Tracking Wildlife 00 Us Fi Rich

Tracking Wildlife by Satellite: Current Systems and Performance
UNITED STATES DEPARTMENT OF THE INTERIOR FISH AND WILDLIFE SERVICE

Fish and Wildlife Technical Report 30
Washington, D.C.
1990
Fish and Wildlife Technical Report

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From
the collection of
International
Bird Rescue Research Center

Cordelia, California
in association with
7 z
m
p
Prejinger
v
Juibrary
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ISSN 0899-3505
Tracking Wildlife by
Satellite:
Current Systems
and Performance
By Richard
B. Harris, Steven G. Fancy, David C. Douglas, Gerald
F.
W. Garner, Steven C. Amstrup, Thomas R.
McCabe, and Larry
Pank
UNITED STATES DEPARTMENT OF THE INTERIOR FISH AND WILDLIFE SERVICE

Fish and Wildlife Technical Report 30
Washinston, D.C.
1990
Contents
Page
Abstract
1
Overview of Argos
Performance
Reliability
in
2
7
Gathering Wildlife Data
7 9
12
21
Efficiency Precision and Accuracy of Locations
Long-term Activity Index

Short-term Activity Index Temperature Sensor
Saltwater Sensor
21
27
28 29
Argos's Location Class Zero (LCO) Service Local User Terminals (LUT's)
30 30
31
Overview and System Description Performance

Cost Comparison
Field Studies
32 32 32 32 34 34 34
35 35
Caribou: Northern Alaska and Yukon

Polar Bear: Beaufort Sea Polar Bear: Chukchi Sea and Bering Sea Muskox: Arctic Slope
Muskox: Greenland
Brown Brown
Bear: Western Brooks Bear: Kodiak Island
Range
Dall Sheep: Brooks Range Elk: Yellowstone National Park
36
39 39
Mule Deer: Southeastern Idaho

Moose: South-central Alaska
Wolf: Northwestern Alaska
Walrus: Bristol Bay, Alaska Application and Sampling Considerations
40 40 42
43 43
Data Processing Sampling Concerns

Cost Comparisons
Directions for Future Research
44 46 47 49 50 50 52
Summary and Conclusions

Acknowledgments
References
Glossary
Tracking Wildlife by
Satellite:
Current Systems and Performance
by
Richard B. Harris
Montana Cooperative
Wildlife
Research Unit
University of Montana
Missoula, Montana 59812
Steven G. Fancy Alaska Fish and Wildlife Research Center 101 12th Avenue, Box 20 Fairbanks, Alaska 99701
David C. Douglas Alaska Fish and Wildlife Research Center 1011 East Tudor Road
Anchorage, Alaska 99503
Gerald W. Garner
Alaska Fish and Wildlife Research Center 1011 East Tudor Road

Steven C. Amstrup
Alaska Fish and Wildlife Research Center 1011 East Tudor Road
Thomas
R.
McCabe
Alaska Fish and Wildlife Research Center 101 12th Avenue, Box 20 Fairbanks, Alaska 99701
and
Larry F. Pank Alaska Fish and Wildlife Research Center 1011 East Tudor Road
ABSTRACT.
Since 1984, the U.S. Fish and Wildlife Service has used the Argos Data Collection and
Location System (DCLS) and Tiros-N series satellites to monitor movements and activities of 10 species of large mammals in Alaska and the Rocky Mountain region. Reliability of the entire system
was generally
high. Data were received from instrumented caribou (Rangifer tarandus) during 9 1
% of
318 possible transmitter-months. Transmitters failed prematurely on 5 of 45 caribou, 2 of 6 muskoxen (Ovibos moschatus), and 1 of 2 gray wolves (Canis lupus). Failure rates were considerably
FISH
AND WILDLIFE TECHNICAL REPORT 30
higher for polar (Ursus maritimus) and brown (U. arctos) bears than for caribou (Rangifer tarandus). Efficiency of gathering both locational and sensor data was related to both latitude and topography.
Mean
animals;
error of locations
was estimated
to be
954
m (median = 543
m)
for transmitters
on captive
90%
of locations were <1,732
from the
true location. Argos's
new
location class zero
processing provided many more locations than normal processing, but mean location error was much higher than locations estimated normally. Locations were biased when animals were at elevations other than those used in Argos's calculations.
Long-term and short-term indices of animal

species, the long-term index
activity
was
correlated with
movement
were developed and evaluated. For several patterns and the short-term index was
calibrated to specific activity categories (e.g., lying, feeding, walking).
reliable
Data processing and sampling considerations were evaluated. Algorithms for choosing the most among a series of reported locations were investigated. Applications of satellite telemetry data
and problems with lack of independence among locations are discussed.
Biotelemetry techniques are used to locate and obtain physiological and behavioral data from free-ranging ani-
sheep (Ovis
dalli) in
mule deer (Odocoileus hemionus)

region.
Alaska and elk (Cervus elaphus) and in the Rocky Mountain
mals and to advance our understanding and management of wildlife. Biologists commonly use radio-tracking
high frequency (VHP) range of the electromagnetic spectrum. However, limited reception range is a drawback of conventional VHP equip-
To provide the reader a more complete understanding of

the
equipment
that operates in the very
Argos system and
we have
the
its applications to tracking animals, included an updated and shortened overview of
Argos system as presented by Fancy
et al. (1988).
ment, particularly for species that

inhabit remote or
is
move
long distances or
Our experience has primarily been with

restricted to those species.
large
mam-
mountainous
areas.
Adequate sampling
malian herbivores and carnivores, and our conclusions are
often constrained by the high cost of locating the animal,
by problems with weather conditions, darkness, safety considerations, and extensive animal movements. The use
of satellites for locating animals and obtaining other data from them has become available with the recent technolo-
Mate (1987) provided a comof experiences using satellite telemetry on various pilation cetaceans. Other researchers have used lightweight, solarpowered
et al.
satellite transmitters to track large birds (Fuller
al.
1984; Strikwerda et
1985, 1986).
The use of
gy
to construct accurate
to
and
reliable transmitters small
satellites to obtain data
enough
be attached to animals.
ing rapidly.
New
on free-ranging animals is expandtechnology and improved equipment are
This report summarizes two years of research and development of satellite telemetry for large mammals by the
continually being developed. However, we have reported the most recent advances with which we are familiar for
Alaska Fish and Wildlife Research Center

