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Inventory, conservation, and management of lava tube caves at El Malpais National Monument, New Mexico

Abstract and Figures

Lava tube caves at El Malpais National Monument have received little scientifi c attention with regard to their bat and arthropod populations. From an all taxa biological inventory of 11 caves, I identified seven new species of cave-dwelling arthropods (including two potential troglobites) and range expansions of two parasitoidal wasps. The presence of unique microhabitats, tree root “curtains” hanging from the ceilings, and moss gardens in cave entrances resulted in higher species richness of arthropods at four caves. For bats, I confi rmed continued use of one large bat hibernaculum cave and one signifi cant bat maternity roost. While several recommendations have been made to better conserve and manage sensitive cave resources, additional research and monitoring will be required for the long-term management and protection of several caves. Finally, I introduce three new terms to cave biology: two for entrance-dwelling animals (eisodophiles and eisodoxenes) and one for animals that hunt deep within or near the entrances of caves (xenosylles).
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Inventory, conservation,
and management of lava
tube caves at El Malpais
National Monument, New
Mexico
By J. Judson Wynne
Figure 1. The author searches for arthropods
beneath the skylights of ELMA-012 cave.
BUREAU OF LAND MANAGEMENT/KYLE VOYLES
RESEARCH REPORTS
45
L
OCATED IN WESTERN NEW MEXICO, EL MALPAIS
National Monument encompasses approximately 1,522 km
2
[~588 mi
2
]. Featuring at least eight major volcanic eruptions
ranging in age from 100,000 to 3,000 years old (Cascadden et al.
1997), the national monument is a dramatic landscape comprising
vast expanses of pahoehoe and ʻaʻā lava fl ows, cinder cones, ice
caves, and at least 290 lava tube caves (fi g. 1, previous page). De-
spite the large number of lava tube caves, this region has received
little scientifi c attention with regard to bat and arthropod popula-
tions that occur within these features.
Bats are often considered keystone species of cave ecosystems.
When bats populate caves in large numbers, they transport a sig-
nifi cant amount of organic material (as guano) from the surface
into the cave. Although bats have been studied throughout most
of the western United States, how these animals use caves remains
underresearched. Bat maternity roosts (sites where female bats
rear their pups) and hibernacula (winter hibernation sites) are
highly sensitive to human disturbance (Brown et al. 1993; Mc-
Cracken 1989; Elliott 2000; Hamilton-Smith and Eberhard 2000).
With the westward advance of white -nose syndrome, a disease
responsible for the mortality of more than fi ve million bats in
eastern North America (USFWS 2012), inventory and monitor-
ing of all roost sites will be critically important to the long-term
management of bats at El Malpais National Monument.
Other animals of high conservation and management value are
arthropods that occur exclusively in caves. Prior to this study, at
least fi ve cave-adapted arthropods (presumed sensitive species)
were known from six lava tube caves at El Malpais (Northup and
Welbourn 1997). Many troglomorphic (cave-adapted) animals
are endemic to a single cave or region (Reddell 1994; Culver et al.
2000; Christman et al. 2005) and are generally characterized by
low population numbers (Mitchell 1970). Additionally, numerous
human-induced impacts threaten subterranean ecosystem health
and the very persistence of cave-obligate species. Many cave-
obligate species are therefore considered imperiled. Nonnative
species introductions (Elliott 1992; Reeves 1999; Taylor et al. 2003;
Howarth et al. 2007), global climate change (Chevaldonné and
Lejeune 2003; Badino 2004), and recreational use (Culver 1986;
Howarth and Stone 1993; Pulido-Bosch et al. 1997) are among the
impacts that present challenges for the long-term management of
cave-obligate arthropod populations at El Malpais.
An all taxa biological inventory focusing on bats, cave-dwelling
arthropods, and other vertebrates was not only important to
characterizing the fauna that use El Malpais lava tubes, but also
was required to provide resource managers with the information
necessary to best conserve and manage these sensitive resources.
My objectives for this study were to (1) catalog all taxa using caves,
including the identifi cation of endemic and sensitive cave-
adapted invertebrates, (2) apply and examine a systematic sam-
pling protocol for inventorying arthropods, (3) draw comparisons
across the national monument to gain inference into patterns of
invertebrate species distributions, biodiversity, biogeography,
and endemism, and (4) provide recommendations to enhance
management of El Malpais lava tube caves. I addressed objectives
1 and 4 in this article and will address objectives 2 and 3 in subse-
quent publications.
Methodology
During 7–15 October 2007 and 8–15 October 2008, research teams
and I conducted two site visits per cave at 10 caves at El Malpais
National Monument and 1 cave on adjacent Bureau of Land
Management lands. We scheduled site visits around deployment
and collection of baited pitfall traps for sampling cave-dwelling
arthropods. At the monument’s request, I used cave codes rather
than actual cave names for all caves on National Park Service
lands. A copy of this report, which includes a table of cave names
with associated cave codes, is on fi le with monument headquar-
Ab
st
ra
ct
Lava tube caves at El Mal
p
ais National Monument have received
little scienti c attention with regard to their bat and arthropod
populations. From an all taxa biolo
g
ical inventor
y
of 11 caves,
I identifi ed seven new species of cave-dwelling arthropods
(including two potential troglobites) and range expansions of two
parasitoidal wasps. The presence o
f
unique microhabitats, tr
ee
root “curtains” hanging from the ceilings, and moss gardens in
cave entrances resulted in higher species richness of arthropods
at four caves. For bats, I confi rmed continued use of one large bat
hibernaculum cave and one signi cant bat maternity roost. While
several recommendations have been made to better conserve
and mana
e sensitive cave resources, additional research and
monitorin
g
will be required
f
or the lon
g
-term mana
g
ement and
protection o
f
several caves. Finall
y
, I introduce three new terms
to cave biolo
gy
: two for entrance-dwellin
g
animals (eisodophiles
and eisodoxenes) and one
f
or animals that hunt dee
p
within or
near the entrances of caves (xenos
y
lles).
Ke
y
word
s
cave, cave biolo
gy
, cave-dwellin
g
arthropods, cave-roostin
g
bats, conservation, eisodo
p
hile, eisodoxene, El Mal
p
ais National
Monument, land mana
g
ement, lava tubes, xenos
y
ll
e
PARK SCIENCE • VOLUME 30 • NUMBER 1 • SUMMER 2013
46
Figure 2. Both moss gardens (top row) and root curtains (bottom
row) are important microhabitats and support new and unique
arthropod species. At least three lava tube caves contain moss
gardens within cave entrances and beneath skylights, and two caves
contain plant root curtains within cave deep zones at El Malpais
National Monument.
BUREAU OF LAND MANAGEMENT/KYLE VOYLES
entrances and beneath skylights (i.e., holes in the ground formed
by the partial collapse of the cave roof), and tree root “curtains”
hanging from the ceilings in cave deep zones (fi g. 2). Within each
unique microhabitat we spent one hour (3 observers × ~20 min-
utes) searching for arthropods. Specifi cally, we searched tree root
curtains hanging from the ceilings in two caves (one hour per
cave) and moss gardens in two caves (one hour per cave).
Arthropod identifi cation
For arthropod groups actively being studied by taxonomic
specialists, I sent either specimens or high-resolution images of
specimens to various taxonomic experts for identifi cation. Oth-
erwise, we used existing keys to identify specimens to the lowest
taxonomic level possible.
Bat sampling
I visited and inventoried three known bat roosts: a Mexican free-
tailed bat (Tadarida brasiliensis) maternity colony, a Townsend’s
big -eared bat (Corynorhinus townsendii) maternity roost, and a
Townsend’s big- eared bat hibernaculum. Additionally, for sites
where bat use was unknown, these caves were surveyed for
midfall use by bats. Research teams scanned ceilings and walls
throughout the length of each cave, and searched for any evi-
ters in Grants, New Mexico, and the National Cave and Karst
Research Institute, Carlsbad, New Mexico.
Arthropod sampling
I used both opportunistic and systematic sampling to search for
arthropods. During each cave visit, a team of three researchers
uniformly applied three techniques: opportunistic collecting,
baited pitfall trapping, and timed searches. For opportunistic
collecting, the team collected invertebrates encountered as they
walked between sampling stations while deploying and removing
pitfall traps and conducting timed searches.
Because I wanted to maximize the number of invertebrate species
detected, we sampled each cave from its entrance (i.e., drip line)
to the back of the cave. Using available cartographic cave maps,
teams applied an interval sampling approach whereby 10% of
each cave’s length was used as the sampling interval (e.g., for a
100 m–long cave [328 ft], sampling interval was every 10 m [33
ft]). All sampling stations were plotted on each cave map. Three
sampling stations (one at either wall and one at the cave center-
line) were established at each sampling interval. Fewer than three
sampling stations per sampling interval were established in only
two cases: (1) when the cave passageway width was ≤5 m (16 ft),
one station was designated in the best available location and (2)
when exposed lava fl oors were encountered and no materials
were available to aid in countersinking the trap, the sampling sta-
tion was skipped.
At each sampling station we deployed one pitfall trap and con-
ducted two timed searches. Traps consisted of two 907 g (32 oz)
plastic containers (13.5 cm high, 10.8 cm diameter rim, and 8.9 cm
base [5.3 in high, 4.3 in diameter, 3.5 in base]) placed inside one
another, with bait (a teaspoon of peanut butter) placed in the
outer container and holes punched in the base of the inner con-
tainer. This design attracts arthropods and keeps most animals
separated from the bait. With the assistance of fi eld technicians, I
buried containers to the rim where possible, built rock ramps to
the trap rim in other cases, and covered all traps with a caprock.
The team conducted timed searches within a 1 m (3.3 ft) radius of
each pitfall trap station for a period of 1 to 3 minutes before traps
were deployed and prior to checking traps (modifi ed from Peck
1976). Each search was concluded after 1 minute if no arthropods
were detected and continued for a total of 3 minutes when arthro-
pods were observed.
Because some caves contain unique microhabitats that support
distinct arthropod communities and endemic populations, I
augmented this sampling protocol by conducting direct intuitive
searches in those areas. Unique microhabitats at El Malpais in-
clude moss gardens (refer to Northup and Welbourn 1997) at cave
RESEARCH REPORTS
47
Coleoptera (8)
Collembola (4)
Diplopoda (2)
Diplura (1)
Diptera (4)
Hemiptera (1)
Hymenoptera (4)
Lepidoptera (9)
Orthoptera (3)
Psocoptera (1)
Siphonaptera (1)
Araneae (14)
Opilliones (1)
Acari (5)
Chilopoda (1)
Figure 3. Number of arthropod
morphospecies detected
by order (including classes
Chilopoda and Diplopoda) at
El Malpais National Monument
in 2007 and 2008. The surveys
were conducted at 11 caves:
ELMA-062, ELMA-008, ELMA-
110, ELMA-262, ELMA-012,
ELMA-054, ELMA-029, ELMA-
303, ELMA-315, ELMA-061, and
Hummingbird.
dence of bats (e.g., guano). When bats were encountered,
I attempted to identify them to species visually. No bats were
handled during this study.
Documenting other vertebrates
Within each cave, I searched for and recorded the presence of all
other vertebrate species. Sign of other vertebrates included direct
observation, scat, feathers, and skeletal remains.
