Arch. Environ. Contam. Toxicol.
40, 112–120 (2001)
DOI: 10.1007/s002440010153
Cyclodiene Insecticide, DDE, DDT, Arsenic, and Mercury Contamination of
Big Brown Bats (Eptesicus fuscus) Foraging at a Colorado Superfund Site
T. J. O’Shea, A. L. Everette, L. E. Ellison
U.S. Geological Survey, Midcontinent Ecological Science Center, 4512 McMurry Avenue, Fort Collins, Colorado 80525-3400, USA
Received: 23 February 2000/Accepted: 27 June 2000
Abstract. Rocky Mountain Arsenal (RMA) National Wildlife
Area, near Denver, Colorado, is a Superfund site contaminated
by past military and industrial uses, including pesticide man-
ufacturing. From an ecosystem standpoint, the most critical
contaminants at RMA are certain cyclodiene insecticides and
metabolites, p,p⬘-DDE, p,p⬘-DDT, arsenic, and mercury. Bats
are important ecosystem components that can be impacted by
persistent contaminants because of their position in the food
chain and their potential longevity and thus duration of expo-
sure. Big brown bats (Eptesicus fuscus) were captured (n ⫽
51) while foraging at RMA in the summers of 1997 and 1998
for determination of concentrations of contaminants of concern
in carcasses, brains, and stomach contents. Adult females (n ⫽
15) were also tracked by radiotelemetry to determine locations
of nearest maternity roosts for sampling of guano for contam-
inant analysis and inspection for potential contaminant-induced
mortality. Bats captured while foraging at RMA had measur-
able quantities of dieldrin and DDE in masticated insect sam-
ples from stomach contents and significantly higher concentra-
tions of dieldrin, DDE, DDT, and mercury (juveniles) in
carcasses than big brown bats (n ⫽ 26) sampled at a reference
area 80 km to the north. Concentrations of dieldrin and DDE in
brains of bats captured while foraging at RMA were also
greater than in bats from the reference area, but not high
enough to suggest mortality. Maximum concentrations of
DDE, DDT, and cyclodienes in brains of big brown bats were
found in adult males from RMA. Guano from the two closest
known roosts had significantly higher concentrations of diel-
drin, DDE, and mercury than guano from two roosts at the
reference area. Dieldrin concentrations in carcasses of bats
from RMA were highest in juveniles, followed by adult males
and adult females. DDE concentrations in carcasses were low-
est in adult females at both sites and highest in adult males at
RMA. No contaminant-related mortality was obvious at the
small maternity colonies near RMA. Big brown bats show
higher contamination than most other mammals previously
sampled at this site. Concentrations and proportions of samples
with detectable residues of dieldrin in carcasses of big brown
bats from RMA were similar to or exceeded reports for this
Correspondence to: T. J. O’Shea
A R C H I V E S O F
Environmental
Contamination
a n d Toxicology
© 2001 Springer-Verlag New York Inc.
species from elsewhere in the United States some 25 years ago,
prior to or just following restrictions on use of this compound.
Impacts of ecosystem contamination on wildlife from persis-
tent chemicals can be difficult to evaluate without experimental
or comparative information for interpretation of chemical res-
idue data. Rocky Mountain Arsenal National Wildlife Area
(RMA) near Denver, Colorado, is a Superfund site where some
of the major contaminants of concern are cyclodiene insecti-
cides, p,p⬘-DDT (2,2-bis-(p-chlorophenyl)-1,1,1-trichloroeth-
ane), p,p⬘-DDE (2,2-bis-(p-chlorophenyl)-1,1,-dichloroethyl-
ene), mercury (Hg), and arsenic (As). Dieldrin (1,2,3,4,10,10-
hexachloro-6,7-epoxy-1,4,4␣, 5,6,7,8,8␣-octahydro-endo-1,4-
exo-5,8-dimethanonaphthalene) is a major concern at RMA
because it has been found in elevated and sometimes lethal
concentrations in the local fauna (Allen and Otis 1998; Cohn
1999), and in general “is one of the most persistent chemicals
ever known” (Matsumara 1985, p. 58). However, because
dieldrin and other contaminants at RMA have long been rec-
ognized globally as pervasive environmental pollutants, there
are past experimental studies and field investigations that can
facilitate interpretation of data on the occurrence of these
contaminants in the biota.
The degree of contamination by dieldrin and other chemicals
has been determined in a variety of mammals at RMA (e.g.,
Close 1996; Allen and Otis 1998; Hoff 1998; Cohn 1999;
Creekmore et al. 1999), but no information has been available
to evaluate exposure and potential impact of these contami-
nants on bats. Bats are important functional components of
many ecosystems and provide a valuable comparative perspec-
tive on contamination patterns in mammals. They are highly
mobile in comparison with most mammals of their size, feed at
relatively high positions in food webs, and are long-lived. All
of these factors can influence degree of exposure to persistent
contaminants. The objectives of our study, therefore, were to
evaluate contamination of big brown bats (Eptesicus fuscus)
that forage at RMA, assess the geographic distances over
which this single terrestrial source might contaminate colonies
of this species, and to compare contamination of big brown
bats with results of studies of other mammals at RMA and
Contamination in E. fuscus at a Superfund Site
studies of bats elsewhere. Big brown bats were the focus of this
study because they are the most abundant bat known to forage
on the refuge (Everette et al. unpublished data), and there have
been previous studies of contaminants in this species that
provide a basis for comparison of results. These include labo-
ratory studies (Clark and Prouty 1977; Clark and Lamont
1976a) and field surveys of contaminant residues (Clark and
Lamont 1976b; Clark and Prouty 1976; Reidinger 1976). Ex-
perimental and field studies of dieldrin, other cyclodienes,
DDT, and DDE in other species of bats also provide additional
critical comparative information (e.g., Clark et al. 1975, 1978a,
1978b; Geluso et al. 1976; Clark and Stafford 1981; Clark and
Prouty 1984). Previous field studies have been based on col-
lecting bats at daytime roosts near contaminant sources without
direct knowledge of foraging areas. In our study we examined bats
captured while foraging at night at the Superfund location, where
there was no prior knowledge about the proximity of roosting
colonies. To our knowledge, only one other study has attempted to
assess contamination of bats resulting from major industrial pol-
lution by organochlorine (OC) insecticides (Clark et al. 1988).