the U.S. Fish
(AFWRC)
of
use by prospective users of the technology.
and Wildlife Service, working in conjunction with the Alaska Department of Fish and Game (ADFG),
Arctic National Wildlife Refuge (ANWR), Idaho Department of Fish and Game, Yellowstone National Park, Cana-
Overview of Argos
The Argos Data Collection and Location System (DCLS) is a cooperative international project of the Centre National d'Etudes Spatiales (CNES) of France, the National Oceanic and Atmospheric Administration (NOAA), and the National Aeronautics and Space Administration (NASA). The primary purpose of Argos is to
collect environmental data (e.g., meteorology, hydrology,
dian Wildlife Service,
Yukon Department of Renewable
Resources, the University of Idaho, and the University of Alaska (Institute of Arctic Biology and Alaska Cooperative Wildlife Research Unit [ACWRU]). Service Argos
(referred to hereafter as Argos)
and Telonics,
Inc.
(Mesa,
Arizona), have participated
in the
technology.
We
present results
development of this of our studies on the re-
liability, accuracy, and precision of the system and on developments in sensor technology and local user termi-
oceanography, ecology). The system consists of transmitters on ocean buoys, glaciers, animals, and other places;
nals (LUT's).
We
Argos system
to obtain locational
summarize our experiences using the and behavioral data on
polar bears (Ursus maritimus), caribou (Rangifer tarandus), muskoxen (Ovibos moschatus), brown bears (Ursus
arctos), gray
equipment on polar-orbiting Tiros-N satellites (currently NOAA- 10 and NOAA- 1 1 ) that receive signals from transmitters during <28 overpasses each day; and a network of
satellite tracking stations and ground and satellite communication links that transfer satellite data to processing cen-
wolves (Canis lupus), moose (Alces alces), Pacific walrus (Odobenus rosmarus divergens), and Dall
ters that distribute results to users
(Argos 1984).
TRACKING WILDLIFE BY SATELLITE
operates a network of satellites for providing data on the earth's environment on a daily basis. global The primary mission of Tiros-N satellites is to obtain data
for weather forecasting. Satellites are launched at an ap-
NOAA
movement of
the source
and receiver. The frequency

satellite is
re-
ceived by instruments on the
higher than the
transmitted frequency (401.650 MHz) as the satellite approaches the transmitter, but becomes lower as it moves
proximate rate of one per year to maintain continuous operation. Additional satellites will enable the program to
continue into the future.
away from the transmitter (Fig.
2).
When the received and
The
near-polar, sun-synchronous orbit of the Tiros-N
transmitted frequencies are equal (the inflection point of the Doppler curve), the position of the transmitter is perpendicular to the satellite ground track. Normal processing by Argos requires four transmissions during an overpass to estimate a location.
images of a particular area to be acquired at approximately the same local solar time each day. To maintain sun-synchronous operation, the orbital plane of the satellite must revolve, or precess, about the earth's
series allows
Each Doppler measurement produces two possible positions for the transmitter that are
symmetrical with re-
polar axis in the same direction and at the same average rate as the earth's annual revolution around the sun. Differ-
spect to the satellite ground track.
two positions
is
The more likely of the determined from previous locations,
ences in the altitudes of the two orbits ensure that the same
is not viewed simultaneously by both Because of the earth's rotation during the approximately 102 min of each orbit, two successive satellite
transmitter velocity, and the earth's rotation. For slow-
location on earth
satellites.
moving
95%
transmitters, the ambiguity can be resolved in of the cases (Argos 1978). Location accuracy is influenced by several factors in-
ground tracks are separated by 25 longitude at the equator, the second ground track being to the west of the first.
cluding the stability of a transmitter's oscillator, the elevation of the transmitter, ionospheric propagation er-
The
satellite orbits are inclined
approximately 98 to the
equatorial plane (8 to the polar axis), so the ground tracks of two successive passes cross each other at 82 latitude,
and errors in satellite orbital data (Le Traon 1987). Errors resulting from differences in actual transmitter elevation and assumed transmitter elevation occur primarily
rors,
and both poles are "seen" by the
satellite
during each
overpass. Therefore, the number of passes over a given location each day is a function of latitude, ranging from 6
in the longitudinal plane. The magnitude of transmitterelevation error also depends on the maximum elevation of
per day over a site on the equator to 28 per day higher than 82 (Fig. 1).
at latitudes
the satellite during the pass (French 1986; Table 1). In April 1987, Argos began categorizing locations
by location quality (LQ) indices. Indices range from

is
The
location of a transmitter
its
is
estimated from the Dop-
pler shift in
carrier frequency.
The Doppler effect
the
perceived change in frequency resulting from the relative
to 3, with 3 being the highest-quality location. Table 2 shows the expected standard deviation of a cluster of locations for LQ 1 to 3 as well as the criteria used in
Fig. 1. Relation between latitude of a study area and the degree of coverage by the
two
satellites
(from Argos 1984).
40
60
Latitude
FISH
Satellite
Orbit
Fig. 2. Doppler shift in frequency as the satellite approaches and then moves away from a
PTT. The slope of the Doppler curve
at the
inflection point determines the distance of the animal from the satellite's ground track.
401.6
Transmitted
Received
Frequency
Frequency
401.6500 MHz
lower than fT
401.640
Doppler Curve
determining the index for each location.

In January 1988,
life
Argos
initiated a
new service for wild-
researchers called location class zero (LCO). In this
special processing, locations are calculated from as few as two Doppler measurements. For all overpasses in which a location fix from LCO processing is obtained, data appear in files separate from those obtained through normal
Argos processing. These locations are generally of lower quality but may still be useful for some wildlife applications.
LCO
processing also contains records for each
normally calculated location and provides the alternate
Table
Effect of
maximum
pass, and
the difference
satellite elevation during a between the assumed and ac-
tual platform transmitter terminal
(PTT) elevation, on
location accuracy of a large, well-insulated transmitter
(adapted from French 1986).