Cave specifi city functional groups
I divided El Malpais cave-dwelling taxa into nine cave specifi city
functional groups. The following functional group terminology
was taken from Barr (1968) and Howarth (1983): (1) troglobites,
obligate cave dwellers who can only complete their life cycle
within the cave environment; (2) troglophiles, species that occur
facultatively within caves and complete their life cycles there, but
also exist in similar surface microhabitats; (3) trogloxenes, taxa
that frequently use caves for shelter but forage on the surface;
and (4) accidentals, morphospecies occurring within caves, but
which cannot survive within the cave environment. Additionally,
because this project sampled cave entrances for arthropods and
documented other vertebrate (i.e., non-bat taxa) use of caves, I
propose three additional groups for categorizing cave-dwellers: (5)
eisodophiles,species facultatively using cave entrances and twilight
zones (areas where light faintly penetrates into the cave, but is
suffi cient for humans to see) that may complete their life cycles
there, but also occur in similar partially sheltered surface environ-
ments; (6) eisodoxenes,animals that frequently use cave entrances
and twilight zones for shelter but return to the surface to forage;
and (7) xenosylles, surface-dwelling animals that hunt deep within
caves or in the cave entrances. For eisodophiles and eisodoxenes,
the etymology of the fi rst half of the terms, eísodo,is from the
Greek word eísodos, “entrance,” while the second halves were
derived from the same naming convention used for functional
groups 2 and 3: philos, Greek for “love” or “desire,” and xenos,
Greek for “guest.” Xenosylle is a combination ofxenosandsyl-
léktis, Greek for “collector.” Finally, (8) parasites, the special-case
group, describes parasitic arthropods detected in caves due to
the presence of their host (e.g., bats or other vertebrates); and (9)
unknown is used for animals for which information is lacking to
reasonably place them in one of the eight other groups.
Results
My work resulted in the identifi cation of at least 66 morphospe-
cies (groups distinguished from others based upon morphological
characteristics), including 59 arthropods (representing at least 13
orders and two classes; fi g. 3), three bats, and four other verte-
brates. Appendix A, available online at http://www.nature.nps.gov
/ParkScience/archive/PDF/Article_PDFs/ParkScience30(1)
Summer2013_A1–A12_Wynne_3653.pdf, shares the entire list of
inventoried species and provides explanations for cave specifi city
functional group designations.
Arthropods
Cave specifi city functional groups for arthropods consisted of
one troglobite, two questionable troglobites, fi ve troglophiles, 18
PARK SCIENCE • VOLUME 30 • NUMBER 1 • SUMMER 2013
48
Figure 5. (A) New species of a potentially troglobitic planthopper
(order Hemiptera: superfamily Fulgoroidea: Fulgoroidea n.sp.?; body
length ~1.5 mm); (B) new species of troglobitic bristletail (order
Diplura: family Campodeidae; from Northup and Welbourn 1997;
length 2.5 mm); (C) new species of trogloxenic Carabid beetle
(Rhadine n.sp., perlevis species-group; length 15 mm); and (D) new
species of trogloxenic cave cricket (Ceuthophilus cf apache n.sp.;
length 25 mm).
questionable troglophiles, seven trogloxenes, one questionable
trogloxene, one accidental, 21 eisodophiles, one parasite, and two
unknowns (fi g. 4). At least seven new species were discovered and
two range expansions were documented. New species discoveries
include one potentially cave-adapted spider (family Theridiidae,
Theridion n.sp.?); a mite (family Histiostomatidae, Histiostoma
n.sp.); two springtails (order Collembola, Drepanura n.sp. and
Pogonognathellus n.sp.); one new cricket species (Ceuthophilus
cf apache n.sp.); one beetle (family Carabidae, Rhadine n.sp.,
perlevis species-group); and a new species of potentially cave-
adapted planthopper (order Hemiptera: superfamily Fulgoroidea;
Fulgoroidea n.sp.?; refer to fi g. 5 for images of select new species).
Additionally, I confi rmed the persistence of the troglomorphic
bristletail (order Diplura: family Campodeidae; Campodeidae
n.sp.) within the deep zone of ELMA-054. Northup and Wel-
bourn (1997) identifi ed this as both a troglobite and an “unde-
termined species.” Working with dipluran taxonomist Dr. R.
Thomas Allen (Academy of Natural Sciences, Drexel University,
Philadelphia, Pennsylvania), we confi rmed this animal as a new
species in 2013. The likely new species of planthopper was de-
tected within the cave deep zones on roots protruding from the
ceiling of ELMA-315 and ELMA-303; this animal has reduced eyes
in its nymphal stage and may be troglomorphic. Also, this work
resulted in the range expansions of two species of parasitoidal
wasps (fi g. 6, next page) (family Tiphiidae, Tiphia andersoni and
T. nona; Allen 1971). Both tiphiids were in a torpor and collected
from beneath rocks within the moss gardens of ELMA-008 and
ELMA-012. Given the season (midfall) and their behavior, I sug-
gest these wasps may have been preparing to hibernate within the
moss gardens.
Figure 4. Cave speci city
functional groups for
arthropods, bats, and other
vertebrates at El Malpais
National Monument.
Troglobite (1)
Questionable Troglobite (2)
Eisodophile (21)
Accidental (1)
Trogloxene (7)
Unknown (2)
Troglophile (5)
Questionable
Troglophile (18)
Parasite (1)
Questionable Trogloxene (1)
Trogloxene (3)
Unknown (1)
Questionable xenosylle (1)
Eisodophile (1)
Trogloxene (1)
Bats
Other Vertebrates
RESEARCH REPORTS
49
A B
C D
A, B, AND D: NORTHERN ARIZONA UNIVERSITY/JUT WYNNE; C: BUREAU OF LAND MANAGEMENT/KYLE VOYLES
Figure 6.
Tiphia andersoni
Allen, 1971. This specimen was collected via
a direct intuitive search of moss gardens (beneath the large skylights)
of ELMA-012. Prior to this work, this parasitoidal wasp was known
to occur in central Mexico, north into southeastern and north-
central Arizona (Allen 1971). Detection of this animal in New Mexico
represents an expansion of its known range.
NORTHERN ARIZONA UNIVERSITY/JUT WYNNE
Caves with the highest arthropod species richness, in rank order,
were ELMA-315 (n = 22), ELMA-012 (n = 16), ELMA-303 (n = 15),
ELMA-008 (n = 15), ELMA-062 (n = 13), and ELMA-262 (n = 11)
(table 1). ELMA-315 and ELMA-303, which had the highest spe-
cies richness, contain extensive root curtains protruding through
ceiling fi ssures within the cave deep zones. For ELMA-012 and
ELMA-008, richness is driven by the large number of species
detected within moss gardens at cave entrances and beneath
skylights. ELMA-062 supports a large Mexican free-tailed bat
maternity roost; because signifi cant nutrients via guano have been
transported into this cave, this likely contributed to the high num-
ber of morphospecies. In 2007, logistical constraints prevented
my team from sampling the moss gardens within the entrance of
ELMA-029 and from further sampling ELMA-110 (which sup-
ports a bat maternity roost). Thus, I suggest both of these caves
likely support more arthropods (in terms of richness and abun-
dance) than are included in this report.
All new species reported here were identifi ed as “new” by taxo-
nomic specialists. Several of these new species ultimately will be
formally described and the results published in scientifi c journals.
Bats
During the 2007 surveys, I observed fi ve hibernating Townsend’s
big-eared bats in the deep zone of ELMA-054 and one torpid
big brown bat (Eptesicus fuscus) roosting near the entrance
(fi gs. 7A and 7B, respectively). Additionally, ELMA-062 contin-
ues to support a maternity colony of Mexican free-tailed bats.
On 10 October 2007, I observed thousands of individuals of this
species roosting approximately 75 m (246 ft) within the cave
(fi g. 7C); however, when I returned four days later to remove
arthropod traps, that number dropped to less than 100. Also in
October 2007, I did not observe any Townsend’s big-eared bats
in residence at ELMA-110; once I arrived, this roost was already
abandoned for the year. Relatively fresh guano in the main section
of the cave (beginning at the northeasternmost skylight, extend-
ing to the northeastern ward) suggests they were using this area
before they relocated to their winter roosts. During an unrelated
study in 2005 and 2006, I observed a maternity roost of this spe-
cies in both the main cave and tunnel segments directly southwest
of the main section of this cave. It seems the colony uses several
areas in the tunnel segments and within the main cave passageway
during the breeding season. Given the sampling period in early
October (after breeding), I was unable to ascertain whether or
not additional summer roosts exist on the national monument.
However, aside from the two known maternity roosts, I did not
observe any signifi cant deposits of fresh guano (suggestive of a
large summer roost) in any of the other caves. Thus, I have no
evidence to suggest additional large summer roosts occur in the
caves sampled. All bats were considered trogloxenes.
Other vertebrates
I documented small-carnivore (questionable xenosylle) scat,
likely ringtail (Bassariscus astutus), skunk (Conepatus sp.), or
raccoon (Procyon lotor), in ELMA-054 and ELMA-110. Skunks
and raccoons often prey upon infi rm bats or bat pups that have
fallen from the ceiling (Winkler and Adams 1972), and ringtails are
commonly known to hunt bats roosting on cave walls. I found a
fully articulated ringtail skeleton near the terminus of the north-
ern extent of ELMA-303. I sent photographs of the skeleton to
Eastern Tennessee State University paleontologist Dr. Jim Mead.
In an e-mail exchange with him on 4 March 2013, he suggested
the remains were between 1,000 and 10,000 years old. This animal
may have entered the cave to hunt bats, became disoriented,
and, unable to fi nd its way back to the entrance, died in the cave.
Given its age, I did not consider this animal’s remains as part of
this inventory. Recent packrat (Neotoma sp.; N.mexicana and/or
N.albigula; refer to Bogan et al. 2007) activity was evident in both
ELMA-062 and ELMA-061; packrats are considered trogloxenes.
Also, I found the carcass of a gopher snake (Pituophis catenifer) in
the twilight zone of ELMA-061. The snake was wrapped around
PARK SCIENCE • VOLUME 30 • NUMBER 1 • SUMMER 2013
50
Figure 7. (A) Hibernating Townsends big-eared bat and (B) big
brown bat aroused from torpor at ELMA-054. (C) Late-season
maternity colony of Mexican free-tailed bats within ELMA-062. Note
that the “rough” surface in this panel is actually tightly clustered
roosting bats. For scale, the average wingspan of Mexican free-
tailed bats is 301 mm (12 in) (refer to Wilkins 1989).
NORTHERN ARIZONA UNIVERSITY/JUT WYNNE
a stick and had several lacerations along the length of its body; I
suggest a park visitor probably killed this animal. Because I do not
know whether the snake was killed in the cave or it was brought
into the cave postmortem, its use of the cave is “unknown.” Final-
ly, a barn owl (Tyto alba; eisodophile) was spooked as my team
entered ELMA-262. This owl was roosting near the entrance and
then fl ew to a skylight where it exited the cave.
Conservation and management
This work resulted in the identi cation of seven new species of
cave-dwelling arthropods (including two potential troglobites),
range expansions of two parasitoidal wasps, and two caves con-
taining signifi cant root curtains hanging from the ceiling. The
presence of root curtains and moss gardens has been shown to
be an important driver of high arthropod richness at El Mal-
pais lava tube caves. For bats, I confi rmed continued use of one
hibernaculum cave and two signifi cant bat maternity roosts.
Although all of these caves will require further study, these fi nd-
ings have been useful in highlighting future management direc-
tions and research needs.
Arthropods
Four of the new species reported here are dependent on caves
for most or all of their life cycle. The potentially new troglomor-
phic spider (Theridion n.sp.?) and the planthopper (Fulgoroidea
n.sp.?) are likely to be restricted to the cave environment, while
the cricket (Ceuthophilus cf apache n.sp.) and beetle (Rhadine
n.sp., perlevis species-group) are trogloxenes. Unfortunately,
only one specimen of Theridion n.sp.? was detected and col-
lected; additional specimens will be required to describe this
animal and determine whether it is indeed troglomorphic. In the
two caves with root curtains, I identifi ed at least one potential
troglobite, a planthopper (Fulgoroidea n.sp.?). Unfortunately,
all specimens collected were nymphs, and adults are required
to confi rm both cave adaptation and whether or not it is a new
species. The remaining three newly discovered species likely
occur in surface habitats as well as caves. The two new springtail
species (Drepanura n.sp. and Pogonognathellus n.sp.) are edaphic
Table 1. Observed morphospecies richness for arthropods,
bats, and other vertebrates at El Malpais National
Monument caves
Cave Arthropods Bats
Other
Vertebrates
μ
ELMA-008 15
μ
ELMA-012 16
ELMA-029 —
β
ELMA-054 2 2 1
ELMA-061 1 2
β
ELMA-0621311
β
ELMA-110 4 1
μ
ELMA-262 11 1
ρ
ELMA-303 15
ρ
ELMA-315 22
Hummingbird 3
Notes: Some species were detected in two or more caves.