Materials and Methods
Study Area, Radiotelemetry, and Sample Collection
RMA National Wildlife Area, 16 km northeast of Denver, Colorado, is a
6,900-ha urban, prairie refuge with scattered cottonwood (Populus sar-
gentii) groves, wetlands and impoundments. It was used as a chemical
production site by the U.S. Army from 1942 to 1969, and from 1947 to
1982 pesticides and other chemicals were commercially produced at
RMA (Allen and Otis 1998; Wiley and Rhodes 1998). It was designated
a Superfund site in 1982. The most persistent and potentially harmful
contaminants with respect to wildlife and food-chain effects include
dieldrin, aldrin (1,2,3,4,10,10-hexachloro-6,7-epoxy-1,4,4␣, 5,8,8␣-hexa-
hydro-1,4-endo, exo-5,8-dimethanonaphthalene), endrin (3,4,5,6,9,9-
hexachloro-1a,2,2a,3,6,6a,7,7a-octahydro-endo-dimethanonaphthalene),
isodrin (1,2,3,4,10,10-hexachloro-1,4,4␣, 5,8,8␣-hexahydro-1,4:5,8-endo,
endo-dimethanonaphthalene), chlordane compounds, p,p⬘-DDT, p,p⬘-
DDE, Hg, and As. The severest contamination is at the center of the
refuge (Cohn 1999). The Rocky Mountain Arsenal National Wildlife
Refuge Act of 1992 designated RMA as the largest urban refuge in the
United States, contingent on remediation of contamination (Wiley and
Rhodes 1998; Cohn 1999).
Exposure of bats to persistent food-chain contaminants can be
estimated by sampling guano from roosts (Clark et al. 1982, 1995).
Analyzing brains of bats found dead at roosts can help determine if
exposure to lethal levels of OC insecticides has occurred, particularly
in young exposed to these lipophilic substances while suckling lipid-
rich milk (Clark et al. 1978b, 1980; Geluso et al. 1976). However,
there was no prior knowledge of bat roosts on or near RMA. To
discover locations of maternity roosts, we captured 15 lactating female
big brown bats as they foraged on RMA and affixed miniature (0.5–
0.7 g) radio transmitters (Holohil Systems, Ltd., Carp, Ontario, Can-
ada) to the interscapular area with Skin-Bond colostomy adhesive
(Smith & Nephew United, Inc., Largo, FL). Transmitters and attach-
ments had a projected longevity of 2 weeks. We searched for signals
while driving a 1.6 ⫻ 1.6-km grid of roads on RMA and thoroughfares
in the area. We found buildings with 12 roosts using this technique, but
none were found on RMA. Locations were 5.8 –13.5 km from the
boundary of RMA (up to 18.8 km from point of capture) in urban areas
to the south. The two closest roosts were 5.8 and 6.2 km away at
Fitzsimons Army Medical Center. Guano was collected at these two
113
attic roosts (Fitzsimons Buildings 110 and 404). No fallen young were
discovered, but three juveniles were sampled from a maternity colony
at Building 110. We conducted limited searches for signals from
tagged bats at night to verify that they repeated their foraging at RMA
subsequent to capture.
We collected bats while they foraged at RMA to determine contam-
inant concentrations in carcasses, brains, and stomach contents. We
did not emphasize sampling bats from roosts because the distances
between roost locations and RMA precluded assumptions on where
bats collected at roosts might forage. We also collected big brown bats
at Fort Collins, Colorado, 80 km north of RMA, as a reference area for
comparison. Fort Collins encompasses an urban and suburban high-
elevation prairie area of about 120,000 people, with no known history
of excessive contamination by cyclodienes, DDT, As, or Hg. At Fort
Collins, we collected bats in flight over water and bats and guano at
two maternity roosts. One roost was in an old barn, and the second was
in the attic of a circa-100-year-old home in the city center.
Specimen Collection and Preparation
Big brown bats were collected in flight with mist nets. On RMA, nets
were placed over drainage canals in areas confined by vegetation, over
a small pond, adjacent to vegetation, and at the edges of small
reservoirs. Collecting at RMA took place on 53 nights during the
summers of 1997 and 1998 using 6 –18-m-long mist nets, for a total of
397 6-m net-nights. Captured bats were categorized as adult or juve-
niles born the summer of capture based on ossification of the phalan-
geal epiphyses (Anthony 1988; Burnett and Kunz 1982). Sex was
determined based on external genitalia. To minimize seasonal effects
on OC concentrations in bats (Clark and Prouty 1976), we avoided
sampling in late summer and autumn when fat deposition increases
prior to hibernation, or in spring when fat reserves are depleted. At
RMA we collected 46 bats between 10 June and 7 August 1998 (30 in
July) and 5 bats from 5 June to 14 July in 1997 (3 in July). We
collected 26 bats in Fort Collins from 12 June to 9 August 1998 (22 in
July). Most of the bats from Fort Collins had empty stomachs because
they were taken as they emerged from roosts, preventing comparisons
with stomach contents of bats collected at RMA. We also determined
OC concentrations in three carcasses, two brains, and a pooled milk
sample from stomachs of two of three juvenile big brown bats col-
lected on 9 July 1998 at Fitzsimons Building 110, and a near-term
fetus of a female collected at RMA on 6 July 1998. Hg was also
determined in carcasses of these additional bats, and As was deter-
mined in carcasses of the three juveniles.
Bats were assigned identification numbers, euthanized by cervical
dislocation, wrapped in aluminum foil, and placed in a chest freezer on
the night of capture. They were later thawed and weighed at room
temperature and prepared by dissection following other studies of OCs
in bats (e.g., Clark and Lamont 1976a, 1976b; Clark and Prouty 1976;
Clark et al. 1975, 1988). Wings and feet were removed, the carcass
was skinned, the gastrointestinal (GI) tract (esophagus, stomach, and
intestines) removed and discarded, and the head severed at the base of
the skull. Stomach contents (masticated insects and milk) were re-
moved for analysis. Brains were removed from the cranium. Major
portions of the head musculature were placed with the carcass. Stain-
less steel dissection tools were rinsed in acetone and hexane between
use with each sample. Contact with plastics was avoided. All person-
nel involved with capture and dissection of bats received pre-exposure
rabies immunizations. Protocols for carcass preparation were similar to
most other studies of mammals at RMA (Foster Wheeler Environmen-
tal Corporation 1996).