Elevation
error (m)
Table
Location quality indices (LQ) and their precision according to Argos. Precision is the standard deviation of the distribution of locations that is, 68% of a series of locations would be expected to fall within this distance (adapted
2.
from LeTraon 1987).
LQ
FISH
Table
4. Specifications
of Argos-certified transmitters
built
by Telonics,
Inc., for terrestrial
mammals.
Transmitter generation
Specification
30
25
NE Alaska
ing length of operation in order to gather intensive data during a particular season. Alternatively, it might be desirable to reduce the
number of animal recaptures
to replace
20
15
the collar, resulting in a duty cycle that sacrifices
number
of locations but extends battery life. Cycling periods that are integer multiples of 24 h will result in locations being
10
5
obtained at approximately the same time each day. For some objectives, this may compromise the randomness or
independence of the sample (Swihart and Slade 1985a). The minimum number of hours of transmission needed
to ensure a location estimate
teristics
lite
depends on
(e.g.,
latitude, charac-
of the
satellite
overpasses
maximum
satel-
NOAA-11
Maine
elevation), and characteristics of the study animal (e.g., its behavior and habitat). Our experiences suggest that a location estimate from PTT's on terrestrial species
NOAA-10
can be expected from about half the satellite overpasses that have a maximum elevation over the study area of 15
or greater.
Performance
Gathering Wildlife Data
in
The following sections present results from nearly 1,000 PTT-months (1 PTT month = 1 PTT operating 1 month) involving 10 mammalian species. First, we summarize
PTT survival
rates.
Second,
we
discuss efficiency
in obtaining locations
and sensor information among
PTT's operating normally. Next, we explore the precision and accuracy of locations obtained through the Argos system, and then we present results of our calibration experiments with activity indices and application of these
experiments to free-ranging caribou.
Reliability
8 10 12 14 16 18
Hour (UT)
2022
PTT
jects
Survival Rates
Generalizing
PTT survival rates across species and probecause duty cycles differed, resulting
Here,
Fig. 3. Satellite coverage at three representative sites in North
was
difficult
America. UT =
Universal Time.
in differing life expectancies.
we
differentiate be-
tween a failure to record locations, which we term location failure, and a failure to receive any data at all, which we computer program. Given a
set
satellite prediction
of orbit
term message
failure.
We
do not
differentiate
among
the
data as a starting point in their calculations, such programs calculate times and characteristics of satellite overpasses.
many
ibou,
possible causes of failure but believe that most
failures
were due
to
premature battery depletion. For car-
The
accuracy of pass predictions decreases as the time

orbit data (available
we used duty cycles that theoretically provided a life Of 45 PTT's deployed

before or after
between the known
from
NASA)
and
expectancy of one year. Reliability of caribou PTT's was

high.
the predicted orbits increases.
However,
predictions six
March 1987,
months
into the future introduced an error of only 3
min
(Fancy et al. 1988). Other considerations for a duty cycle must be dictated by study objectives. For example, it may be worth sacrific-
only 5 failed: 3 experienced message mediately, 1 failed within 3 months, and another failed
within 8.5 months. All other collars functioned for a
full
failures almost im-
year and were
still
operating
when removed. As of
late
FISH
May
1988, 289 PTT-months of data were
removed from
possible 318 PTT-months (91%). We do not know why this group of PTT's had a lower survival rate: not 1 of the
Eleven PTT's were deployed on brown bears in Alaska during summer 1987. Eight of these were expected to
function through May 1988, and three were expected to function through September 1988. One bear shed its collar
in August 1987. No data were received following den entrance from 9 of the 10 remaining bears. We initially assumed that PTT signals were blocked by the dens. How-
10 collars deployed in March 1987 lasted for a full year, and mean operating time for this group of PTT's was
approximately 33 weeks. One of four deployments on muskoxen in northern
Alaska experienced message failures almost immediately. It was redeployed after being refurbished and has since
operated continuously for 12 months.
ever, following
den emergence, we received data from
One
of two muskfail-
only one PTT and it ceased functioning within 3 weeks. Three of the eight PTT's were designed to cease transmitting during
oxen collared
in
Greenland experienced a location
denning but resume
ure just after the investigator departed. This PTT was deployed on a bull, and we suspect that abuse incurred
programming
in spring. An error in the duty cycles for these three prevented
during the rut was responsible for
its failure.
This
PTT
after
resumption of transmissions. Between spring 1985 and spring 1988, 109 PTT's were
began operating again

deployment.
by
itself
some
months
deployed on polar bears in the Beaufort, Bering, and Chukchi seas (Garner et al. 1989). Five models of PTT's
powered by C-size batteries and was deployed on a wolf in the ANWR; it failed after 1 .5 months. However, a similar PTT placed on another wolf transmitted for 15 months, 7 months longer than expected. As of October 1988, none of the PTT's deployed on elk, mule deer, or Dall sheep had failed before expected battery depletion. One of two PTT's deployed on moose experienced message failure after apfirst
1
The
small
PTT
were used. Versions
A and B of second-generation PTT's

C of third-
weighing
.2
kg
differed in their duty cycles. Versions A, B, and
generation PTT's each contained various hardware and software improvements over previous models.
cases, location failures preceded message failBoth of the two generation 2A PTT's failed before the expected battery life of 288 days. Location failures occurred after 197 and 283 days; message failures ocures.
In
most
proximately 10 months.
Eight PTT's have been deployed on walruses since summer 1987. The longest operation time for any of the 8 was 4.5 months; the others have experienced either location or message failures within 4 months. Although reasons for failure are still unknown, it seems that current hardware configurations for walruses are not capable of providing the > 1-year expected life span typical of most
terrestrial species applications.
curred after 244 and 283 days. All five generation 3A PTT's experienced location failures before the end of their
expected 414-day battery life, although one location failure occurred on day 411. Three of the five exceeded the expected battery
life
for
messages only.
Survival rates were similar for the 30
2B and 30 3B
models
(Fig. 4); however, 3B PTT's appeared to perform slightly better than the 2B PTTs. More 3B PTT's than 2B
PTT's provided both location and sensor data throughout
120 100
O)
Fig. 4. Survival
Expected Battery
Life
curves of Model
2B and 3B
CO
i
platform transmitter terminals (PTT's) on polar bears in the Beaufort, Chukchi, and
0)
Bering seas, 1986-88.
O
CO
Q.
Model 2B Location
Model 2B Data only Model 3B Location
Model 36 Data only
100
200
300
400
500
Days since Deployment
most 10-day
intervals.
The 2B versions
failed at higher
resulted in biased data (S. Tomkiewicz, Telonics, Inc.,
rates early in their
deployments, although 3B
versions
failed at higher rates as
deployments neared the expected