μ
Moss gardens occurred beneath skylights and within entrances of these caves.
β
Confirmed bat roosts.
ρ
Caves with extensive root curtains protruding through the ceiling within the deep zone.
RESEARCH REPORTS
51
A B
C
(soil-dwelling) organisms. Histiostoma n.sp. is a very small mite
(600–900μm [0.6–0.9 mm] in length) and is found in association
with other insects. Because the deutonymph (early life stage of
mites) hitchhikes on larger-bodied insects for dispersal between
habitats, this mite may have been transported into the cave by an-
other animal. Both springtails and mites will require further study
to determine their affi nities for caves and the ecological roles they
play in the cave environment.
To address questions concerning population dynamics and
distribution patterns of these new arthropod species, additional
surveys at caves known or likely to support these animals will be
required. This information will be necessary to develop resource
management plans to best protect these species and their habitats.
All of these new species should be considered important fi nds
because they expand our knowledge of the natural history of El
Malpais National Monument and, by extension, the state of New
Mexico.
Bats
ELMA-054 supports the largest known hibernaculum of
Townsend’s big-eared bats on the monument, while ELMA-110
supports the largest known maternity roost of this species. Wynne
(2006) counted 100 Townsend’s big-eared bats hibernating in the
deepest section of ELMA-054. ELMA-110 supports a maternity
roost of Townsend’s big-eared bats, estimated at 50 individuals in
2006 (Wynne, unpublished data). ELMA-110 has been closed to
park visitors for several years while bats are in residence. As a re-
sult of this study and the 2006 site visit, ELMA-054 is now closed
during the hibernation period (October through mid-April).
Based on our knowledge of Townsend’s big-eared bats in other
areas, I suggest the same population uses both roosts. In Okla-
homa, movements of these bats between maternity roosts and hi-
bernacula averaged 11.6 km (7.2 mi) (range 3.1 to 39.7 km [1.9–24.7
mi], n = 3 individuals; Humphrey and Kunz 1976). Dobkin et al.
(1995) documented Townsend’s big-eared bats traveling distances
ranging from 5 to 24 km (3–15 mi) from summer roost to foraging
sites in Oregon. Additionally, Pierson et al. (1999) suggested that
this species was in decline throughout its range. The straight-line
distance from ELMA-110 to ELMA-054 is 10.5 km (6.5 mi).
Given that this species is likely to be the most aff ected by white-
nose syndrome on the monument, knowledge of this bat’s habits,
movements, and roost locations will enhance its management and
protection. I recommend conducting a radio tagging and telem-
etry study of Townsend’s big-eared bats and their use of these
two roosts. For such a project, radio tagging of bats should occur
late in the maternity season (late August to early September).
This project would enable monument personnel to (1) establish
baseline estimates of population size and structure to begin
monitoring this species and its two known roosts, (2) determine if
the same population is using both ELMA-110 and ELMA-054, (3)
potentially identify additional Townsend’s big-eared bat roosts by
tracking bat movements with telemetry, and (4) make informed
decisions regarding potential cave closures and protection of this
species.
Scientists and managers know little about the winter habitat
requirements of year-round bat residents at El Malpais. Thus,
more surveys are needed, particularly winter bat inventories, to
identify additional hibernacula. I recommend annual to biennial
monitoring of ELMA-054, as well as newly identifi ed hibernacula
and long-term microclimatic monitoring in caves supporting
hibernating bats. In light of the westward advance of white-nose
syndrome and global climate change, this information may be
useful in guiding management decisions to protect bat popula-
tions in the future. Additionally, information gathered by such
an endeavor may be informative for developing similar monitor-
ing strategies for other management units of the National Park
System in the southwestern United States.
Deep zones and unique habitats
All deep zone environments that support or have the potential to
support cave-adapted animals should be considered high-priority
sites for conservation and management. Deep zones are charac-
terized where environmental conditions (e.g., complete dark-
ness, temperature, relative humidity, moisture, airfl ow) remain
relatively stable over time (refer to Howarth 1980 and 1982). When
nutrients are added to this equation (via root curtains protruding
from the ceiling, bat guano, or dissolved organic material perco-
lating through rock), these areas should be intensively sampled
for troglomorphic arthropods. For example, Howarth et al.
(2007) stressed the importance of roots in caves for conserving
troglomorphic arthropods in Hawaiian lava tubes.
Three caves on the monument meet these criteria. ELMA-315
and ELMA-303 contain deep zones with extensive root curtains
hanging from the ceiling. During the arthropod sampling period,
these caves were among the warmest on the monument (ELMA-
315: mean temperature 12.4°C [54.3°F], standard deviation 0.5°C
[0.9°F]; ELMA-303: mean temperature 11.9°C [53.4°F], standard
deviation 0.7°C [1.3°F]). I know of no other caves in the region
that support this microhabitat type. Additionally, ELMA-110
has the most extensive deep zone microhabitat known on the
monument. At the terminus of this cave, water percolates through
ssures into the cave chamber. I recommend conducting addi-
tional surveys in all of these caves using a bait sampling and direct
intuitive search sample design (sensu Wynne 2010 and Wynne et
al. 2012). These inventories, conducted during the most produc-
PARK SCIENCE • VOLUME 30 • NUMBER 1 • SUMMER 2013
52
tive times of year (i.e., spring and summer), would likely result in
the detection of additional troglomorphic arthropods.
Because cave-adapted animals are cryptic, they are often diffi cult
to detect and researchers must conduct numerous site visits to
obtain even a baseline knowledge of community composition.
For example, Krejca and Weckerly (2007) reported that 10 to 22
site visits were required to detect three endangered arthropods
known to occur in Texas caves. Although it is not directly ap-
plicable to terrestrial cave-dwelling invertebrates, Culver et al.
(2004) reported that Sket (1981) discovered a new stygobite (an
aquatic cave-adapted animal belonging to a new genus) after
more than 100 collecting trips to a cave in Slovenia. During this
study, I identifi ed two potential troglobites and detected only one
of ve troglobites originally identifi ed by Northup and Welbourn
(1997). This not only underscores the ineffi ciency in our abilities
to eff ectively detect cave-adapted animals but also emphasizes the
need for additional inventory work in deep zone microhabitats.
ELMA-054 is home to a troglomorphic bristletail (order Diplura:
family Campodeidae). It has been detected on the mud fl oors
of a small chamber at the terminus of this cave. This animal may
prove to be a narrow-range endemic (occurring in this cave and
nowhere else on the planet). To best protect this animal and its
habitat, in 2013 monument personnel permanently closed the
deepest section of ELMA-054 to all recreational use.
Another important and highly sensitive microhabitat is moss gar-
dens. These areas have been identifi ed as relict habitats of the last
glacial maximum (approximately 20,000 years ago) and support
species now restricted to this environment at both El Malpais
(Northup and Welbourn 1997) and in Oregon (Benedict 1979).
Species richness for both ELMA-012 and ELMA-008 was driven
by the large number of species detected within moss gardens at
cave entrances and beneath skylights. Roughly 25% of the arthro-
pods detected during the Northup and Welbourn (1997) study
were found within moss gardens.
Because moss gardens are considered relict habitats and have
been shown to support large numbers of species, this micro-
habitat should be aff orded the highest level of protection. In 2013
ELMA-012 was closed to recreational use. Moss gardens within
ELMA-008 have been roped off and signage has been posted in-
dicating the fragility of these habitats. Based on my observations
of both caves since 2005, this approach seems to be deterring foot
traffi c in these areas. However, some of the posts supporting the
rope have fallen. More frequent maintenance of the posts and
ropes, and adding more signage in ELMA-008
, are relatively inex-
pensive measures that may serve to better protect these important
microhabitats. Should ELMA-012 reopen in the future, I recom-
mend using the same management and maintenance approach
described for ELMA-008.
I did not detect any arthropods in ELMA-029 because I did not
have an opportunity to sample the moss gardens in the cave en-
trance (as Northup and Welbourn [1997] did during their work).
I observed no signs of recent human use or visitation when I was
there in 2007. Given its remote location (approximately 1.6 km
[1 mi] from an unmaintained dirt road), this cave and its moss
gardens are likely well protected.
ELMA-029 also contains the most signifi cant ice deposit on the
monument, with a meters-thick ice sheet extending from near
the entrance to the back of the cave. Cave interior and deep zone
temperatures fl uctuated from near to below freezing (mean tem-
perature = 0.141°C [32.25 °F], standard deviation = 1.21°C [2.18°F])
during the arthropod sampling period. Although this cave is not
suitable habitat for most arthropod species, it is possible that ice
crawlers (order Notoptera: family Grylloblattidae) occur there
and in other ice caves on private lands adjacent to the monument.
These animals are known to occur in caves at both Oregon Caves
and Lava Beds National Monuments (Jarvis and Whiting 2006)
and would be a signifi cant discovery at El Malpais. If ice crawlers
exist within this cave, these animals would likely be relict species
of the last glacial maximum. I suggest surveys for ice crawlers be
conducted at ELMA-029, as well as at other ice caves in the El
Malpais region.
This work resulted in the identifi cation of seven new species of cave-dwelling
arthropods (including two potential troglobites), range expansions of two parasitoidal
wasps, and two caves containing signifi cant root curtains hanging from the ceiling.
RESEARCH REPORTS
53
Future directions
The information presented here provides a solid foundation on
which to continue building knowledge of cave natural resources
on the national monument, and has already proven useful in man-
aging these resources. Additional studies targeting the use of lava
tubes by cave-roosting bats, the distributional extent of known
troglomorphic arthropods in caves or groups of caves, additional
sampling for several of the new species discussed here, and fur-
ther study of cave deep zones, root curtains, moss gardens, and
cave ice sheets will be required to obtain the data necessary for
optimal conservation and management of lava tube cave biologi-
cal resources at El Malpais National Monument. My hope is that
some of the protocols presented here and the recommendations
made will be useful in the development and implementation of a
monitoring framework that may be used to gauge the response of
sensitive cave-dwelling animals to recreational use, invasive spe-
cies, global climate change, and white-nose syndrome.
Acknowledgments
Special thanks to Kayci Cook Collins, David Hays, and Dana Sul-
livan for their guidance and support of this research. Ara Kooser,
Jessica Markowski, Peter Polsgrove, and Kyle Voyles provided
assistance in the fi eld. Kyle Voyles codeveloped the arthropod
sampling protocol. I extend much gratitude to Jeff Alford and his
family at Bandera Ice Caves for providing us with a secure camp-
site during both site visits. Frank Howarth, David Hays, Dale Pate,
Je Selleck, and Stefan Sommer provided comments leading to
the improvement of earlier versions of this manuscript. This proj-
ect was funded through a Colorado Plateau–CESU cooperative
agreement between El Malpais National Monument and North-
ern Arizona University.
Literature cited
Allen, H. W. 1971. A monographic study of the genus
Tiphia
of western
North America. Transactions of the American Entomological Society
97:201–359.
Badino, G. 2004. Cave temperatures and global climate change.
International Journal of Speleology 33:103114.
Barr, T. C., Jr. 1968. Cave ecology and evolution of troglobites. Pages 35
102
in
T. Dobzhansky, M. Hecht, and W. Steere, editors. Evolutionary
biology, Volume 2. Appleton-Century-Crofts, New York, New York,
USA.