We sampled guano at roosts because it provides a measure of
contaminant exposure of an entire colony, with a single sample in-
cluding feces from multiple bats deposited over days or weeks (Clark
et al. 1982). Guano was placed in chemically clean jars as it was
114
collected from the surface layer of accumulations. Guano samples
were not oven-dried in the laboratory. Attics and the barn loft were hot
and dry when samples were collected in July 1998, and guano was
presumed to be as uniformly dry as if it had been placed in desiccators,
as observed in some previous studies (Clark et al. 1982; Clark and
Prouty 1976). To test this presumption, we determined percent mois-
ture in 18 guano samples collected simultaneous to those submitted for
chemical analysis. These were stored in sealed jars for ⬍90 days after
laboratory analysis. After a month in desiccant (CaSO4) samples had
lost only 0.79 ⫾ 0.27% moisture (maximum 1.4%).
We attempted to quantify relationships between presumed relative
age of bats and contaminants in samples. However, true age of adults
has not been determined in any study of bats and contaminants. Instead
tooth wear measured as canine tip width (CTW) has been used as a
surrogate index of age (Clark et al. 1975, 1978a; Clark and Prouty
1976; Clark and Lamont 1976a, 1976b). This index has not been
calibrated. Relationships between CTW and incremental lines in den-
tine and cementum in sections of canine teeth have been demonstrated
for big brown bats (Christian 1956). However, these sections were
made on bats of unknown age and reliability of interpretation as annuli
is questionable, although tooth wear can be of some utility as an age
index in bats if used with caution (Anthony 1988). In addition, OCs are
well known to increase with age in some mammals, particularly males,
and concentrations or burdens of OCs correlate with tooth wear in
some other studies of bats (Clark et al. 1975; Clark and Prouty 1976).
We therefore measured CTW to the nearest 0.3 mm on both upper
canines of adult big brown bats with an ocular micrometer and a
dissecting microscope, using CTW of the least worn tooth in calcula-
tions if wear was uneven. We investigated relationships between CTW
and dieldrin and DDE concentrations or total ␮g in carcasses for all
adults and in adult males or females at RMA.
Contaminant Analysis
Samples were analyzed for the following OC insecticides and metab-
olites: aldrin, dieldrin, endrin, isodrin, ␥-chlordane, ␣-chlordane, p,p⬘-
DDE, and p,p⬘-DDT. Other OC pesticides or polychlorinated biphe-
nyls (PCBs) were not investigated. Each carcass, guano, and stomach
contents sample was homogenized and a 1.0 ⫾ 0.1-g aliquot was
extracted with methylene chloride. Brains and the pooled milk sample
were analyzed in their entirety or in approximately 0.1-g aliquots. The
extract was cleaned up by fractionation by gel permeation chromatog-
raphy. Extract was solvent-exchanged into hexane, and analyzed by
dual capillary-column gas chromatography with a Perkin-Elmer Au-
tosystem equipped with two Ni-63 electron-capture detectors and two
fused silica capillary columns. For carcass, stomach contents, and
guano samples detection limits were 0.020 ␮g/g for most cyclodienes
and p,p⬘-DDE, and 0.025 ␮g/g for isodrin and p,p⬘-DDT. Detection
limits for all compounds in the small (0.12– 0.28 g) brain samples were
about 0.1 ␮g/g. Quality control procedures included the use of stan-
dard reference materials, calibration standards, method blanks, solvent
blanks, surrogate standards, and spiked samples. Calibration standards
for all compounds originated from commercially available stock that is
NIST or EPA traceable. Lipid content of samples was determined
gravimetrically. Moisture content was not determined.
Arsenic in carcass and guano samples was determined by graphite
furnace atomic absorption spectroscopy. Sample aliquots were di-
gested with 1:1 nitric acid using microwave-assisted techniques, and
the digestate cooled and diluted to volume. Samples were analyzed on
a Perkin-Elmer Model 4100ZL Atomic Absorption Spectrophotometer
with Zeeman background correction and an As electrodeless discharge
lamp. Minimum reporting level for As was 0.194 ␮g/g.
Hg in carcasses and guano was determined as total Hg by cold vapor
atomic absorption spectroscopy. An aliquot was solubilized with con-
centrated sulfuric and nitric acids and oxidatized at 60°C for 1 h with
T. J. O’Shea et al.
potassium permanganate and potassium persulfate. Hg in the digested
sample was reduced to elemental Hg with stannous chloride and
measured using a Perkin-Elmer Model FIMS-100. Lower limit of
detection for Hg was 0.02 ␮g/g.
For As and Hg, daily calibration solutions, independent reference
standards, and spiked method proficiency samples were routinely
analyzed during quality control procedures. All analyses were per-
formed by PBS&J Environmental Laboratories, Orlando, Florida, un-
der contract to the U.S. Army. Average (⫾ SD) percent recoveries in
spiked laboratory control standards (n ⫽ 11) were 100 ⫾ 9% for
DDT, 82 ⫾ 9% for dieldrin, 71 ⫾ 11% for aldrin, 83 ⫾ 10% for
endrin, 61 ⫾ 7% for ␣-chlordane (n ⫽ 10), 95 ⫾ 21% for As (n ⫽
7), and 95 ⫾ 0% for Hg (n ⫽ 2). Average (⫾ SD) percent recoveries
in spiked bat, stomach, and guano samples (n ⫽ 7, brains were too
small to spike) were 96 ⫾ 11% for p,p⬘-DDE, 82 ⫾ 12% for p,p⬘-
DDT, 77 ⫾ 9% for dieldrin, 73 ⫾ 8% for ␣-chlordane, 75 ⫾ 7% for
␥-chlordane, 86 ⫾ 12% for aldrin, 69 ⫾ 8% for endrin, 97 ⫾ 13% for
isodrin, 52 ⫾ 27% for As, and 86 ⫾ 18% for Hg. Concentrations were
not adjusted for percent recovery.
Statistical Analyses
Residue concentration data were transformed to log10 prior to statis-
tical analysis. Concentrations reported as lower than the detection level
were entered into computations as one-half that level. For each con-
taminant, residue concentrations in guano samples (␮g/g air dry
weight) were analyzed with a balanced analysis of variance (ANOVA)
by location (Fitzsimons or Fort Collins) and roost within location (Fort
Collins barn or attic, Fitzsimons Building 110 or Building 404).
Concentrations (␮g/g wet weight) of OCs in stomach contents of
adults and juveniles were contrasted because of potential bias if
younger bats had milk mixed with masticated insects in stomachs.