recently deployed generation
rate
personal communication). PTT's deployed on caribou at other times were not tested for mercury switch malfunc-
battery life (Fig. 4).
The
3C
of 8.3% (3 PTT's) during the first 140 days following deployment, suggesting improvement over the 3B version.
PTT's had a message failure
based on the very low activity counts obtained during visual observation (D. Vales, personal communication), one of two PTT's on elk seemed to have
tions, although,
Duty Cycles
988) reported a few shifts or errors in duty of second-generation PTT's. Among the 56 PTT's cycles placed on caribou in 1987-88, we noted no similar shifts
had a similar problem. The activity sensor on one of two PTT's deployed on muskoxen in Greenland also malfunctioned after two months of operation.
Fancy et al.
( 1
Efficiency
or errors. All PTT's

cles
programmed
for diverse duty cyal
on 4 muskoxen, 2 elk, 2 mule deer, 2 moose, 1 Dall sheep, 1 1 brown bears, and 1 gray wolf operated without
The quantity and quality of data received from individuPTT's varied among projects. We examined hypotheses
was influenced by
lati-
that efficiency of data collection
detectable errors or shifts.
To lengthen expected transmitter survival time from 414 to 648 days (Table 5), we altered duty cycles of PTT's deployed on polar bears in 1988. We also documented the reduction in location frequency resulting from the reduction in transmission hours per 72-h cycling period.
tude, season, presence of topographic relief in the study

area, longevity of the
deployment, and species.
We
de-
fined two monthly performance indices that provided standardized measures of efficiency across projects, species,
Mean
(standard error; SE) locations per PTT within 72-h periods were 4.38 (0.4 1 ) for 1 2/60 duty cycles, 2.74 (0. 1 1 ) for 8/64
PTT was
duty cycles, and so forth. The message index for each defined as the number of times at least one
duty cycles, and 2.27 (0.18) for 7/65 duty cycles. We considered a duty cycle successful for polar bears
it
if
yielded an average of at least one location per 72-h period. Compared with 88% (SE 3%) for 8/64 PTT's and
that PTT each month, divided by the total number of transmission hours during that month; the location index for each PTT was the total number of unique location estimates each month divided
message was received from
82% (SE 3%) for 7/65 PTT's, PTT's with the 12/60 duty cycle were successful during 91% (SE 7%) of cycling periods. Reducing transmission hours per cycling period
produced 3 and 10% reductions in the proportions of success. The degree to which altering duty cycles succeeded in extending PTT longevity is not yet
by the total number of transmission hours during that month. The latter index was a rough estimate of probability of obtaining a location during each hour of transmission time. Both indices adjusted for differences in duty
by 33 and
42%
cycles
known.
Activity Sensor Malfunctions
In late 1987, because of unusually
efficiencies for 9 species in 12 study areas are summarized in Table 6. Walrus data are not included because transmitters could only operate when the
among PTT's. Mean data collection
animals were surfaced, thus there were no set expected
hours of transmission.
low
activity indices,
Mean message indices varied from low of 0.37 for elk in Yellowstone National Park to a high
to suspect that some activity sensors in PTT's on caribou had malfunctioned. On inspection deployed following sensor removal (for refurbishing), 4 of 12 mercury switches inspected were found to have cracks that
we began
of 1.16 for muskoxen in Greenland (indices > 1.0 were possible where satellite overpasses occurred more than
once per hour). Monthly location performance indices varied from a low of 0.08 for Kodiak brown bears to a high
Table
5.
and Bering seas during 1985-88. Sample

Sp = spring, Fa =fall.
Platform transmitter terminals (PTT's) deployed on polar bears fUrsus maritimus) in the Beaufort, Chukchi, sizes ofSC PTT's with different duty cycles appear in parentheses. Dates:
PTT
generation
10
FISH
Table
6.
Performance indices (with their coefficients of variation [CV]
in
transmitter terminals (PTT's) on various species, April 1987'-September 1988. months used in calculations.
parentheses) and study locations for platform Sample sizes are the number ofPTT-
Species and general location
11
X
0)
_c
g E
CO
Fig. 5. Location performance indices for four PTT's deployed on caribou dur-
ing October 1987.
.0 "CO
o o
J
S
by
others, suggested that
Sea. However, these patterns were not consistent, even among polar bears. Figure 7B shows the pattern of performance from four PTT's deployed on polar bears in the
Our
data, as well as studies
topography affected efficiency. Specifically, efficiency was lower when animals were in valley bottoms, espehigh topographic relief. In one experiment, a PTT was placed in a north-facing talus-slope gully within the Brooks Range Dall sheep study area (68 north
cially in areas of
latitude).
Beaufort Sea in 1987.
The surrounding rock walls were about 200
1.61
Message
Index
high, and the gully was approximately 50 m across. During > 17 h of transmission time, no messages were re-
ceived from the PTT.

tions for a bear
On Kodiak
Island, only
two
loca-
1.2
were calculated during the entire month of
0.8 H
August 1987. This bear was primarily using the stream bottom of a deep canyon from which radio signals might
have had difficulty reaching the
satellite (V.
Barnes,
o
T3
<D
0.4
n U
1.6
personal communication). Keating (unpublished report, on file at Glacier National Park, West Glacier, Montana) tested a Telonics PTT at 23 mountainous locations in Glacier National Park. Locations were
classified as being valley, midslope, or
AFWRC,
mountain peak.
Keating defined the index
R5
as the proportion of over-
1-21
Location Index
passes yielding locations, and reported that R 5 was significantly higher (P < 0.00 1 ) for mountain peak locations than
for midslope locations,
and significantly lower (P < 0.00 1 )

and Craighead (1987) also
0.8
for valley locations. Craighead
0.4
reported lower efficiency from PTT's on caribou in mountain valleys than in open terrain. Because efficiency is
related to terrain, investigators
interpret the
may be
misled
if
they
number of
location estimates as representa-
30
40
50
60
Mean
Fig. 6. Relation
Latitude of
latitude
80 Study Area ()
70
90
tive of the time spent by the