Benedict, E. M. 1979. A new species of
Apochthonius
Chamberlin from
Oregon (Pseudoscorpionida, Chthoniidae). Journal of Arachnology
7:7983.
Bogan, M. A., K. Geluso, S. Haymond, and E. W. Valdez. 2007. Mammal
inventories for eight national parks in the Southern Colorado Plateau
Network. Natural Resource Technical Report NPS/SCPN/NRTR-
2007/054. National Park Service, Fort Collins, Colorado, USA.
Brown, P., R. Berry, and C. Brown. 1993. Bats and mines: Finding solutions.
Bats 11:12–13.
Cascadden, T. E., A. M. Kudo, and J. W. Geisman. 1997. Discovering the
relationships in a family of volcanoes: Cerro Candelaria, Twin Craters,
Lost Woman Crater and Lava Crater. New Mexico Bureau of Mines and
Mineral Resources 156:41–52.
Chevaldonné, P., and C. Lejeune. 2003. Regional warming-induced species
shift in northwest Mediterranean marine caves. Ecology Letters 6:371–
379.
Christman, M. C., D. C. Culver, M. K. Madden, and D. White. 2005. Patterns
of endemism of the eastern North American cave fauna. Journal of
Biogeography 32:1442–1452.
Culver, D. C. 1986. Cave faunas. Pages 427–443
in
M. Soulé, editor.
Conservation biology. Sinauer Associates, Sunderland, Massachusetts,
USA.
Culver, D. C., M. C. Christman, B. Sket, and P. Trontelj. 2004. Sampling
adequacy in an extreme environment: Species richness patterns in
Slovenian caves. Biodiversity and Conservation 13:12091229.
Culver, D. C., L. L. Master, M. C. Christman, and H. H. Hobbs III. 2000.
Obligate cave fauna of the 48 contiguous United States. Conservation
Biology 14:386401.
Dobkin, D. S., R. D. Gettinger, and M. G. Gerdes. 1995. Springtime
movements, roost use and foraging activity of Townsends big-eared
bat (
Plecotus townsendii
) in central Oregon. Great Basin Naturalist
55:315321.
Elliott, W. R. 1992. Fire ants invade Texas caves. American Caves Winter:13.
. 2000. Conservation of the North American cave and karst biota.
Pages 665669
in
H. Wilkens et al., editors. Subterranean ecosystems,
ecosystems of the world, Volume 30. Elsevier, Amsterdam, the
Netherlands.
Hamilton-Smith, E., and S. Eberhard. 2000. Conservation of cave
communities in Australia. Pages 647–664
in
H. Wilkens et al., editors.
Subterranean ecosystems, ecosystems of the world, 30th edition.
Elsevier, Amsterdam, the Netherlands.
Howarth, F. G. 1980. The zoogeography of specialized cave animals: A
bioclimatic model. Evolution 34:394406.
. 1982. Bioclimatic and geological factors governing the evolution
and distribution of Hawaiian cave insects. Entomologia Generalis
8:17–26.
PARK SCIENCE • VOLUME 30 • NUMBER 1 • SUMMER 2013
54
. 1983. Ecology of cave arthropods. Annual Review of Entomology
28:365–389.
Howarth, F. G., S. A. James, W. McDowell, D. J. Preston, and C. T. Imada.
2007. Identifi cation of roots in lava tube caves using molecular
techniques: Implications for conservation of cave arthropod faunas.
Journal of Insect Conservation 11:251–261.
Howarth, F. G., and F. D. Stone. 1993. Conservation of Hawaii’s
speleological resources. Pages 124126
in
W. R. Halliday,
editor. Proceedings of the Third International Symposium on
Volcanospeleology, Bend, Oregon, 1982. International Speleological
Foundation, Seattle, Washington, USA.
Humphrey, S. R., and T. H. Kunz. 1976. Ecology of a Pleistocene relict, the
western big-eared bat (
Plecotus townsendii
), in the southern Great
Plains. Journal of Mammalogy 57:470494.
Jarvis, K. J., and M. F. Whiting. 2006. Phylogeny and biogeography of
ice crawlers (Insecta: Grylloblattodea) based on six molecular loci:
Designating conservation status for Grylloblattodea species. Molecular
Phylogenetics and Evolution 41:222–237.
Krejca, J. K., and B. Weckerly. 2007. Detection probabilities of karst
invertebrates. Pages 283–289
in
W. R. Elliott, editor. Proceedings
of 18th National Cave and Karst Management Symposium (St.
Louis, Missouri, USA, 812 October 2007). Texas Parks and Wildlife
Department, Austin, Texas, USA.
McCracken, G. 1989. Cave conservation: Special problems of bats. National
Speleological Society Bulletin 51:49–51.
Mitchell, R. W. 1970. Total number and density estimates of some species
of cavernicoles inhabiting Fern Cave, Texas. Annales de Spéléologie
25:7390.
Northup, D. E., and W. C. Welbourn. 1997. Life in the twilight zoneLava
tube ecology, natural history of El Malpais National Monument. New
Mexico Bureau of Mines and Mineral Resources, Bulletin 156:69–82.
Peck, S. B. 1976. The effect of cave entrances on the distribution of cave-
inhabiting terrestrial arthropods. International Journal of Speleology
8:309–321.
Pierson, E. D., M. C. Wackenhut, J. S. Altenbach, P. Bradley, P. Call, D. L.
Genter, C. E. Harris, B. L. Keller, B. Lengus, L. Lewis, B. Luce, K. W.
Navo, J. M. Perkins, S. Smith, and L. Welch. 1999. Species conservation
assessment and strategy for Townsend’s big-eared bat (
Corynorhinus
townsendii townsendii
and
Corynorhinus townsendii pallescens
). Idaho
Conservation Effort, Idaho Department of Fish and Game, Boise, Idaho,
USA.
Pulido-Bosch, A., W. Martín-Rosales, M. López-Chicano, C. M. Rodríguez-
Navarro, and A. Vallejos. 1997. Human impact in a tourist karstic cave
(Aracena, Spain). Environmental Geology 31:142–149.
Reddell, J. R. 1994. The cave fauna of Texas with special reference to the
western Edwards Plateau. Pages 31–50
in
W. R. Elliott and G. Veni,
editors. The caves and karst of Texas. National Speleological Society,
Huntsville, Alabama, USA.
Reeves, W. K. 1999. Exotic species of North American caves. Pages 164
166
in
K. Henderson, editor. Proceedings of the 1999 National Cave and
Karst Management Symposium. Chattanooga, Tennessee, USA.
Sket, B. 1981.
Niphargobates orophobata
n.g., n.sp. (Amphipoda,
Gammaridae s.l.) from cave waters in Slovenia (NW Yugoslavia). Bioloski
Vestnik 29:105–118.
Taylor, S. J., J. Krejca, J. E. Smith, V. R. Block, and F. Hutto. 2003.
Investigation of the potential for red imported fi re ant (
Solenopsis
invicta
) impacts on rare karst invertebrates at Fort Hood, Texas: A
eld study. Illinois Bexar County Karst Invertebrates Draft Recovery
Plan Natural History Survey, Center for Biodiversity Technical Report
2003(28):1–153.
U.S. Fish and Wildlife Service (USFWS). 2012. North American bat death
toll exceeds 5.5 million from white-nose syndrome. News release, 17
January 2012. Accessed 1 February 2012 from http://www.fws.gov
/whitenosesyndrome/pdf/WNS_Mortality_2012_NR_FINAL.pdf.
Wilkins, K. T. 1989.
Tadarida brasiliensis
. Mammalian Species 331:1–10.
Winkler, W. G., and D. B. Adams. 1972. Utilization of southwestern bat caves
by terrestrial carnivores. American Midland Naturalist 87:191–200.
Wynne, J. J. 2006. Cave trip report and Junction Cave bat hibernacula, 4
February 2006. Unpublished report submitted 15 February 2006 to
El Malpais National Monument, National Park Service, Grants, New
Mexico. 1 page.
. 2010. Preliminary results of arthropod baiting and surface
sampling at Grand Wash Cave, Grand CanyonParashant National
Monument, Arizona. On fi le with National Park Service, Grand Canyon
Parashant National Monument, Saint George, Utah. 11 pages.
Wynne, J. J., L. Pakarati, and C. Tambley. 2012. Artrópodos cavercolas
en zonas profundas en cavernas en Rapa Nui. On fi le with Corporacíon
Nacional Forestal (CONAF), Parque Nacional Rapa Nui, Easter Island,
Chile. 11 pages.
About the author
J. Judson “Jut” Wynne is a research ecologist with the Colorado
Plateau Biodiversity Center and Colorado Plateau Research Station
and a PhD candidate in biological sciences at Northern Arizona
University, Flagstaff. For more than 10 years, he has studied and
published on cave ecosystems and microclimates of Belize, Chile,
Easter Island, Hawaii, and throughout the western United States.
He can be reached via http://www.jutwynne.com.
APPENDIX A IS AVAILABLE ONLINE AT
HTTP://WWW.NATURE.NPS.GOV/PARKSCIENCE
/ARCHIVE/PDF/ARTICLEPDFS/PARKSCIENCE30(1)
SUMMER2013_A1–A12_WYNNE_3653.PDF
RESEARCH REPORTS
55
Editor’s note: The following is an online-only supplement to the research report “Inventory, conservation, and
management of lava tube caves at El Malpais National Monument, New Mexico, by J. Judson Wynne. It can be
cited as Wynne, J. J. 2013. Appendix A: Annotated list of cave-dwelling taxa. [Online supplement.] Park Science
30(1)Appendix A:1–12. Available online at http://www.nature.nps.gov/ParkScience/archive/PDF/Article_PDFs
/ParkScience30(1)Summer2013_A1-A12_Wynne_3653.pdf.
Author’s notes: In cases where members of a given morphospecies were detected only in entrances and twi-
light zones, I erred cautiously and referred to them as “eisodophiles.” In cases where both the location of the
detection and known information concerning the morphospecies supported the likelihood of an animal being
“troglophillic,” but I was still uncertain, I categorized the animal as a “questionable troglophile.” Additionally,
when a morphospecies was found only in the deep zone of a cave (or several individuals of a morphospecies
occurred only within the deep zone) but troglomorphic characters were lacking, I also referred to it as “question-
able troglophile.
T
HERE WERE SEVERAL CASES WHERE INDIVIDUALS EVADED CAPTURE BUT WERE BELIEVED TO
represent a distinct arthropod morphospecies for a given cave. Because this information is of limited value
in this article, arthropod morphospecies groups for which specimens are lacking were not included.
However, this information has been integrated into a larger El Malpais morphospecies database and will be ana-
lyzed and the results reported in additional publications.
For arthropod groups actively being studied, I either sent specimens or high-resolution images of specimens to
taxonomic specialists for identification or verification of my identifications. These experts include Rolf Aalbu,
Department of Entomology, California Academy of Sciences, San Francisco, California (Coleoptera:
Tenebrionidae); R. Thomas Allen, The Academy of Natural Sciences of Drexel University, Philadelphia,
Pennsylvania (Diplura); Max Barclay, Natural History Museum, London (Coleoptera), and Thomas Barr (deceased),
formerly with Department of Biology, University of Kentucky, Lexington, Kentucky (Coleoptera: Carabidae); Ernest
Bernard, Department of Entomology, The University of Tennessee, Knoxville (Collembola); Jostein Kjaerandsen,
Museum of Zoology, Lund University, Lund, Sweden (Diptera: Mycetophilidae); Sarah Oliveira, Department of
Biology, University of São Paulo, Brazil (Diptera: Mycetophilidae); Theodore Cohn (deceased), formerly with
Department of Zoology, San Diego State University, California (Orthoptera: Rhamphidophoridae); Lynn Kimsey,
Department of Entomology, University of California, Davis (Hymenoptera: Tiphiinae); Robert Johnson, School of
Life Sciences, Arizona State University, Tempe (Formicidae); Edward Mockford, Department of Biology, University
of Illinois, Normal (Psocoptera); Glené Mynhardt, Department of Evolution, Ecology, and Organismal Biology, The
Ohio State University, Columbus (Coleoptera: Ptinidae); Barry O’Connor, Department of Ecology and Evolutionary
Biology, University of Michigan, Ann Arbor (Acari); Stewart Peck, Department of Biology, Carleton University,
Ottawa, Ontario, Canada (Coleoptera: Leiodidae); Pierre Paquin, Cave and Endangered Invertebrate Research,
SWCA Environmental Consultants, Austin, Texas (Araneae); William Shear, Department of Biology, Hampden-
Sydney College, Hampden Sydney, Virginia (Myriapods and Opiliones); and Harald Schillhammer, Department of
Entomology, Naturhistorische Museum, Vienna, Austria (Coleoptera: Staphylinidae). For all other specimens,
Colorado Plateau Museum of Arthropod Biodiversity staff and I identified the specimens to the lowest taxonomic
level possible using available taxonomic keys.