Concentrations of individual OCs in carcasses and brains were ana-
lyzed by location (RMA or Fort Collins), and by age/sex category
(adult male, adult female, or juveniles) with two-way ANOVA (Zar
1984, p. 206), including tests for interaction. Because OCs in bat
carcasses can vary with the amount of fat (Clark et al. 1980), the
percent lipid in carcasses was also contrasted by location and age/sex
category using ANOVA. Only the data for As and Hg in carcasses
failed to meet assumptions for ANOVA. The nonparametric Kruskal-
Wallis test (Zar 1984, p. 176) was therefore used to test for differences
in these metals among locations and age/sex categories. Relationships
between tooth wear and OC concentrations were explored using re-
gression, but a lack of correlation (see below) made adjustment of
statistical comparisons of groups by this index of age unnecessary. We
also used regression to examine relationships between DDE and diel-
drin in brains and carcasses as a function of percent lipid in carcass.
All statistical analyses were performed using SAS for Microsoft Win-
dows (version 7, SAS Institute, Cary, NC) with an ␣ ⱕ 0.05 consid-
ered significant.
Results
Contamination Patterns in Guano and Stomach Contents
Dieldrin was significantly higher in guano from the roost in
Building 404 ( p ⬍ 0.001) near RMA, whereas dieldrin was
not detected in other samples (Table 1). Cyclodienes other than
dieldrin were not detected in guano, other than ␣-chlordane at
0.03 ␮g/g and ␥-chordane at 0.05 ␮g/g in guano from Fitzsi-
mons Building 110. Concentrations of DDE were significantly
higher at the roosts nearest RMA ( p ⫽ 0.0005), with no roost
Contamination in E. fuscus at a Superfund Site
Table 1. Geometric means and ranges of concentrations (␮g/g air dry weight) of dieldrin, p,p⬘-DDE, p,p⬘-DDT, arsenic, and mercury in gua-
no of big brown bats (Eptesicus fuscus) from two maternity roosts closest to Rocky Mountain Arsenal (RMA) National Wildlife Area and
from two reference roosts at Fort Collins, Coloradoa
Location and Roost Dieldrin
Near Rocky Mountain Arsenal
Fitzsimons Bldg. 110 ND
(n ⫽ 6)
Fitzsimons Bldg. 404 0.07
(n ⫽ 6) (0.04–0.21)
Fort Collins
Attic (n ⫽ 6) ND
Barn (n ⫽ 6) ND
a
␣-chlordane and ␥-chlordane were also detected at a roost near RMA (see text). ND ⫽ not detected
effect within locations ( p ⫽ 0.57, Table 1). Concentrations of
DDT did not differ between sites ( p ⫽ 0.07), although
maximum concentrations were found near RMA (Table 1).
Concentrations of Hg in guano were higher in roosts near RMA
( p ⫽ 0.003), with a significant roost effect ( p ⫽ 0.009) and
higher means and maxima in Building 404 (Table 1). Arsenic
concentrations in guano were higher in roosts near RMA in
comparison with Fort Collins ( p ⫽ 0.01); however, there was
also a roost effect within locations ( p ⬍ 0.0001). Highest
mean concentrations of As were found in Building 404, but
concentrations were also relatively high at the roost in the Fort
Collins barn (Table 1).
Stomach contents of bats from RMA contained measurable
quantities of dieldrin and DDE, with other cyclodienes and
DDT not detected (As and Hg analyses were not conducted).
Geometric mean (0.05 ␮g/g) and maximum (0.42 ␮g/g) con-
centrations of DDE were slightly higher than dieldrin (0.03
␮g/g geometric mean, 0.24 ␮g/g maximum). The pooled milk
sample from two juveniles collected at the nearest roost had
0.38 ␮g/g DDE, comparable to the maxima in the masticated
insects in stomachs of bats feeding on RMA. There were no
significant differences between adults and juveniles captured
foraging at RMA in DDE ( p ⫽ 0.73) or dieldrin ( p ⫽ 0.95)
concentrations in stomach contents.
Contamination of Bats: Carcasses and Brains
Concentrations of dieldrin in carcasses of bats collected while
foraging at RMA were higher ( p ⬍ 0.0001) than in those
from Fort Collins, the latter being uniformly at or near the
limits of detection (Table 2). Dieldrin concentrations in car-
casses varied by age and sex group ( p ⫽ 0.007) and were
highest in juveniles, followed by adult males (Table 2). The
most contaminated specimens at RMA were an adult male and
an adult female at 2.7 and 2.6 ␮g/g dieldrin, respectively
(Table 2). No statistical interactions occurred between site and
age/sex group in dieldrin concentrations ( p ⫽ 0.28). Concen-
trations of other cyclodienes were lower than the limits of
detection in carcasses from both areas, with the exception of
␣-chlordane at 0.03 and 0.06 ␮g/g in two adult males from
RMA. DDT concentrations in carcasses were higher at RMA
than the reference area ( p ⬍ 0.0001) and varied by age/sex
115
DDE DDT Hg As
0.08 0.04 0.12 1.4
(ND–0.26) (ND–0.33) (0.03–0.34) (0.93–2.4)
0.07 0.02 0.26 2.9
(0.02–0.31) (ND–0.10) (0.20–0.43) (2.2–3.7)
0.02 ND 0.11 0.42
(ND–0.04) (0.10–0.13) (0.21–2.3)
0.02 0.02 0.08 2.5
(0.02–0.03) (ND–0.04) (0.07–0.10) (1.9–3.8)
category ( p ⫽ 0.01) with adult females ⬍ males or juveniles.
No statistical interactions occurred between site and age/sex
group ( p ⫽ 0.21) in DDT concentrations in carcasses. DDE
concentrations in carcasses varied with site ( p ⬍ 0.0001) and
age/sex group ( p ⬍ 0.0001), with concentrations higher in
bats at RMA, lowest in adult females at both sites, and highest
in adult males at RMA (Table 2). The maximum DDE con-
centration in carcasses was 16 ␮g/g in an adult male from
RMA, with maximum DDE in juveniles also ⬎ adult females
(Table 2). No statistical interactions occurred between site and
age/sex group ( p ⫽ 0.60) in DDE concentrations in carcasses.
Hg in carcasses (Table 2) showed significant effects by location
only in juveniles, which were higher at RMA ( p ⫽ 0.003).
Arsenic in carcasses was above detection limits in one adult
female (0.19 ␮g/g) and an adult male (0.21 ␮g/g) from RMA,
and one adult female (0.23 ␮g/g) from Fort Collins.
The proportion of carcasses from RMA with detectable (⬎
0.02 ␮g/g) residues of dieldrin was 100% for juveniles (12 of
12), 96% for males (22 of 23) and 75% for females (12 of 16),
whereas dieldrin was below detection limits in all adults and 7
of 11 juveniles from the reference area. The proportion of
carcasses with greater than 0.1 ␮g/g dieldrin (important for
comparison with other studies) at RMA was 56% for adult
males (13/23), 25% for adult females (4/16), and 75% for
juveniles (9/12). None of 26 bats from Fort Collins exceeded
0.1 ␮g/g dieldrin, whereas the 3 juveniles from Fitzsimons
roosts exceeded this concentration.
