any other known species in the genus and the only known species without eyes. Descriptions of the epigean (surface) and hypogean (cave) environments are provided. Observations on the biology and ecology of A. anophthalmus also are presented.