“Det.” following each species or morphospecies designation is the abbreviation for the Latin dēterminēvit or
“determined by.”
APPENDIX A
Annotated list of cave-dwelling taxa
By J. Judson Wynne
PARK SCIENCE • SUMMER 2013 • VOLUME 30 • NUMBER 1 Appendix A-1
Research Report
PARK Science
ISSN 1090-9966 (online)
Published by
U.S. Department of the Interior
National Park Service
Natural Resource Stewardship & Science
Of ce of Education and Outreach
Lakewood, Colorado
Phylum Arthropoda
Class Arachnida
Order Araneae
Family Araneidae
Metellina mimetoides
Chamberlin & Ivie, 1941. Det. P. Paquin. Eisodophile.
One adult female was collected via timed search in the twilight zone of ELMA-262. Additionally,
one juvenile specimen that may represent this species was collected via timed search at the
entrance of ELMA-262.
Family Linyphiidae
Note:
Numerous troglobitic and troglophillic forms of this family are known globally (e.g., Ruzicka
1998; Deltshev and Curcic 2002; Miller 2005).
Linyphiidae sp. Det. P. Paquin. Eisodophile.
One juvenile specimen was collected opportunistically near the entrance of ELMA-262. Another
juvenile was collected via timed search in the twilight zone of ELMA-303.
Lepthyphantes
sp. Det. P. Paquin. Eisodophile.
Two female specimens were collected by timed searches at the entrance of ELMA-012; one female
specimen was collected using direct intuitive searches in the moss gardens of ELMA-008.
Porrhomma
sp. 1. Det. P. Paquin. Troglophile?
One female specimen was collected using direct intuitive searches within root curtains in the
deep zone of ELMA-315.
Porrhomma
sp. 2. Det. P. Paquin. Troglophile?
One female specimen was collected using direct intuitive searches within root curtains in the
deep zone of ELMA-303. P. Paquin (personal communication, e-mail, 23 March 2007) suggests it
differs from
Porrhomma
sp. 1.
Family Liocranidae
Liocranidae sp. Det. P. Paquin. Troglophile?
One juvenile specimen was collected via timed search in the deep zone of ELMA-012.
Family Nesticidae
Note:
Nesticidae has an impressive cave fauna globally (Hedin 1997; Cokendolpher and Reddell
2001; Snowman et al. 2010).
Nesticidae sp. Det. P. Paquin. Eisodophile.
One juvenile specimen was collected opportunistically from the twilight zone of ELMA-012.
Eidmanella pallida
(Emerton, 1875). Det. P. Paquin. Troglophile.
Three females were collected using direct intuitive searches within root curtains in the deep zone
of ELMA-315. Two juvenile specimens (identified as Nesticidae sp.) were collected using direct
intuitive searches within root curtains in the deep zone of this cave. While unconfirmed, these
juveniles may also be
Eidmanella pallida
.
Note:
Reddell and Cokendolpher (2004) consider this species a troglophile in Texas caves.
Appendix A-2
PARK SCIENCE • VOLUME 30 • NUMBER 1 • SUMMER 2013
Family Pholcidae
Note:
Both troglophillic and troglobitic forms of this family are known globally (e.g., Gertsch and
Peck 1992; Deeleman-Reinhold 1993; Chen et al. 2011; Ferreira et al. 2011).
Psilochorus
sp. 1. Det. P. Paquin. Troglophile?
One male and two females were collected using direct intuitive searches within root curtains in
the deep zone of ELMA-303. Three males were collected via timed searches (n = 2) and pitfall
trapping (n = 1) at the entrance and beneath the skylights of ELMA-008. One female specimen
was identified via timed search in the twilight zone, directly below the entrance of ELMA-315.
Two males were collected by timed searches in both the entrance and deep zone of ELMA-012,
and two individuals (1 male, 1 female) were collected via timed search from the entrance in
Hummingbird Cave.
Psilochorus
sp. 2. Det. P. Paquin. Troglophile?
Two individuals were collected opportunistically in the deep zone near the bat maternity roost in
ELMA-062. One adult female specimen was designated as a different species from
Psilochorus
sp.
1 (P. Paquin, personal communication, e-mail, 4 December 2009). Additionally, one juvenile speci-
men identified as
Psilochorus
was collected from the same cave. I suggest it is probably the same
morphospecies because an adult female was identified in the same area as the juvenile specimen.
Family Theridiidae
Achaearanea porteri
(Banks, 1896). Det. P. Paquin. Troglophile.
Two females were collected using timed searches, one near the entrance and the other in the twi-
light zone of ELMA-303. Three females were collected via direct intuitive searching (n = 2) in root
curtains and with timed searches (n = 1) in the deep zone of ELMA-315.
Note:
Cokendolpher and Reddell (2001) consider this species a troglophile in Texas caves.
Nesticodes rufipes
(Lucas, 1846). Det. P. Paquin. Troglophile.
Three adult females were collected using direct intuitive searches in root curtains from the deep
zone of ELMA-315.
Note:
Because all were located within the same location and none had characters suggestive of
troglomorphism, I consider this spider a troglophile. Additionally, theridiid spiders have been
widely documented globally as being both troglophiles and troglobites (e.g., Ferreira and Martins
1998; Ruzicka 1998; Dippenaar-Schoeman and Myburgh 2009).
Steatoda
sp. Det. P. Paquin. Unknown.
One juvenile specimen was collected from a pitfall trap within the twilight zone of ELMA-062.
Theridion
n.sp.? Det. P. Paquin. Troglobite?
One adult female was collected via timed search in the twilight zone of ELMA-262. P. Paquin
(personal communication, e-mail, 23 March 2007) suggests this may be a new species, and
potentially has cave-adapted characteristics.
Appendix A-3
RESEARCH REPORTS
Order Opiliones
Family Sclerosomatidae
Leiobunum townsendii
Weed, 1893. Det. W. Shear. Trogloxene.
This harvestman (n = 13) was identified from ELMA-012, ELMA-062, ELMA-008, ELMA-262,
ELMA-303, ELMA-315, and Hummingbird Cave. It was collected via direct intuitive search in the
moss gardens of ELMA-008 and ELMA-012 and in root curtains in the deep zone of both ELMA-
303 and ELMA-315. It was collected both opportunistically and via timed search in the entrances
and twilight zones of ELMA-062 and Hummingbird Cave. W. Shear (personal communication,
e-mail, 12 April 2009) suggests this group in western North America requires major revision. It is
possible multiple species exist across the southwestern United States, or greater North America.
However, until it is revised, the accepted name provided here will be used.
Subclass Acari
Order Sarcoptiformes
Family Histiostomatidae
Histiostoma
n.sp. Det. B. O’Connor. Troglophile?
Two deutonymphs were collected during timed searches in the deep zone of ELMA-315. B.
O’Connor indicates this is an undescribed species. This animal is similar to H. pierrestrinati
described from Carlsbad Cavern (B. O’Connor, personal communication, e-mail, 3 August 2012).
Order Trombidiformes
Family Bdellidae
Bdellidae sp. Det. B. O’Connor. Troglophile?
One specimen was collected by direct intuitive searches in the deep zone of ELMA-303. The
palpi were damaged during collection, so lower-level taxonomic identification was not possible.
B. O’Connor (personal communication, e-mail, 3 August 2012) indicates this family contains
predators of soil, leaf litter, and littoral zones.
Family Erythraeidae
Erythraeus
sp.? Det. B. O’Connor. Eisodophile.
Five specimens were captured via pitfall trapping from the twilight zone of ELMA-008.
B. O’Connor (personal communication, e-mail, 3 August 2012) indicates this genus is known from
the Southwest, but no species are described. Additional analysis will be required to identify these
specimens to a lower taxonomic level.
Family Rhagiididae
Rhagiididae sp. Det. B. O’Connor. Troglophile?
One specimen was collected by direct intuitive searches in the dark zone of ELMA-012. The speci-
men was damaged and could not be identified beyond family level.
Family Smarididae
Phanolophus
sp. Det. B. O’Connor. Unknown.
One specimen was collected via pitfall trapping at the entrance of ELMA-012. This family of pred-
atory mites has not been studied in North America (B. O’Connor, personal communication, e-mail,
3 August 2012).
Appendix A-4
PARK SCIENCE • VOLUME 30 • NUMBER 1 • SUMMER 2013
Subphylum Myriapoda
Class Chilopoda
Order Lithobiomorpha
Family Gosibiidae
Gosibiidae sp. Det. B. Shear. Troglophile?
One specimen was collected using direct intuitive searches from root curtains in the deep zone of
ELMA-303. Additional specimens will be required to identify this centipede beyond the family
level (W. Shear, personal communication, e-mail, 9 October 2009).
Class Diplopoda
Order Chordeumatida
Family Contylidae
Austrotyla
sp.? Det. W. Shear. Eisodophile.
This specimen (n = 1), identified to genus level by W. Shear, was collected via direct intuitive
search from the moss gardens of ELMA-008. Additional specimens will be required to identify this
animal to a lower taxonomic level.
Austrotyla
cf
coloradensis
(Chamberlin, 1910). Det. W. Shear. Troglophile?
One specimen was collected using direct intuitive searches from root curtains in the deep zone of
ELMA-315. This is a tentative species designation because of a lack of material. Additional speci-
mens will be required to confirm this species designation.
Class Entognatha
Order Collembola
Note:
The two new collembolan species will be included in a paper describing several new cave-
dwelling Collembola species from the southwestern United States.
Family Entomobryidae
Drepanura
n.sp. Det. E. Bernard. Troglophile?
One specimen was collecting using pitfall trapping near the entrance of ELMA-008. E. Bernard
(personal communication, e-mail, 15 July 2010) indicates this specimen represents a new species.
Entomobrya guthriei
Mills, 1931. Det. E. Bernard. Troglophile?
Five specimens were collected via pitfall trapping from the twilight zone to the deep cave zone of
ELMA-110.
Entomobrya zona
? Christiansen & Bellinger, 1980. Det. E. Bernard. Troglophile?
All specimens were collected in the entrances and twilight zones of ELMA-012 (n = 28) and
ELMA-008 (n = 4). Seven specimens were collected using direct intuitive searches from moss gar-
dens beneath the skylights of ELMA-012. All of the remaining specimens were captured using pit-
fall trapping. They likely represent
E. zona
. E. Bernard (personal communication, e-mail, 15 July
2010) made this tentative species designation, but indicated the specimens are not a “sure fit”
for this species.
Family Tomoceridae
Pogonognathellus
n.sp. Det. E. Bernard. Eisodophile.
All specimens were collected via direct intuitive searches from the moss gardens of ELMA-008
(n = 10) and opportunistic collecting of ELMA-012 (n = 2). E. Bernard (personal communication,
e-mail, 15 July 2010) suggests these specimens represent a new species.
Appendix A-5
RESEARCH REPORTS
Order Diplura
Family Campodeidae
Campodeidae n.sp. Det. J. Wynne and T. Allen. Troglobite.