The proportion of the carcass as lipids did not differ between
big brown bats from RMA and Fort Collins ( p ⫽ 0.37), but
varied by age/sex category ( p ⫽ 0.022) with adult males
highest (Table 3). No significant correlations were found be-
tween CTW and concentrations or total body burdens of DDE
or dieldrin in carcasses of adult males or adult females from
RMA, Fort Collins, or the combined sample of adults. The only
detectable concentrations of contaminants in the single fetus
from RMA were 0.06 ␮g/g DDE and 0.04 ␮g/g Hg. The
maternal carcass had 1.07 ␮g/g DDE, 0.05 ␮g/g DDT, and 0.09
␮g/g Hg, with no other contaminants detected.
Concentrations of dieldrin in brains of bats collected while
foraging at RMA were higher than in brains of bats collected in
Fort Collins ( p ⫽ 0.038) but did not differ by age/sex cate-
gory ( p ⫽ 0.38). Interaction was found to be not significant
( p ⫽ 0.73). Concentrations of DDE in brains of bats at RMA
116 T. J. O’Shea et al.

Table 2. Geometric mean (GM) and maximum (MAX) concentrations of dieldrin, DDT, DDE, Hg, and As (␮g/g wet weight) in carcasses of
big brown bats (Eptesicus fuscus) from Rocky Mountain Arsenal and Fort Collins, Colorado, summer 1998 (ND ⫽ not detected)
Dieldrin DDT DDE Mercury Arsenic
Location Age/Sex n GM MAX GM MAX GM MAX GM MAX GM MAX
Rocky Mountain Arsenal Males 23 0.13 2.7 0.04 0.28 1.2 16 0.10 0.19 ND ND
Females 16 0.06 2.6 0.02 0.09 0.25 2.9 0.08 0.16 ND 0.19
Juveniles 12 0.22 1.4 0.04 0.73 1.1 3.3 0.04 0.28 ND ND
Fitzsimons Bldg. 110 Juveniles 3 0.20 0.54 0.21 0.24 0.96 1.04 0.03 0.04 ND ND
Fort Collins Males 4 ND ND ND 0.03 0.34 0.63 0.10 0.11 ND 0.23
Females 11 ND ND ND 0.03 0.08 0.58 0.07 0.12 ND ND
Juveniles 11 0.02 0.03 ND 0.04 0.59 1.60 0.01 0.03 ND ND

Table 3. Arithmetic means (⫾1 standard deviation) and ranges (in parentheses) of percent lipid in carcasses and estimated total carcass bur-
dens (␮g) of dieldrin, DDE, and DDT in big brown bats (Eptesicus fuscus) captured at Rocky Mountain Arsenal National Wildlife Area, a
nearby maternity roost, and a reference area near Fort Collins, Colorado
Location Age/Sex n Percent Lipid in Carcass Dieldrin (total ␮g) DDE (total ␮g) DDT (total ␮g)
Rocky Mountain Arsenal Males 23 12.2 ⫾ 6.85 2.9 ⫾ 5.1 17.7 ⫾ 26.3 0.49 ⫾ 0.51
(3.5–28.8) (⬍ 0.1–20.3) (⬍ 0.1–127.4) (⬍ 0.1–2.1)
Females 16 9.7 ⫾ 4.32 2.1 ⫾ 6.3 5.2 ⫾ 7.6 0.20 ⫾ 0.16
(3.3–17.6) (⬍ 0.1–26) (0.3–25.2) (⬍ 0.1–0.6)
Juveniles 12 7.8 ⫾ 2.0 2.6 ⫾ 2.8 9.0 ⫾ 5.6 0.65 ⫾ 1.1
(5.3–11.9) (0.1–9.9) (4.3–22.1) (⬍ 0.1–4.2)
Fitzsimons Bldg. 110 Juveniles 3 7.9 ⫾ 4.3 1.5 ⫾ 1.3 5.5 ⫾ 0.33 1.2 ⫾ 0.14
(2.9–10.5) (0.7–3.0) (5.1–5.7) (1.0–1.3)
Fort Collins, Colorado Males 4 14.3 ⫾ 13.9 0.1 ⫾ 0.02 3.9 ⫾ 2.2 0.1 ⫾ 0.03
(6.9–35.1) (⬍ 0.1–0.1) (0.8–6.0) (⬍ 0.1–0.2)
Females 11 7.2 ⫾ 3.3 0.1 ⫾ 0.01 1.3 ⫾ 1.5 0.1 ⫾ 0.01
(3.8–13.7) (⬍ 0.1–0.1) (0.3–4.7) (0.1–0.15)
Juveniles 11 9.3 ⫾ 3.9 0.1 ⫾ 0.06 4.5 ⫾ 2.6 0.1 ⫾ 0.05
(2.7–18.4) (⬍ 0.1–0.2) (2.4–12.0) (⬍ 0.1–0.2)

were also higher than in brains of bats collected in Fort Collins
( p ⫽ 0.0005) and also differed by age/sex category ( p ⫽
0.019), with highest geometric mean concentrations in adult
males and juveniles (Table 4). Interaction was found to be not
significant ( p ⫽ 0.18). Concentrations of DDT, aldrin, endrin,
isodrin, ␣-chlordane, and ␥-chlordane were below minimum
detectable levels in brains of all bats. Dieldrin was below limits
of detection in brains of all bats from Fort Collins, but was
above detection limits in six bats from RMA, with a maximum
of 0.70 ␮g/g in an adult female (Table 4). DDE was above
limits of detection in 30 of the 51 brains of bats from RMA
with a maximum of 1.6 ␮g/g in an adult male (Table 4),
whereas DDE was above detection limits in brains of just 3 of
the 26 bats from the reference area. The brain of one of the two
juveniles from the nearby maternity roost at Building 110 had
detectable concentrations of DDE (0.45 ␮g/g) and dieldrin
(0.27 ␮g/g), higher than those of juveniles from Fort Collins,
but comparable to those of juveniles captured while foraging at
RMA (Table 4). No significant relationships were found be-
tween percent lipid in carcasses and concentrations of DDE or
dieldrin in carcasses or brains.