ic
animal in different topographin Table
types.
Efficiencies
text
summarized
6 are from PTT's
between
and PTT performance. See
deployed
in the field. Efficiencies
achieved by PTT's dur-
for explanation of indices.
ing experiments before deployment were
much
higher
12
FISH
0.5-1
Chukchi Sea
except brown bears). PTT's were subjected to considerable abuse, inherent with an animal that lives in such a cold climate, moves in and out of icy water,
slightly, as in other applications
In addition, polar bear
and
it
is
animals that weigh hundreds of pounds. Still, possible that the large body mass of both species of
kills
bears contributed to the attenuation of the transmitted

signal
(VSWR effect).
Projects that experienced relatively high efficiency in
obtaining locations also received a higher proportion of

better quality locations than did those with
efficiency. This result
low overall
was not
is
surprising, because the
quality of the location estimate
positively related to the
number of messages. The northern Alaska muskoxen study was the only one in which LQ1 (poorest quality)
locations were outnumbered by LQ2 locations (Table 7). Those projects where efficiency was low such as the mule deer study in Idaho and the brown bear study on
Kodiak Island
also had the highest proportions of the lowest quality (LQ1) locations.
Precision and Accuracy of Locations
9 10
11
12
30-day Period
Fig 7. Location performance indices for PTT's deployed on polar bears: A. Three bears in the Chukchi Sea, showing decreasing performance with time; B. Four bears in the Beaufort Sea,
Some study objectives depend on the system's ability to maintain an acceptably small magnitude of location error. Factors that may have contributed error to Argos's esti-
PTT oscillator instability, changes in PTT animal movement, insufficient number of transelevation,
mate included
missions reaching the
al data,
satellite,
and errors
in satellite orbit-
showing variable performance with time.
computational algorithms, or
mapping methods.
We
addressed the following hypotheses concerning loca-
tion precision:
Fluctuating
PTT temperature (assumed to affect oscillareduced precision from that achieved using
the satellite as
it
than those shown. For example, during tests designed for assessing accuracy and precision of PTT's (see next section), the location
tor stability)
PTT
at constant temperature.
performance index of PTT's was 0.94 when placed on fenceposts and buildings but dropped to
The elevation of
approach
to the
made
its
nearest
in the
PTT was related to the precision
when placed on nearby captive caribou. Similarly, Keating (unpublished report) reported a 43% reduction in R s for a PTT deployed on a female bighorn sheep (Ovis
0.65
resulting locations.
Animal PTT's (with
canadensis) compared to similar fixed locations. The poorer performance of PTT's when placed on
short antennas mostly-encased in did not achieve the level of precision the collar) achieved by larger, fixed PTT's with long, external an-
tennas (the type used in Argos's
own
estimates of the
animals
is likely
caused by the proximity of the antenna
body and the resulting effect on the voltage standing wave ratio (VSWR). The VSWR effect results in reduction of effective radiated power from the
to the animal's
system's precision). Third-generation PTT's produced locations of greater precision than second-generation models.
Individual PTT's displayed detectable variation in

precision.
antenna.
Polar bear PTT's were the least efficient of

cies
all
spe-
we
tested. The antennas of

(as
all
polar bear PTT's

to protruding
Deployment of PTT's on animals reduced locational precision from that achieved at fixed stations (e.g., roofs
of buildings, trees, posts).
were encased within the collar
opposed
13
Table
7. Proportion of locations in each of the 3 location quality index (LQ) categories for platform transmitter terminals (PTT's) on various species, April 1987-September 1988. LQ3 locations are the best quality, followed by LQ2 andLQl Sample sizes are the total number of locations receivedfrom all PTT's during the period. LQO locations
.
are excluded.
Species and general location
14
FISH
Table
8.
terminals (PTT's)
Precision of locations (in meters) obtained from transmitters at known locations. Fixed platform transmitter had long antennas and were not miniaturized. Only locations coded with quality indices 2 and 3
quality indices were not used by
were obtained from fixed transmitters. Telonics PTT's were the type used on caribou (Rangifer tarandusj. Location Argos before April 1987.
Location
15
c o
600 550 500
(56)
g o
maximum
pass.
sizes.
Fig. 8. Relation between location error and satellite elevation during an over-
CD 13
Numbers
in parentheses are
sample
450
(104)
o
it-
400
(117)
CD
O C
CO
(150)
350
10
CO
b
(Table 9). Location estimates calculated for two PTT's
30 Maximum
20
40
50
60
70
()
80
90
Satellite Elevation
In a crossover experiment, however,
we found
indica-
deployed on captive caribou consistently had greater error

than did location estimates calculated simultaneously (same overpass) for two similar PTT's on an adjacent
tions that the reduction in precision associated with de-
ployment on an animal may be complicated by a 3-way interaction between overpass, attachment status (on vertion
one each on a fencepost (Fig. 11). (The remaining PTT's contributed too few locations to caribou and a fencepost
Movement of the caribou within the pen could have contributed, at most, about 100 of this error. Locations from the same overpass were estimated from signifibe of use.)
sus off the animal), and the specific PTT. third-generaPTT that had been tested on a caribou later produced location estimates with standard deviations 33 and 58%
cal
lower when transmitting from the fencepost during identidaytime hours (Wilcoxon LMest of ranked straight-line
cantly fewer messages for the caribou PTT's than for the PTT's on the fencepost (paired-f = 2.58, df = 1 8, P < 0.02). Apparently, signal attenuation (VSWR effect) and subtle
errors, z
= 3.237, P< 0.001; Figs. 12Aand 12B). However, when a second-generation PTT was also tested on both the
caribou and the fencepost, errors were not different (Wilc-
changes
receive
temperature or orientation of the PTT attached to the caribou contributed to the failure of the satellite to
in
some messages, which resulted in decreased preci-
sion of location estimates.
oxon t/-test, z = 1.383, P > 0.15; Figs. 12C and 12D). These two PTT's did not differ from each other in variability of location estimates when both were on the = 1 133, P > 0.25), but they fencepost (Wilcoxon U- test, z
.
500 1
Fig. 9. Improvement in mean location error for a FIT at Nome (solid line) as overpasses
(grouped by
maximum
elevation in 10
LLJ
450
blocks) are progressively excluded from the total sample. Dashed line shows the proportion of overpasses remaining as
each block
is
C 400
03 0)
excluded.
Numbers above dashed line are the