This animal was first reported by Northup and Welbourn (1997). Five specimens were collected
using direct intuitive searches from the “mud room” at the terminus of ELMA-054. Dipluran tax-
onomist Dr. Thomas Allen has these specimens and has confirmed this as a new species (personal
communication, e-mail, 5 May 2013). I will be working with him to describe this new species.
Class Insecta
Order Coleoptera
Family Carabidae
Rhadine
n.sp.
perlevis
species-group. Det. T. Barr. Trogloxene.
These carabid beetles (n = 25) were identified primarily by pitfall trapping (but also with opportu-
nistic collecting and timed searches) from ELMA-062, ELMA-110, ELMA-262, ELMA-303, and
ELMA-315. This animal was observed from the twilight to deep zones of most caves. These speci-
mens were initially sent to Dr. Thomas Barr for identification. T. Barr (personal communication,
e-mail, 12 June 2009) suggested the specimens represent a new species and they belong to the
perlevis
species-group of
Rhadine
. Dr. Barr passed away in April 2011. The specimens are now at
the Carnegie Museum of Natural History in Pittsburgh, Pennsylvania, and are awaiting formal
description. Dr. Kipling Will, Essig Museum of Entomology, University of California, Berkeley, is
coordinating this effort.
Family Cryptophagidae
Cryptophagidae sp. Det. M. Barclay. Eisodophile.
One specimen was collected via pitfall trapping from the entrance of ELMA-062. Additional work
will be required to identify this specimen to a lower taxonomic level.
Family Leiodidae
Dissochaetus arizonensis
Hatch, 1933. Det. S. Peck. Accidental.
This leiodid beetle was collected from cave entrances of ELMA-012 (n = 1) and ELMA-062 (n = 1),
while specimens from ELMA-315 (n = 2) were detected in the cave deep zone; all were captured
using baited pitfall traps.
Note:
S. Peck (personal communication, e-mail, 28 February 2013) suggests this species is an
accidental because there are no data to suggest it is a regular cave dweller or that it reproduces
in caves.
Family Melyridae
Listrus
sp. Det. M. Barclay. Eisodophile.
This coleopteran was captured via pitfall trapping (n = 1) in the twilight zone of ELMA-262. This
specimen will require further study.
Family Ptinidae
Niptus ventriculus
LeConte, 1859. Det. G. Mynhardt. Troglophile.
Five spider beetle specimens were collected via pitfall trapping in ELMA-008 (n = 1), ELMA-012
(n = 2), and ELMA-262 (n = 1) and by opportunistic collecting in ELMA-062 (n = 1). Four speci-
mens were collected in the cave entrances, while one specimen was collected in the twilight
zone.
Note:
Spilman (1968) documented this species in packrat middens, while Aalbu (2005) indicated
Appendix A-6
PARK SCIENCE • VOLUME 30 • NUMBER 1 • SUMMER 2013
larvae and potentially adults feed on the scat of packrats. Given that habitat exists for these spi-
der beetles and that they complete a portion of their life cycle underground, I consider this animal
a troglophile.
Family Staphylinidae
Staphylinidae sp. Det. J. Wynne. Eisodophile.
One individual was collected from the entrance of ELMA-012 during time searches. Additional
work will be required to identify this specimen to a lower taxonomic level.
Subfamily Tachyporinae
Sepedophilus
sp. Det. H. Schillhammer. Eisodophile.
Three individuals were collected from the twilight zone of ELMA-062 (n = 2) and entrance of
ELMA-315 (n = 1). Each specimen was detected using a different technique from the others:
opportunistic collecting, timed searches, and pitfall traps. H. Schillhammer (personal communica-
tion, e-mail, 19 April 2013) suggests this genus is generally not associated with caves.
Family Tenebrionidae
Neobaphion planipennis
(LeConte, 1866). Det. R. Aalbu. Trogloxene.
Four individuals were collected opportunistically and via timed search from ELMA-062 (n = 3)
and using direct intuitive searches in ELMA-303 (n = 1). In ELMA-062 this species was observed
in the dark zone and beneath a skylight entrance; the individual in ELMA-303 was collected from
the deep zone.
Note:
Aalbu et al. (2012) consider this species an occasional trogloxene in ELMA-062.
Order Diptera
Family Culicidae
Culicidae sp. Det. J. Wynne. Trogloxene.
One culicid fly was collected opportunistically from the entrance of ELMA-012 and one via timed
search in the deep zone of ELMA-315. Additional work will be required to identify this specimen
to a lower taxonomic level.
Note:
Reeves et al. (2000) and Makiya and Taguchi (1982) identified mosquitoes as trogloxenes.
Family Mycetophilidae
Mycetophila
sp. Det. J. Kjaerandsen and S. Oliveira. Trogloxene?
One specimen was collected using direct intuitive searches from the root curtains in the deep
zone of ELMA-303. Additional work will be required to identify this specimen to a lower taxo-
nomic level.
Note:
Peck (1981) considered a morphospecies of this genus and five morphospecies of this family
to be trogloxenes from two caves (>2,134 m [7,000 ft] elevation) in the Uinta Mountains, Utah.
Additionally, from caves in Grand Canyon National Park, Peck (1980) considered a morphospecies
of this genus to be a trogloxene.
Family Phoridae
Phoridae sp. Det. J. Wynne. Eisodophile.
Eight specimens were collected from pitfall traps at the entrance of ELMA-062 (n = 7) and in the
twilight zone of ELMA-008 (n = 1). One individual was collected using direct intuitive searches in
the moss gardens beneath skylights of ELMA-012. Additional work will be required to identify
Appendix A-7
RESEARCH REPORTS
these specimens to a lower taxonomic level.
Family Sciaridae
Sciaridae sp. Det. J. Wynne. Eisodophile.
Twenty-one specimens were collected via opportunistic collecting, pitfall trapping, and timed
searches from the entrance to the middle of ELMA-062; one specimen was collected using direct
intuitive searches from the moss gardens beneath a skylight of ELMA-008; and three specimens
were collected opportunistically from the entrance of ELMA-061. Additional work will be required
to identify these specimens to a lower taxonomic level.
Order Hemiptera
Infraorder Fulgoromorpha
Superfamily Fulgoroidea
Fulgoroidea n.sp.? Det. J. Wynne. Troglobite?
Nymphal-stage planthoppers were collected using direct intuitive searches in root curtains from
the deep zones of ELMA-303 and ELMA-315. Adults will be required to confirm troglomorphism,
identify to a lower taxonomic level, and determine new species status.
Order Hymenoptera
Family Formicidae
Liometopum
sp. Det. R. Johnson. Eisodophile.
One undetermined
Liometopum
specimen was collected using direct intuitive searches in the
moss gardens of ELMA-008.
Pheidole
sp. Det. R. Johnson. Eisodophile.
Two minor workers (R. Johnson, personal communication, e-mail, 10 December 2010) were col-
lected via pitfall trapping near the entrance and at close proximity to the moss gardens of ELMA-
008.
Family Tiphiidae
Note:
All specimens of both tiphiid wasp species were found in a torpor beneath rocks; given the
time of season, I suggest these individuals were in the early stages of hibernation and were likely
using moss gardens as winter habitat.
Tiphia andersoni
Allen, 1971. Det. L. Kimsey. Eisodophile.
One female specimen was collected using direct intuitive searches in moss gardens (beneath
large skylights) of both ELMA-012 and ELMA-008. Historically, this wasp is known to occur in
central Mexico as well as southeastern and north-central Arizona (Allen 1971). This animal was
not known to occur in New Mexico and thus represents a range expansion.
Tiphia nona
Allen, 1965. Det. L. Kimsey. Eisodophile.
One female specimen was collected using direct intuitive searches in the moss gardens of ELMA-
008. Previously it was known from central Mexico, southeastern Arizona to the southern extent
of the Mogollon Rim, and one locality in southwestern Kansas (Allen 1971). This animal was not
known to occur in New Mexico and thus represents a range expansion.
Appendix A-8
PARK SCIENCE • VOLUME 30 • NUMBER 1 • SUMMER 2013
Order Lepidoptera
Note:
None of the larval specimens were reared in the lab and I was unable to locate a key for
Lepidoptera larvae. Thus, all lepidopteran specimens have been sorted into operational taxonomic
units, and further identifications were not possible before this article was published. This level of
identification is acceptable for community-level as well as other analyses, which will be the sub-
ject of additional scientific publications.
Lepidoptera sp. 1. Det. J. Wynne. Troglophile?
Three larval specimens were collected with pitfall traps (n = 2) and via direct intuitive searches
(n = 1) from the root curtains within the deep zone of ELMA-315.
Lepidoptera sp. 2. Det. J. Wynne. Troglophile?
Four larval specimens were collected using direct intuitive searches of the root curtains within the
deep zone of ELMA-315 (n = 3) and ELMA-303 (n = 1).
Lepidoptera sp. 3. Det. J. Wynne. Troglophile?
One larval specimen was collected using direct intuitive searches of the root curtains within the
deep zone of ELMA-315.
Lepidoptera sp. 4. Det. J. Wynne. Troglophile?
One larval specimen was collected using direct intuitive searches of the root curtains within the
deep zone of ELMA-315.
Lepidoptera sp. 5. Det. J. Wynne. Troglophile?
One larval specimen was collected using direct intuitive searches of the root curtains within the
deep zone of ELMA-303.
Lepidoptera sp. 6. Det. J. Wynne. Eisodophile.
One adult moth was collected during a timed search in the entrance of ELMA-262.
Lepidoptera sp. 7. Det. J. Wynne. Eisodophile.
One adult moth (different from Lepidoptera sp. 6) was collected during a timed search in the
entrance of ELMA-012.
Family Tenididae
Tenididae sp. 1. Det. J. Wynne. Eisodophile.
One micro-lepidopteran was collected opportunistically in ELMA-262.
Tenididae sp. 2. Det. J. Wynne. Eisodophile.
One micro-lepidopteran (different from
Tenididae
sp. 1) was found in a pitfall trap in the twilight
zone of ELMA-008.
Appendix A-9
RESEARCH REPORTS
Order Orthoptera
Family Rhaphidophoridae
Ceuthophilus
sp. Det. T. Cohn. Trogloxene.
One juvenile male was captured via pitfall trapping from the entrance of ELMA-010. Given this
animal’s immature state, it was not possible to identify it to a lower taxonomic level.
Ceuthophilus
cf
apache
n.sp. Det. T. Cohn. Trogloxene.
T. Cohn (personal communication, e-mail, 21 March 2011) indicated this was a new
Ceuthophilus
species, which is similar to
Ceuthophilus
cf
apache
. We collected one adult male and one adult
female from ELMA-062, two adult males from ELMA-303, and one adult male from ELMA-315.
This morphospecies was detected using opportunistic collecting, pitfall trapping and timed
searches, and occurred from the entrances to each cave’s dark/deep zone.
Ceuthophilus
(
Geotettix
)
polingi
Hubbell, 1936. Det. T. Cohn. Trogloxene.
T. Cohn and A. Swanson identified all specimens in this group. We collected two adult females
and four adult males from ELMA-262, one adult male from Hummingbird Cave, one adult male
from ELMA-012, one adult male from ELMA-054, one adult female and two adult males from
ELMA-303, and two adult females from ELMA-315. This species was detected using opportunistic
collecting, pitfall trapping, and timed searching, and occurred from the entrances to each cave’s
dark/deep zone. T. Cohn (personal communication, e-mail, 21 March 2011) suggested this animal
was considered rare until recently; we now know it is widespread in its range, but probably
restricted to caves and animal burrows.
Order Psocoptera
Family Psyllipsocidae
Psyllipsocus ramburii
Selys Longchamps, 1872. Det. E. Mockford. Troglophile.
This species was identified from ELMA-062 (n = 2), ELMA-262 (n = 1), and ELMA-315 (n = 6).