Discussion
This study shows that despite their mobility and relatively large
ranges in comparison with terrestrial small mammals, bats that
roost at distances from confined but concentrated sources of
persistent food chain contaminants can bioaccumulate these
substances through exposure during foraging. Radio-tracking
of females showed that by utilizing the Superfund site as a
foraging area, bats from at least 12 colony sites located up to 18
km away were subject to contamination. Concentrations of
dieldrin, DDE, As, and Hg in guano collected at roosts of big
brown bats closest to RMA were significantly higher than those
from roosts in the reference area, implicating RMA as the
source of these contaminants. Comparisons of maximum con-
centrations of contaminants in guano at RMA with other stud-
ies is also instructive. Maximum concentrations of dieldrin in
guano samples from RMA were an order of magnitude higher
than at the reference area and higher than those reported in 7 of
12 previous maxima from other species at other locations,
including studies conducted when the use of dieldrin and aldrin
(which is metabolized to dieldrin) as pesticides was largely
unrestricted (Reidinger 1972; Clark et al. 1975, 1988, 1995;
Clark and Prouty 1976; Reidinger and Cockrum 1978). In the
majority of previous cases where maxima exceeded those re-
ported near RMA, concentrations of dieldrin in guano ex-
ceeded 0.38 ␮g/g, a threshold that, based on a variety of
evidence, is considered to be associated with mortality of bats
(Clark et al. 1978b, 1982, 1988; Clawson and Clark 1989).
This threshold was not exceeded in guano of the big brown bat
colonies we sampled near RMA. However, Clawson and Clark
(1989) have noted that mortality from dieldrin poisoning can
Contamination in E. fuscus at a Superfund Site
Table 4. Concentrations of dieldrin and DDE (␮g/g wet weight) in brains of big brown bats (Eptesicus fuscus) captured at Rocky Mountain
Arsenal National Wildlife Area, a nearby maternity colony (Fitzsimons), and a reference area at Fort Collins, Colorado (ND ⫽ not detected)
Location Age/Sex n GM
Rocky Mountain Arsenal Males 23
Females 16
Juveniles 12
Fitzsimons Bldg. 110 Juveniles 2 0.18
Fort Collins, Colorado Males 4 ND
Females 11
Juveniles 11
occur in some juvenile gray bats (Myotis grisescens) in colo-
nies where dieldrin concentrations in guano may be ⬍ 0.1
␮g/g, a level that was exceeded in big brown bat guano near
RMA.
Maximum DDE concentrations in guano of bats collected
near RMA, although higher than in samples collected at the
reference area, were comparable to reports in other studies
where there has been no indication of associated mortality of
bats. However, most of these were cases where guano was
sampled prior to or just following restrictions on use of the
parent compound DDT (Reidinger 1972; Clark et al. 1975,
1982, 1995; Clark and Prouty 1976; Reidinger and Cockrum
1978; Clawson and Clark 1989). DDE concentrations in guano
from bat colonies with possible DDE mortality or elevated
ecosystem contamination with DDE have all been an order of
magnitude higher than in guano we sampled near RMA (Clark
et al. 1988, 1995). Thus the concentrations of DDE present in
guano from bats roosting near RMA are not elevated enough to
suspect mortality from this chemical (Clark et al. 1982, 1988).
Interestingly, however, maximum concentrations of DDT in
guano from a roost near RMA were higher than previous
reports of DDT in guano of other species of bats elsewhere. An
absence of significant differences between RMA and the ref-
erence site in the other cyclodienes in guano indicates lack of
entry into the food webs utilized by big brown bats at RMA,
perhaps in part due to higher degradation rates in the environ-
ment (e.g., aldrin, endrin). This is also consistent with their
absence in masticated insect samples from stomachs of bats
from RMA. Maximum Hg concentrations in guano of big
brown bat colonies near RMA were higher than those reported
for guano samples from Brazilian free-tailed bats (Tadarida
brasiliensis) from an area in Arizona with ecosystem contam-
ination from Hg released to the atmosphere during smelter
operations (Petit and Altenbach 1973).
Although geometric mean concentrations of OCs in stomach
contents of bats captured while foraging at RMA were uni-
formly low, it is of note that maximum concentrations of
dieldrin and DDE in these samples exceed those reported in
studies of bats in the United States from periods before or
shortly after usage restrictions (Clark et al. 1975, 1983; Clark
and Prouty 1976; Clark and Lamont 1976b). Big brown bats
consume a variety of insects, but small beetles are usually the
major component of their diet (Kurta and Baker 1990; Whi-
taker 1995). It has been shown that beetles at RMA are more
contaminated with dieldrin than other insects and are higher in
OCs than many vertebrates at RMA. The geometric mean
117
Dieldrin DDE
Range GM Range
0.12 ND–0.50 0.23 0.11–1.6
0.09 ND–0.70 0.10 0.10–1.4
0.08 ND–0.23 0.15 0.10–0.67
ND–0.27 0.24 ND–0.45
ND 0.07 0.11–0.17
ND ND 0.08 0.10–0.27
ND ND 0.09 0.10–0.23
concentration of dieldrin in 20 pooled beetle samples collected
at RMA in 1995 was 0.19 ␮g/g (maximum 5.5 ␮g/g), whereas
DDE in beetle samples was about 0.02 ␮g/g (maximum 0.18
␮g/g) (Foster Wheeler Environmental Corporation 1996), only
slightly lower than in stomach contents. Thus, dietary exposure
of big brown bats that forage at RMA and elevated concentra-
tions of dieldrin and DDE in carcasses and brains are consistent
with residue surveys of one of their principal prey groups.
DDE was the only OC insecticide detected in the pooled
milk sample and was lower than or equivalent to results from
past studies elsewhere (Clark and Lamont 1976b). However,
DDE in milk of migratory free-tailed bats in Texas during 1973
was about an order of magnitude higher than in big brown bats
from near RMA (Clark et al. 1975), whereas DDE in milk of
gray bats from northern Alabama was two orders of magnitude
greater (Clark et al. 1988). In the latter case, however, the
source of this DDE may have been an unusual case of regional
industrial contamination (O’Shea et al. 1980; Clark et al.
1988). In our study dieldrin was not detected in milk. This is
typical for bats, except for gray bats from Missouri during the
1970s, when elevated dieldrin concentrations from regional
agricultural uses were clearly associated with direct mortality
(Clark et al. 1978b, 1983). The presence of DDE in milk and
the patterns of dieldrin, DDT, and DDE concentrations in
carcasses by age/sex categories (higher in adult males and
juveniles than in adult females, see below) imply lactational
transfer of these OCs, and the single fetus demonstrated trans-
placental transfer. These mechanisms of OC transfer to young
have also been demonstrated in other studies of bats (e.g.,
Clark et al. 1975; Clark and Lamont 1976a, 1976b; Clark 1978;
Clark and Krynitsky 1978; Thies and McBee 1994).