elevations of excluded blocks.
maximum
350 300
Locations Excluded (10
blocks)
16
FISH
PIT A
PTT B
PTT C
Fig. 10. Location error for 3 third-generation PTT's at the same location on a rooftop dur-
k_
O
u_
LLI
ing 10 overpasses. Each group of error bars represents a satellite overpass.
Overpasses
did differ
when both were on

it
captive caribou (z
3.024,
P<
appears that certain PTT's are more susceptible to increased error from deployment on animals; also, animal orientation may influence the number
0.003). Thus,
of messages received during an overpass. Transmissions from a PTT on a free-ranging Dall sheep appeared to vary
in strength
depending on the orientation of the animal (M. Hansen, personal communication, June 1988). Accuracy. Mean location estimates from fixed PTT's
diverged slightly from the true location, primarily along

the longitudinal axis.
Fancy
et al.
(1988) reported that
Table
9.
Performance of Te Ionics platform transmitter
terminals (PTT's), three deployed on caribou (Rangifer tarandus) in a small enclosure and three on afencepost
adjacent to the enclosure.

Standard deviation of locations
17
Keating (unpublished report) were similarly unable to find
Fencepost
A.
significant correlations
between
PTT
elevation and error
magnitude within a single data set. However, comparison among data sets confirmed the findings of French ( 1 986) that locations were biased when calculated using an incor13). The practical effect of Argos locations using sea level when PTT's were at estimating higher elevations was an increase in the spread of location
rect elevation (Fig.
c o o
estimates in the longitudinal directions. Two independent studies of location error at high elevations supported this
_D
conclusion. Squires and Anderson (unpublished data) calculated a mean error of 2,722 from fixed locations in
^
1 o
.2
CO
8
7
Caribou
Yellowstone and Glacier national parks, a much higher error for PTT's at lower elevations than reported by Fancy
B.
et al.
(1988) or this report. Although the national park
6 5
transmitter elevations were not reported, they were considerably above sea level. Keating (unpublished data) cal-
culated a median error of 2,325

(n
m from 23
test locations
= 691)
at
elevations ranging from 1,463 to 3,052
above sea
level.
H
Satellite
Fig. 11. Location errors for a
JJI
Overpass
Elevation-related errors were primarily longitudinal be-
cause the
satellites travel in nearly
north-south orbits.
When
signals
come from PTT's

Argos
that are higher than the
assumed
lite
elevation,
interprets
them as coming from
locations that are closer than they actually are to the satel-
along
its
across-track direction (Fig. 14). Thus,
when
FIT placed on a fencepost (A), and
PTT's were above sea

direct line
level, errors tended to be along a
enclosure.
another simultaneously placed on a captive caribou (B) in an Each bar from (A) corresponds with the one below
from the PTT to the satellite as it passed over. found highly significant (P < 0.01) circular correlations between the azimuth of error and that of the satellite
We
at its nearest
approach to the
PTT
in
both the Brooks
location; nor was any correlation found between the longitudinal or latitudinal components of error and slope (Table
12). Similarly,
Range
tional
(radio-collared Dall sheep) and Yellowstone
Na-
no correlations were found between the
azimuth of location error and the aspect of the elk's location (rank circular correlation, Batschelet 198 1 1 87), both
:
Keating (unpublished data) and Squires and Anderson (unpublished data) also found significant relations between azimuths. All these studies
(elk; Fig. 15).
Park
were
at relatively
high elevations.
We
found
that the coritself
when
positions were considered and when only locations with > slope were considered. Keating (personal
all
relation of error
azimuth with
satellite
azimuth was
communication, 1988) also found no correlation between error and aspect of known locations in Glacier National Park, Montana.
correlated with the elevation of the PTT, strengthening with increasing elevation (Fig. 16). Keating (personal
communication) observed a similar strengthening of the azimuth-azimuth relation as PTT elevation increased.
We did not test the effects of vegetative cover on accuracy or precision. However, Squires and Anderson (Wyoming Department of Fish and Game, personal communi-
The magnitude of error arising from a discrepancy between actual and assumed PTT elevation was related to the
maximum elevation of the satellite overpass. As suggested

by the sea-level data (Fig. 8), errors were greatest at very low and very high satellite elevations, and least at intermediate
satellite elevations.
PTT's in Yellowstone and Glacier national parks, found no association between location error and vegetative covers classed as open meadow,
cation), in a test of fixed
These errors intensified con-
and high-density conifers. medium-density we were unable to demonstrate a correlation Finally, between the elevations of the 44 known elk locations and
conifers,
their magnitude
data were from an elevation considerably siderably above sea level. Keating fit a second-order polynomial regression to data collected from a
different elevations; he
when
PTT
of error (Table
2).
Fancy
et al.
( 1
988) and
found
that
almost
he had placed at 53% of the vari-
18
FISH
19
Table 10. Bias in locations from three fixed platform transmitter terminals (PTT's) before and after correction from NAD27 to WGS84 mapping systems. Sample sizes are shown in Table 8.
20
FISH
3
E
3,500i
3,000-
2,000^
Fig. 13. Relation between mean longitudinal error and FIT elevation for PTT's at four
locations.
~
O)
1,500 1,000
Number of location estimates (and

at
number of PTT's)
(5);
each
site
o
0)
of increasing elevation: 113
(4);
were, in order 48 (3); 101
140
(5).
500^
200
400
600
800
1,000
Location Elevation (m)
Fig. 14. Schematic representation of location error resulting from a PTT being at a higher
elevation than that assumed by Argos when making calculations. The PTT is estimated to
be
at the intersection
of the cone with the
earth surface (vector B), when in fact, it is farther from the satellite in the longitudinal direction but above the assumed earth sur-
and face in elevation (vector A). Vectors are of equal length. The resulting error
B
is
generally along a direct line from the true location to the satellite at its closest
approach.
360
300
D E
21
0.8
Fig. 16. Relation
study
sites
between the elevation of nine and the strength of the correlation
C o
(Batschelet 1981:187) between satellite azimuth at closest approach and the azimuth of error. Locations, in order of increasing
elevation, are Inuvik, Northwest Territorities; Nome, Alaska; Fairbanks, Alaska
0.6
O O
o
0.4
1
Chatham Dome, AK; Murphy Dome, Alaska; Galbraith Lake, Alaska; Gardiner-Mammoth area of Yellowstone National Park. Argos's calculations assumed that all PTT's were at sea level.
(three locations);
0.2
500
1POO
100
2,000
Location Elevation (m)
Long-term Activity Index
angles were so extreme that motion was never detected because the mercury never moved back and forth, while
other angles produced high counts from subtle ments, such as those from respiration.
We
did not calibrate 24-h sensors to specific behavior