With the exception of one individual collected opportunistically, all were detected in pitfall traps
and from cave entrances to the dark/deep zones.
Note:
This species is known to occur in caves globally (E. Mockford, e-mail, 1 February 2013). E.
Mockford and I (unpublished data) recently confirmed this species on Easter Island, South Pacific
Ocean, as well as from a cave on Grand Canyon–Parashant National Monument, Arizona.
Order Siphonaptera
Family Pulicidae
Pulicidae sp. Det. J. Wynne. Parasite.
Nine specimens were collected from ELMA-315. I found no evidence of recent rodent activity
within either cave. However, the presence of fleas suggests recent vertebrate use. Additional
work will be required to identify these specimens to a lower taxonomic level.
Phylum Chordata
Subphylum Vertebrata
Class Reptilia
Family Colubridae
Pituophis catenifer
(Blainville, 1835). Det. J. Wynne. Unknown.
A gopher snake carcass was found in the twilight zone of ELMA-061. This individual had numer-
ous lacerations along the length of its body. A park visitor probably killed the snake. Because I
am uncertain whether the snake was killed in the cave or brought into the cave postmortem, its
functional group status is “unknown.
Appendix A-10
PARK SCIENCE • VOLUME 30 • NUMBER 1 • SUMMER 2013
Class Mammalia
Order Chiroptera
Family Vespertilionidae
Corynorhinus townsendii
Cooper, 1837. Det. J. Wynne. Trogloxene.
This bat has been documented hibernating in ELMA-054 since 2005 (Wynne 2006). A maternity
roost exists at ELMA-110. This maternity roost has been documented both in the tunnel section
prior to the main section of the cave and in the twilight zone of the cave’s main section.
Eptesicus fuscus
(Palisot de Beauvois, 1796). Det. J. Wynne. Trogloxene.
One torpid big brown bat was observed near the entrance of ELMA-054.
Family Molossidae
Tadarida brasiliensis
(I. Geoffroy, 1824). Det. J. Wynne. Trogloxene.
A long-established maternity roost of Mexican free-tailed bats exists in ELMA-062. We observed
bats in residence during the October 2007 work.
Order Rodentia
Family Muridae
Neotoma
sp. Det. J. Wynne. Trogloxene.
Evidence of
Neotoma
sp. was documented at both ELMA-062 and ELMA-061. Both
N.mexicana
and
N.albigula
have been confirmed on the monument (Bogan et al. 2007). Either or both of
these species likely use these caves.
Order Carnivora
Unknown family, genus, and species. Xenosylle?
Small carnivore scat was observed at the entrance of ELMA-054 and in the twilight zone of
ELMA-110. Because we neither observed small carnivores nor saw them hunting bats within
either cave,questionable xenosylle” is most appropriate.
Class Aves
Order Tytonidae
Tyto alba
(Scopoli, 1769). Det. J. Wynne. Eisodophile.
A barn owl was spooked as the team entered ELMA-262. The animal was observed within the
main entrance and flew deeper into the cave toward the next collapse pit entrance, where it exit-
ed the cave.
Literature cited
Aalbu, R. L. 2005. Holey dung: Can you fi nd
Niptus
? Western Cave Conservancy Newsletter 2:1–2.
Aalbu, R. L., A. D. Smith, and C. A. Triplehorn. 2012. A revision of the
Eleodes
(subgenus
Caverneleodes
) with new
species and notes on cave breeding
Eleodes
(Tenebrionidae: Amphidorini). Annales Zoologici 62:199–216.
Allen, H. W. 1971. A monographic study of the genus
Tiphia
of western North America. Transactions of the
American Entomological Society 97:201–359.
Bogan, M. A., K. Geluso, S. Haymond, and E. W. Valdez. 2007. Mammal inventories for eight national parks in the
Southern Colorado Plateau Network. Natural Resource Technical Report NPS/SCPN/NRTR-2007/054. National
Park Service, Fort Collins, Colorado, USA.
Chen, H.-M., F. Zhang, and M.-S. Zhu. 2011. Four new troglophilous species of the genus
Pholcus
Walckenaer
(Araneae, Pholcidae) from Guizhou Province, China. Zootaxa 2922:51–59.
Appendix A-11
RESEARCH REPORTS
Cokendolpher, J. C., and J. R. Reddell. 2001. New and rare nesticid spiders from Texas caves (Araneae: Nesticidae).
Texas Memorial Museum, Speleological Monographs 5:25–34.
Deeleman-Reinhold, C. L. 1993. Description of a new cave-dwelling pholcid spider from northwestern Australia,
with an identifi cation key to the genera of Australian Pholcidae (Araneae). Records of the Western Australian
Museum 16:323–329.
Deltshev, C., and B. P. M. Curcic. 2002. A contribution to the study of the genus
Centromerus
Dahl (Araneae:
Linyphiidae) in caves of the Balkan Peninsula. Revue Suisse de Zoologie 109:167–176.
Dippenaar-Schoeman, A. S., and J. G. Myburgh. 2009. A review of the cave spiders (Arachnida: Araneae) from
South Africa. Transactions of the Royal Society of South Africa 64:5361.
Ferreira, R. L., and R. P. Martins. 1998. Diversity and distribution of spiders associated with bat guano piles in
Morrinho Cave (Bahia State, Brazil). Diversity and Distributions 4:235–241.
Ferreira, R. L., M. F. V. R. Souza, E. O. Machado, and A. D. Brescovit. 2011. Description of a new
Eukoenenia
(Palpigradi: Eukoeneniidae) and
Metagonia
(Araneae: Pholcidae) from Brazilian caves, with notes on their
ecological interactions. Journal of Arachnology 39:409419.
Gertsch, W. J., and S. B. Peck. 1992. The pholcid spiders of the Galápagos Islands, Ecuador (Araneae: Pholcidae).
Canadian Journal of Zoology 70:11851199.
Hedin, M. C. 1997. Molecular phylogenetics at the population/species interface in cave spiders of the southern
Appalachians (Araneae: Nesticidae: Nesticus). Molecular Biology and Evolution 14 (3):309–324.
Makiya, K., and I. Taguchi. 1982. Ecological studies on over winteringpopulations of
Culex pipiens pallens
3.
Movement of the mosquitoesin a cave during over wintering. Japanese Journal of SanitaryZoology 33:335
343.
Miller, J. A. 2005. Cave adaptation in the spider genus
Anthrobia
(Araneae, Linyphiidae, Erigoninae) Zoologica
Scripta 34:565–592.
Northup, D. E., and W. C. Welbourn. 1997. Life in the twilight zoneLava tube ecology, natural history of El
Malpais National Monument. New Mexico Bureau of Mines and Mineral Resources, Bulletin 156:69–82.
Peck, S. B. 1980. Climate change and the evolution of cave invertebrates in the Grand Canyon, Arizona. National
Speleological Society Bulletin 42:5360.
. 1981. The invertebrate fauna of the caves of the Uinta Mountains, northeastern Utah. Western North
American Naturalist 41:201–206.
Reddell, J. R., and J. C. Cokendolpher. 2004. The cave spiders of Bexar and Comal Counties, Texas. Texas Memorial
Museum. Speleological Monographs 6:75–94.
Reeves, W. K., J. B. Jensen, and J. C. Ozier. 2000. New faunal and fungal records from caves in Georgia, USA.
Journal of Cave and Karst Studies 62:169179.
Ruzicka, V. 1998. The subterranean forms of
Lepthyphantes improbulus
,
Theonoe minutissima
and
Theridion
bellicosum
(Araneae: Linyphiidae, Theridiidae). Pages 101–105
in
P. A. Selden, editor. Proceedings of the 17th
European Colloquium of Arachnology, Edinburgh, Scotland, 1997.
Snowman, C. V., K. S. Zigler, and M. C. Hedin. 2010. Caves as islands: Mitochondrial phylogeography of the cave-
obligate spider species
Nesticus barri
(Araneae: Nesticidae). Journal of Arachnology 38:49–56.
Spilman, T. J. 1968. Two new species of
Niptus
from North American caves (Coleoptera: Ptinidae). Southwestern
Naturalist 13:193–200.
Wynne, J. J. 2006. Cave trip report and Junction Cave bat hibernacula, 4 February 2006. Unpublished report
submitted 15 February 2006 to El Malpais National Monument, National Park Service, Grants, New Mexico.
1 page.
Appendix A-12
PARK SCIENCE • VOLUME 30 • NUMBER 1 • SUMMER 2013
... Early on, Barber (1931) and Valentine (1941) favoured baited pitfall traps for capturing omnivorous and carrion beetles due to the quick return rate, while deemphasizing visual searching due to low returns. Because not all arthropods attracted to pitfall traps will be captured, searches around traps prior to removal have been applied by previous workers (Campbell et al., 2011;Martín & Oromí, 1986;Poulson & Culver, 1969;Wynne, 2013;Wynne et al., 2014). For sampling previously unstudied caves in Hawai'i and Australia, Howarth (1980Howarth ( , 1988 applied a variety of techniques including intensive direct intuitive searches (in promising microhabitats), deploying bait stations (using naturally occurring organic material, tubers, rotting meat, cheese and grains) and to a lesser extent baited pitfall traps-the latter included both live and kill pitfalls. ...
... Because caves are highly heterogeneous in their distribution and occurrence of microhabitats, it is not possible to apply an interval sampling approach without either missing or ineffectively sampling areas that may support unique arthropod communities. We encountered moss gardens within entrances and beneath skylights of two (ELMA-0008 and ELMA-0012; Wynne, 2013;Wynne & Shear, 2016) and root curtains from two (ELMA-0303 and ELMA-0315; Wynne, 2013) ELMA caves. Fern-moss gardens occurred within entrances and beneath cave skylights of five of six lava tube caves on Rapa Nui . ...
... For most caves containing moss or fern-moss gardens, we conducted direct intuitive searches (DIS) with an unlimited search radius in which we intensively searched for arthropods by examining moss and ferns, looking on and beneath cobbles and boulders and inspecting cave sediment. For root curtains, we gently searched root masses protruding from cracks within cave ceilings of cave deep zones (Wynne, 2013). DIS were conducted for 1 hr (20 min × 3 observers searching) within each of these habitats. ...
Article
Aim: Identify the optimal combination of sampling techniques to maximize the detection of diversity of cave-dwelling arthropods. Location: Central-western New Mexico; northwestern Arizona; Rapa Nui, Chile. Methods: From 26 caves across three geographically distinct areas in the Western Hemisphere, arthropods were sampled using opportunistic collecting, timed searches, and baited pitfall trapping in all caves, and direct intuitive searches and bait sampling at select caves. To elucidate the techniques or combination of techniques for maximizing sampling completeness and efficiency, we examined our sampling results using nonmetric multidimensional scaling (NMDS), analysis of similarity (ANOSIM), Wilcoxon signed-rank tests, species richness estimators and species accumulation curves. Results: To maximize the detection of cave-dwelling arthropod species, one must apply multiple sampling techniques and specifically sample unique microhabitats. For example, by sampling cave deep zones and nutrient resource sites, we identified several undescribed cave-adapted and/or cave-restricted taxa in the southwestern United States and eight new species of presumed cave-restricted arthropods on Rapa Nui that would otherwise have been missed. Sampling techniques differed in their detection of both management concern species (e.g., newly discovered cave-adapted/restricted species, range expansions of cave-restricted species and newly confirmed alien species) and specific taxonomic groups. Spiders were detected primarily with visual search techniques (direct intuitive searches, opportunistic collecting and timed searches), while most beetles were detected using pitfall traps. Each sampling technique uniquely identified species of management concern further strengthening the importance of a multi-technique sampling approach. Main conclusions: Multiple sampling techniques were required to best characterize cave arthropod diversity. For techniques applied uniformly across all caves, each technique uniquely detected between ~40% and 67% of the total species observed. Also, sampling cave deep zones and nutrient resource sites was critical for both increasing the number of species detected and maximizing the likelihood of detecting management concern species.