Patterns of contamination in guano and stomach contents of
big brown bats were also seen in carcasses and brains. Maxi-
mum dieldrin concentrations in carcasses of big brown bats
collected foraging on RMA are typically one to two orders of
magnitude higher than in those from Fort Collins and exceed
previous maxima for dieldrin in carcasses of this species from
elsewhere in the United States in the late 1960s and 1970s
(Reidinger 1976; Clark and Lamont 1976a, 1976b; Clark and
Prouty 1976; Henny et al. 1982) prior to or just following
restrictions on use of this compound. Maximum concentrations
of DDE and DDT in carcasses of big brown bats from RMA
were generally one to two orders of magnitude higher than in
big brown bats from the reference area, but with the exception
of an elevated case from Arizona (Reidinger 1976), were
within an order of magnitude of those reported for this species
118
in the 1970s (Clark and Lamont 1976a, 1976b; Clark and
Prouty 1976; Henny et al. 1982). The only other compound
detected in carcasses at RMA was ␣-chlordane. Traces of this
compound have been previously reported in a few individuals
of this species at two other U.S. localities (Clark and Prouty
1976; Clark and Lamont 1976b).
The proportions of big brown bats sampled at RMA with
detectable residues of dieldrin in carcasses are similar to or
exceed reports for this species from elsewhere in the United
States about 25 years ago, prior to or just following restrictions
on use of this compound. Henny et al. (1982) found none of 25
big brown bats sampled in eastern Oregon from 1975–1977
with ⬎0.1 ␮g/g dieldrin, whereas proportions of samples ⬎0.1
␮g/g at RMA were similar to those found in 1973 in Maryland
and West Virginia (Clark and Lamont 1976b; Clark and Prouty
1976). Geometric means and maxima for dieldrin and geomet-
ric means for DDE in carcasses of big brown bats from RMA
were also higher than in 81 pipistrelles (Pipistrellus pipistrel-
lus) from agricultural areas of Spain in 1988 –1990, about
15–20 years after bans or restrictions on dieldrin and DDT in
that country (Guillén et al. 1994). Dieldrin concentrations in
big brown bat carcasses from RMA were also higher than or
comparable to dieldrin in carcasses of 20 Schreiber’s bent-
winged bats (Miniopterus schreibersi) and 19 greater horse-
shoe bats (Rhinolophus ferumequinum) from five localities in
Spain collected over the same period (Hernández et al. 1993).
Although concentrations of dieldrin and DDE in carcasses
and brains of big brown bats at RMA are elevated, they do not
suggest lethality. Unfortunately, there are no data on sublethal
effects of these contaminants on bats, so effects such as endo-
crine disruption and immunosuppression cannot be addressed
here. Diagnostic lethal concentrations of dieldrin in brains of
gray bats have been estimated, with a lower 95% confidence
limit for geometric means at 6.7 ␮g/g (Clark et al. 1978b,
1980). For DDE, this limit has been established experimentally
as 519 ␮g/g in little brown bats (Myotis lucifugus) (Clark and
Stafford 1981) and 483 ␮g/g in Brazilian free-tailed bats (Clark
and Kroll 1977), with estimates for DDT at 18.1 ␮g/g in brains
of little brown bats (Clark et al. 1978a). Maximum dieldrin
concentrations were about 10% of diagnostic lethal concentra-
tions in our sample of 51 brains. Maximum concentrations of
DDE in brains were less than 1% of lethal values, and DDT
was not detected (Table 4). Dieldrin is the major contaminant
of concern at RMA and was the compound found closest to
lethal concentrations in bat brains at this site. Occasional
mortality of big brown bats from dieldrin poisoning at RMA
cannot be entirely ruled out. Our sample was based on presum-
ably healthy bats captured while actively foraging, and it is
conceivable that individuals with much greater exposure may
die at unknown roost locations and escape detection by our
sampling methods. Bats with low body fat would be at most
risk. In general, this would be at the end of hibernation or in
individual bats forced to lose fat by disease, disturbance of
torpor cycles, or other factors. However, based on our sam-
pling, these would be unrepresentative of most of the big
brown bats that utilize RMA as a foraging site. A low likeli-
hood of mortality based on our sample is also supported by
experimental findings on captive little brown bats. These bats
were fed 0.38 ␮g/g wet weight dieldrin for up to 52 days
without reaching lethal brain concentrations, and only 6 of 47
had measurable dieldrin in brains, ranging from 0.51 to 1.3
T. J. O’Shea et al.
␮g/g (Clark and Prouty 1984). The maximum concentration
reached in carcasses in this nonlethal feeding study was 2.5
␮g/g wet weight. Dosages and residues were comparable to or
lower than those we observed in stomach contents, brains, and
carcasses of big brown bats at RMA. Guano of captive little
brown bats fed dieldrin at 0.38 ␮g/g ranged 0.08 – 0.17 ␮g/g
dieldrin (Clark and Prouty 1984), similar to the ranges we
detected in guano samples from a roost near RMA.
Bats in our sample also had sufficient lipid reserves (Table 3)
to prevent mobilization of significant quantities of DDE into
the brain at the season in which they were collected. Mobili-
zation may not occur in bats until decreasing carcass lipids
approach about 2–3% or less of carcass weight (Clark and
Kroll 1977; Clark and Prouty 1977, 1984). In addition, total
amounts of DDE in carcasses suggest that even under extreme
mobilization of fat reserves quantities present in big brown bats
at RMA available to enter the brain are insufficient to cause
mortality. Based on feeding studies, captive big brown bats
were estimated to require mobilization of about 148 –171 ␮g of
DDE into the brain to cause death; the amount of DDE in the
carcass associated with this level in the brain at low carcass
lipid concentrations was about 6,000 –7,000 ␮g (Clark and
Prouty 1977; Clark and Krynitsky 1983). The maximum body
burden of DDE in our sample of big brown bats was about 2%
of this amount and less than the total required to reach the brain
to achieve lethal levels. The modest concentrations of dieldrin
and DDE probably also contribute to the lack of correlation of
dieldrin or DDE in carcasses or brains with the proportion of
lipid in carcasses. Such correlations have been found in exper-
imental samples with ranges from about 2 to 90 ␮g/g (wet
weight) dieldrin in carcasses (Clark et al. 1980), 500 to 17,000
␮g/g (lipid weight) or 6,000 to 14,000 total ␮g DDE in car-
casses (Clark and Prouty 1977), and 1,200 to 71,000 ␮g/g
(lipid weight) DDE in carcasses (Clark and Stafford 1981).