it
move-
keep animals under constant surveillance. However, from extensive experience in the caribou project, we found that successive
patterns because
was
difficult to
Methods. All experiments were conducted with a speto adcially designed collar that allowed the investigator
just the inclination of the
counts of zero meant that the animal had died or the collar
mercury tip-switch for recording
had been shed. In some cases,
this indicator
quickly re-
sulted in aerial searches to find the animal,
which allowed
activity data (Pank et al. 1985, 1987; Fancy et al. 1988). The collar was otherwise identical to other second-genera-
a better chance to determine the cause of death and retrieve

the collar.
tion collars
we
used.
at
Data were obtained
60-s intervals by using a Telonics
Results from caribou studies supported the concept that the 24-h index was related to the amount of activity. Studies
uplink receiver (Beaty et al. 1987) that received transmissions from active PTT's within a 2-km radius. Captive
of the Porcupine and Central Arctic caribou herds
animals were
fitted
with the experimental collar and ob-
revealed a strong (P
rates
< 0.0001)
correlation
between the
mean monthly 24-h activity index and monthly movement

(Fancy
et al. 1989).
had
their lowest values in midwinter.
Both indices peaked in July and It also seemed that
_
O
LU
6i
5-
the 24-h index
was
useful in identifying
for a caribou
when
the caribou
cows bore young:
known
to be pregnant, a
notable drop in the index indicated calving (Fig. 19).
Short-term Activity Index

Calibration of the Short-term Activity Index
Calibration studies were conducted with captive caribou, elk,
15
mule
deer,
and moose. The purpose was
to
associate counts or series of counts with gross activity categories (e.g., inactive, walking, feeding) and apply the
interpretation to free-ranging animals. Calibration in the
10
20
30 Maximum
40
50
60
()
70
Satellite Elevation
wild was performed only for elk. For each species, preliminary experiments were conducted to determine the
Fig. 17. Relation between longitudinal errors of location estimates and maximum elevation of the satellite for 20 known
mercury switch that resulted in the best discrimination between activity types. Some switch
inclination of the
positions of Dall sheep in Alaska. Argos's calculations assumed that the PTT was at sea level. Data courtesy of M.
Hansen, University of Alaska.
22
FISH
23
24
FISH
A.
Running
(o
19)
B.
Running
(a = 4)
Walking (0- 206)
Walking
(o
&)
O
CD
Feeding (o- 555)
Feeding
(/?
135)
C7
CD
Lying ("- 385)
Lying (o
53)
20
40
60
20
40
60
D.
C.
Moving
(a
7)
Running
Feeding (o- 125)
CD
Lying (n* 230)
8
LL
20
25
26
FISH
Overpass 1 Score - 1.631
Running
D.
0.2
,-
Overpass 1 Score 2.000
10
20
40
50
60
10
20
30
40
50
Short-term Activity Index

Fig. 22.
Comparison of short-term activity indices (scaled as proportions) for a free-ranging caribou during two satellite overpasses with indices obtained from captive caribou engaged in four different activities. A. Lying. B. Feeding. C. Walking. D. Running. = 0.097), whereas counts for overpass 2 overlapped Activity counts for overpass 1 overlapped most with lying (A, lowest score
most with running (D, score = 0.999).
anomalies in each mercury switch, sampling

other causes.
error, or
sions using a conventional tip-switch motion detector.
However, they also pointed out
that standing behavior is
The 60-s motion detector has potential as a short-term activity index when behavioral categories
Discussion.
are coarsely defined.
generally uncommon in the wild for most ungulates; thus, the confounding of inactive and active periods would
We
are skeptical that attempts to
between categories of activity (e.g., browsing versus grazing) would be productive. Fancy et al. (1988) found high overlap in sensor counts with a finer delineation of activities. Gillingham and Bunnell
discriminate
finely
more
probably be minor. Discrimination accuracy for captive animals was generally
high
overpass.
when using the entire series of counts from an Our simulation experiments for the caribou data
(1985) concluded that conventional tip-switch and variable-pulse collars were not highly accurate in discriminating
analysis
tivity
suggested that classification error was quite small. This assumed that successive counts from each ac-
among
nine behavior categories.
We have noted
lim-
itations in the present system; for
example, in not one of
category were independent; examination of our data supported this assumption. The assumption that animals did not change behavior during the course of an overpass
the four species tested could
from bedded
(inactive).
We
distinguish standing still can see no resolution for this
we
was also implicit in this categorization system. Although we had no method to assess the behavior of the categorization system
problem; in a motion-sensing device, the effect of standing motionless cannot be expected to differ from lying.
Beier and McCullough (1988) reported similar conclu-
when animals switched

it is
activity types during
an overpass,
likely that the selected type will
that actually occurred,
be one most often the predominant one.
27
bedded. Orienting the switch to be less sensitive might well result in low counts during prolonged grazing, when
the neck
is
held down, or perhaps even during trotting,
when the neck is held high. The captive female moose seemed to not be bothered by the size and weight of the collar. Feeding by the moose
Lying
Feeding Running
(browsing on
tion
0.5
Walking
tall shrubs) produced slightly better separafrom inactive behaviors than did feeding by the elk (ground-level grazing of grasses and forbs).
General activity patterns for free-ranging caribou, as

Mean number
of
messages
for
overpasses yielding locations
(7.8)
predicted by the short-term activity index, correlated well with information about the movement patterns of these animals. However, the proportion of time estimated in
feeding activity was considerably lower and the time spent inactive higher than documented by observational
studies of Alaskan caribou.
It
remains to be seen which
2 3 4 5 6 7 8 9 10
11
12 13 14 15 16
M essages / Over pass