... Within the Bandera Volcanic Field of western New Mexico, moss gardens occurring in cave entrances and beneath skylights represent unique and important microhabitats (Lindsay 1951;Northrup & Welbourn 1997;Wynne 2013) in an otherwise xeric landscape (Figs. 5,6). ...
... El Malpais National Monument (ELMA), located in Cibola County, New Mexico, encompasses approximately 1,522 km 2 in the western part of the state. Featuring evidence of at least eight major volcanic eruptions ranging in age from 100,000 to 3,000 years old (Cascadden et al. 1997), the national monument comprises vast expanses of pahoehoe and ʻaʻā lava flows, cinder cones, ice caves, and at least 290 lava tube caves (Wynne 2013). Biological inventories were focused on caves in close proximity to trails and roads and/or known to support sensitive biological resources, including bat roosts. ...
... Today, moss gardens are particularly suitable habitat for these organisms. This habitat supports high biological diversity (Wynne 2013) and unique faunal assemblages, including a presumed relict species of spider, Lepthyphantes turbatrix ((O. Pickard-Cambridge, 1877); Family Linyphiidae; Lightfoot et al. 1994) and A. awishoshola. ...
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Austrotyla awishoshola n. sp. is described from the moss gardens of one lava tube cave in El Malpais National Monument, Cibola Co., New Mexico. Most chordeumatidans require mesic conditions, and these environments are limited to moss gardens in several cave entrances and beneath cave skylights in El Malpais. Presently, this species is known from the moss gardens of a single of cave in the monument. We suggest A. awishoshola may be a climatic relict, having become restricted to the cave environment following the end of the Pleistocene. We discuss the importance of cave moss gardens as refugial and relictual habitats. Recommendations are provided to aid in the conservation and management of A. awishoshola and these habitats.
... nov., discovered within a cave entrance vegetation community. As similar cave entrance vegetation communities have been identified as either supporting distinct relict plant communities and/or plant species in southern China (Monro et al. 2018), Easter Island, Chile ), and west-central New Mexico, USA (Lindsay 1951, Northrup and Welbourn 1997, Wynne 2013, their importance in supporting cave-restricted arthropod populations demonstrated (Northrup and Welbourn 1997, Wynne 2013, Wynne and Shear 2016, and widespread land cover conversion of both lowlands and uplands has occurred in China since 1958, this finding warrants both additional research into this species distribution, as well as a larger scale examination of other potential 'disturbance relict' arthropod species within cave entrance vegetation communities of the SCK. ...
... nov., discovered within a cave entrance vegetation community. As similar cave entrance vegetation communities have been identified as either supporting distinct relict plant communities and/or plant species in southern China (Monro et al. 2018), Easter Island, Chile ), and west-central New Mexico, USA (Lindsay 1951, Northrup and Welbourn 1997, Wynne 2013, their importance in supporting cave-restricted arthropod populations demonstrated (Northrup and Welbourn 1997, Wynne 2013, Wynne and Shear 2016, and widespread land cover conversion of both lowlands and uplands has occurred in China since 1958, this finding warrants both additional research into this species distribution, as well as a larger scale examination of other potential 'disturbance relict' arthropod species within cave entrance vegetation communities of the SCK. ...
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We synthesized the current knowledge of cave-dwelling millipede diversity from Guangxi Zhuang Autonomous Region (Guangxi), South China Karst, China and described six new millipede species from four caves from the Guilin area, northeastern Guangxi. Fifty-two cave-dwelling millipedes are known for the region consisting of 38 troglobionts and 14 troglophiles. Of the troglobionts, 24 are presently considered single-cave endemics. New species described here include Hyleoglomeris rukouqu sp. nov. and Hyleoglomeris xuxiakei sp. nov. (Family Glomeridae), Hylomus yuani sp. nov. (Family Paradoxosomatidae), Eutrichodesmus jianjia sp. nov. (Family Haplodesmidae), Trichopeltis liangfengdong sp. nov. (Family Cryptodesmidae), and Glyphiulus maocun sp. nov. (Family Cambalopsidae). Our work also resulted in range expansions of Pacidesmus trifidus Golovatch & Geoffroy, 2014, Blingulus sinicus Zhang & Li, 1981 and Glyphiulus melanoporus Mauriès & Nguyen Duy-Jacquemin, 1997. As with many hypogean animals in Southeast Asia, intensive human activities threaten the persistence of both cave habitats and species. We provide both assessments on the newly described species’ distributions and recommendations for future research and conservation efforts.
... To accomplish this, we recommend applying a sampling strategy similar to the approach developed by Wynne et al. (2018). If this is not possible, we minimally suggest systematically sampling these caves using baits (Howarth et al. 2007;Wynne et al. 2018), pitfall traps (Wynne 2013;Wynne et al. 2018), and direct intuitive searches (Wynne 2013;Wynne et al. 2018) within cave deep zones. We confirmed the presence of the springtail O. gevorum at four additional caves, which expanded its range by 5.73 km to the east and ~1 km north of Sima Gesm, the type locality. ...
... To accomplish this, we recommend applying a sampling strategy similar to the approach developed by Wynne et al. (2018). If this is not possible, we minimally suggest systematically sampling these caves using baits (Howarth et al. 2007;Wynne et al. 2018), pitfall traps (Wynne 2013;Wynne et al. 2018), and direct intuitive searches (Wynne 2013;Wynne et al. 2018) within cave deep zones. We confirmed the presence of the springtail O. gevorum at four additional caves, which expanded its range by 5.73 km to the east and ~1 km north of Sima Gesm, the type locality. ...
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.
... Para llegar a conseguir este objetivo se recomienda aplicar la estrategia de muestreo desarrollada por Wynne et al. (2018b). Si esto no fuera posible, sugerimos que, al menos, se realice un muestreo sistemático en estas cuevas utilizando cebos (Wynne et al. 2018b), trampas de "pitfall", y búsquedas intuitivas directas (Howarth et al. 2007;Wynne 2013;Wynne et al. 2018b), dentro de las zonas profundas de las cuevas. Se ha confirmado la presencia del colémbolo O. gevorum en cuatro cavidades adicionales, lo que amplía su rango en 5.73 km hacia el este y ~1 km hacia el norte de Sima GESM, que es la localidad tipo, ampliando hasta cinco las localizaciones en la Sierra de las Nieves. ...
Article
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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.
... 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. ...
Article
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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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Abandoned mines provide habitat for bats, but their importance to other wildlife is less understood. This descriptive study was designed to answer the following questions with an emphasis on carnivores: are wildlife species other than bats visiting abandoned mines, is wildlife entering abandoned mines, does wildlife visitation at abandoned mines differ seasonally, and does wildlife visitation differ at individual mines? To address these questions, we monitored 50 abandoned mines using remote cameras in the northern Sangre de Cristo Mountains, Colorado, USA, for 25,201 camera days from May 2017 to August 2020. We monitored mines in 2 phases. During phase 1 (May 2017–May 2019), we monitored 30 randomly selected mines to gather baseline data on carnivore visitation and to model carnivore visitation. During phase 2 (May 2019–August 2020), we monitored 27 mines to test the visitation model and to determine if carnivores visited multiple mines as they traveled across the landscape. We observed >48 species of vertebrates at mines, including 11 of 14 carnivore species known to occur in the Sangre de Cristo Mountains. Carnivores ranged in size from ringtails (Bassariscus astutus) to American black bears (Ursus americanus). Pumas (Puma concolor) visited mines most frequently and we observed pairs of adult pumas entering mines, presumably during courtship and mating.We also observed American black bears, pumas, and common gray foxes (Urocyon cinereoargenteus) visiting and entering mines with young. Carnivores visited mines at low levels throughout the year and visitation differed by season, temperature, and carnivore species, size, and family. Our most parsimonious generalized linear models identified mine elevation, entrance (portal) size, land cover type, tree cover, and aspect as significant predictors of visitation. Our top models explained ≥78% of the variation in carnivore visits and indicated that carnivores in the Sangre de Cristo Mountains were most likely to visit small horizontal mines at lower elevations in dense piñon (Pinus edulis)—juniper (Juniperus sp.) woodlands. We encourage resource managers to monitor abandoned mines for ≥1 year prior to closing or gating mines to understand which wildlife species might be affected by closures.
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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.
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The importance of ants as elements in cave ecology has been mostly unrecognized. A global list of ant species recorded from caves, compiled from a review of existing literature, is presented. This paper also reviews what is currently known about ants occurring in Arizona (USA) caves. The diversity and distribution represented in these records suggests ants are relatively common cave visitors (trogloxenes). A general utilization of caves by ants within both temperate and tropical latitudes may be inferred from this combined evidence. Observations of ant behavior in Arizona caves demonstrate a low level and sporadic, but persistent, use of these habitats and their contained resources by individual ant colonies. Documentation of Neivamyrmex sp. preying on cave-inhabiting arthropods is reported here for the first time. Observations of hypogeic army ants in caves suggests they may not penetrate to great vertical depth in search of prey, but can be persistent occupants in relatively shallow, horizontal sections of caves where they may prey on endemic cave animals. First cave records for ten ant species are reported from Arizona caves. These include two species of Neivamyrmex (N. nigrescens Cresson and Neivamyrmex sp.; Formicidae: Dorylinae), four myrmicines (Pheidole portalensis Wilson, Pheidole cf. porcula Wheeler, Solenopsis aurea Wheeler and Stenamma sp. Westwood), one dolichoderine (Forelius keiferi Wheeler) and three formicines (Lasius arizonicus Wheeler, L. sitiens Wilson, and Camponotus sp. Mayr).
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Populations of cave invertebrates are generally considered to be food-limited. The cave entrance is a major source of food input into the community in the form of decaying organic matter. Thus, the densities of scavenging terrestrial cave invertebrates should be related to the distance from the cave entrance because this represents a measure of food abundance. A test showed this expectation to be true in Crossings Cave, Alabama. A population density peak occurred 10 m inside the cave where the dark zone and detritus infall regions meet. The greatest population peak occurred at 100 m where densities of crickets and their guano are highest. The pattern should hold for most caves, but the actual distances will vary in each site depending on its circumstances. When the fauna was removed from the cave, the remnant had not regained community equilibrium a year later. Removal of the dominant scavenger, a milliped, allowed other species populations to expand because of decreased competitions.
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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.
Conference Paper
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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
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An ecological study was carried out on distribution, movement and change in the body weight of Culex pipiens pallens Coquillett during their overwintering. Overwintering population of Cx. p. pallens was distributed aggregatively in a cave (air-raid shelter). More mosquitoes were found in the inner room of a 3-room shelter and gathered on the lower part of the wall. The degree of such aggregations changed with fluctuation in the temperature and humidity inside the room, and became maximum in January. The movement of mosquitoes was influenced by air temperature : It was least frequent in February when the temperature was lowest. The movement was least frequent on the lower part of the wall. The mobility rate (percentage of mosquitoes which changed their resting positions) was about 70% on average. Mean body-weight of mosquitoes before and after overwintering was 1.271mg and 0.708mg, the reduction rate being 44.3%.
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Five species of spiders from the families Theridiidae (Nesticoides rufipes), Sicariidae (Loxosceles similis and Sicarius tropicus), Oecobidae (Oecobius annulipes) and Corinnidae were found on bat guano piles in the Morrinho cave (Bahia state, Brazil). Species richness of spiders was positively correlated with the area of the guano piles and silverfish abundance, and negatively correlated with the distance from the cave entrance. The positive relationship found between spider richness and diversity with area of the piles is presumably because prey abundance is positively correlated with pile size. The relationship between distance from the cave entrance and spider richness may be due to different colonization abilities of each spider family. Spider diversity was positively correlated only with pile area, while pH of the piles (which may be indicative of age) did not show correlation with any other parameters.
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