Modest dieldrin and DDE concentrations coupled with a nar-
row range of tooth wear (0.13 to 0.64 mm) in big brown bats
from RMA probably also contributed to lack of correlation
between CTW and concentrations of these contaminants in
carcasses. Lack of correlation with CTW and dieldrin or DDE
concentrations in carcasses has also been noted in some other
studies of bats (Clark et al. 1975; Clark and Prouty 1976; Clark
and Krynitsky 1978).
Contamination of bats is generally higher than that reported
for other mammals at RMA. Dieldrin concentrations have been
determined in carcasses of deer mice (Peromyscus manicula-
tus) from RMA collected at areas of highest soil contamination,
and in areas of very low and intermediate contamination (Allen
and Otis 1998). These data were expressed as the proportion of
deer mice collected at each of two sites in these categories that
exceeded 0.015 ␮g/g. These proportions were 0.000 at a ref-
erence site (n ⫽ 3 mice), 0.125 and 0.222 at intermediate sites
(n ⫽ 26), and 0.600 and 0.765 at the highly contaminated sites
(n ⫽ 22). These carcasses included skins, which could also be
contaminated by residues in soil. DDE was not detected in
these deer mice (Allen 1996). The proportion of big brown bats
collected while foraging at RMA that exceeded 0.020 ␮g/g
dieldrin (our minimum detection limit) in carcasses was 0.902
(46/51, all age and sex categories combined), whereas those
from our reference area was 0.269 (7/26). Although not limited
to foraging at small core areas, a higher proportion of big
brown bats showed contamination than terrestrial deer mice
Contamination in E. fuscus at a Superfund Site
from very contaminated sites. Similarly low concentrations of
dieldrin were reported in deer mice, cottontail rabbits (Sylvila-
gus sp.), and pocket gophers (Geomys sp.) from RMA (Frank
1997). Pocket gophers are completely fossorial, but just 22% of
the brains of those captured in the most contaminated soils at
RMA had detectable concentrations of dieldrin, and none had
detectable residues of DDE (Close 1996). Furthermore, 70% of
the pocket gopher carcasses from these sites had detectable
residues of dieldrin with the highest reported on any site less
than half the maximum in big brown bats. Lower proportions
of samples with detectable concentrations of dieldrin and low
maxima were also reported for kangaroo rats (Dipodomys
ordii), grasshopper mice (Onychomys leucogaster), and prairie
voles (Microtus ochrogaster) from the most contaminated ar-
eas of RMA (Allen 1996).
Large mammals at RMA are also generally less contami-
nated than big brown bats. Dieldrin was detected in just 1 of 18
brains of deer (Odocoileus virginianus and Odocoileus hemio-
nus) (Creekmore et al. 1999), at ⬍ 3% of the maximum
concentration found in brains of big brown bats. Dieldrin was
detected in fat of 9 of 17 deer, with a maximum of 0.72 ␮g/g,
whereas the maximum dieldrin concentration in carcass lipids
of big brown bats was 22 ␮g/g. Badgers (Taxidea taxus) and
coyotes (Canis latrans), both carnivores, are the only mammals
at RMA that thus far can show higher contamination with
dieldrin. At least two badgers were fortuitously found dead
with dieldrin concentrations in adipose tissue of 29 and 75 ␮g/g
and up to a potentially lethal 5 ␮g/g in the brain (Hoff 1998).
One coyote was reported with a concentration of 2.7 ␮g/g
dieldrin in the brain (U.S. Fish and Wildlife Service 1997).
However, the greater likelihood of discovering badger or coy-
ote carcasses than those of bats may bias this comparison. For
example, the highest concentrations of dieldrin in 36 adipose
tissue biopsies of 29 free-ranging badgers (more comparable to
free-flying bats) at RMA from 1994 –1996 was 7.6 ␮g/g, with
11 of 33 (33%) samples and 8 of 29 (28%) individuals below
limits of detection (Hoff 1998). Dieldrin concentrations in
brains and adipose tissue of five radio-tagged badgers which
died from various unrelated factors in the mid-1990s were
lower than in big brown bats (Hoff 1998). Unlike the preva-
lence of contamination in big brown bats, DDE was not de-
tected in the brains of these badgers, and reached a maximum
of 0.039 ␮g/g in adipose tissue (Hoff 1998). DDE was also not
detected in brains of three coyotes found dead in 1995 (U.S.
Fish and Wildlife Service 1997). Departure from the pattern of
higher concentrations of some contaminants in bats than in
other mammals is only evident where data were from terrestrial
species (primarily deer mice and prairie dogs, Cynomys ludovi-
cianus) collected in 1988 –1990 on specific sites at RMA
thought a priori to be most highly contaminated (Stollar and
Associates 1992). Even in that comparison, maxima and fre-
quency of occurrence of DDE, DDT, and Hg were higher in big
brown bats than in prairie dogs and deer mice. Dieldrin was
also lower in prairie dogs, but concentrations of dieldrin in deer
mouse samples were equivalent to or higher than in bats. Thus,
depending on the nature of the comparisons, our overall con-
clusion is that big brown bats are generally more contaminated
with dieldrin and DDE than other mammals that inhabit RMA.
This can be attributable to long life spans (to at least 19 years
[Kurta and Baker 1990]), their position in the food chain,
relative contamination of their most likely prey (Coleoptera),
119
and their high food intake rates needed to sustain metabolic
requirements of flight and build up fat reserves for hibernation
(Clark 1988). Although the potential for contamination is fa-
cilitated by these life-history factors, it is remarkable that
contaminants from a local but intense source of industrial
pollution are detectable in big brown bats and their guano,
despite long distances to roosts that doubtless dilute exposure
by facilitating foraging at other locations.
Acknowledgments. L. Stone, J. McCance, and K. Castle, Midconti-
nent Ecological Science Center (MESC) helped in the field. C. Henry,
J. Griess, M. Sattelberg, and S. Skipper of the U.S. Fish and Wildlife
Service facilitated activities at RMA. P. Smith of MESC assisted with
data organization and office support. For manuscript review we thank
D. Clark Jr. and M. Sattelberg. J. Wimsatt served as consulting
veterinarian. The Griffiths and Doster families in Fort Collins and the
Fitzsimons Army Medical Center near Denver provided access to bat
colonies in their buildings. Specimens were collected under Colorado
Division of Wildlife permit TR-738.
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