Fig. 23. Relation between the probability of misclassifying a caribou's activity based on short-term activity counts and the
factors contribute to a bias in favor of the inactive category
expense of the three active categories. Such biases should be identified, quantified, and corrected by calibraat the
tion under wild conditions.
We conclude that although the
number of messages (i.e., activity counts) received during an overpass. Each data point was estimated from 50,000
computer
iterations.
using PTT's are reliable for behavior types, they should not be used quantitatively for energetic studies at this time but rather as an index for
general patterns
assessing gross seasonal trends and herd-specific differ-
we found by
The experimental collar seemed to be too heavy (1.6 kg) and too large (7 cm wide) for female mule deer. The deer frequently attempted to remove the collar and seemed
even after wearing it for 3 days. A shape more contoured to the slender neck of female mule deer would provide activity data with
uncomfortable with
it,
ences in activity budgets.
Temperature Sensor
All PTT's
we deployed
since 1985 have included tem-
lighter collar with a
perature sensors. We have not tested all PTT's for accuracy, but a spot check of 4 third-generation PTT's that had
less disturbance to the animal.
For elk, considerable overlap between counts from bedding and feeding activity occurred, even at the best setting of the mercury tip-switch (+2). Elk frequently graze with their necks pointed downward; they also trot
of +2 with their heads held high, pointed upward. An orientation succeeded in generating intermediate counts for
been deployed on caribou during 1987 showed that temperatures were reliably reported with little variation
among the four. The overall mean slope relating reported temperature to ambient temperature was 0.901; differences among the four PTT's were nonsignificant
(P > 0.05).
feeding and high counts for walking-running, but also resulted in occasional intermediate and high counts while
However, we are uncertain what the temperature data gathered under wild conditions represent. The temperature sensor reflects ambient temperatures under controlled
Table 14. Correspondence ofactivity predicted by 50, 000 simulation
trials with the known (observed) caribou (Rangifer tarandusj activity that produced the distribution (sample sizes given for each count distribution). In all cases, the length of the count series (overpass length) followed the distribution for northern Alaskan caribou.
Predicted activity
Observed
activity
28
FISH
Inactive
Feeding
Inactive
Feeding Running
Walking
Running
Walking
cu
1i
c
.0
*1_
O
Q.
O N D
1986
Fig. 24.
A M J 1987
June
Fig. 25.
July
Mean proportion of time spent in four activities by freeranging caribou in northern Alaska, October 1986-October 1987, as estimated by the short-term activity index.
Mean proportion of time spent in four activities by six free-ranging caribou in northern Alaska during June and July, 1987 as estimated by the short-term activity index.
conditions, but other data suggest that these relations break down somewhat under field conditions. The temperature sensor cannot respond to rapid changes in ambient temperature because the canister and internal PTT
mal's body and by the electronic circuitry
itself.
Pank et al.
(1985) were able to explain only ambient temperature (measured in a shaded area 50-100
59%
of the variance in
m away) and PTT temperature when collars were attached

to captive caribou.
one experiment, components have an moved from ambient tem4 third-generation PTT's were
insulating effect. In
PTT
temperatures were most often

relation
warmer than ambient temperatures. The
between
peratures of roughly 4
to
24
C. At the end of the
PTT
experiment (50 min), the temperature sensors read between 13.5 and 14.5 C (Fig. 26). These sensors seem to require > 1 h to register such an extreme change in an
animal's microclimate. In addition to the time lag, other
factors that
temperature and ambient vary among species, seasons, and PTT placement on the animal's body (e.g., Johnsen et al. 1985).
temperature will probably
Saltwater Sensor
may cause the PTT temperature to differ from
ambient temperature include possible warming by the ani-
14
Ambient
Air
Temperature
Two prototype PTT's equipped with saltwater sensors were deployed on polar bears in spring 1987, one each in the Chukchi Sea and Beaufort Sea. An internal clock counted the number of seconds of immersion within each
72-h duty cycle. An additional counter recorded the ber of times the PTT was immersed in salt water
during each 72-h cycle.
num> 5 s
Transmitted
PTT Temperatures
more time
27).
much According to the data, the Chukchi Sea bear spent in saltwater than the Beaufort Sea bear (Fig.
We recorded only 9 immersions for the Beaufort bear but 1,522 for the Chukchi bear during the same period. Reasons for the time differences in saltwater immersions between the two bears are unclear. Independent evidence
20 30 10 Minutes since Temperature
40
Change
50
have had more access suggests the Chukchi Sea bear may malto open water than the Beaufort Sea bear; however, sensors cannot functions or inconsistencies between the
be ruled out.
Fig. 26. Delayed response of temperature sensors within four
third-generation PTT's to a sudden change in ambient temperature from 5 to 25 C.
29
Beaufort Sea
mates with
failing to
LQ =
(Table 15).
As
expected, locations
in
300
Chukchi Sea
meet Argos's criteria (and thus appearing only
250
i
(u
200150
had considerably greater error. who wish to increase sample size of locaInvestigators tion estimates may include LCO locations with prothe separate data file)
CO
gressively lower LI values but will progressively increase

the error of the resulting data set. Tradeoffs
may
be made
4->
100
50
between increasing sample size and decreasing precision. Table 16 summarizes the LCO locations, and within those,
the proportion in each of the LI categories
jects (except caribou, for
from our pro-
which we did not use
LCO proc-
M
Fig. 27.
essing) from February through September 1988.
O
in salt
As
stated previously, assessing error in locations estiin test situations
Time spent
bears one in determined by saltwater sensors.
water during each month by two polar the Beaufort Sea, and one in the Chukchi Sea as
mated from PTT's

errors are
can be misleading;
be greater when PTT's are acon animals. For example, 14 LCO location tually deployed estimates of free-ranging elk at known positions in Yel-
more
likely to
lowstone National Park had a mean error of 16.65
km
Argos's Location Class Zero (LCO) Service

The Argos Location Class Zero (LCO)
service can be
(median 12.7 km), more than five times the mean error from normally processed location estimates during the
same
used by those who wish locations to be estimated when normal processing fails. Even if the number of locations
estimated by normal processing
is
period. In another example, normally processed lo-
deemed sufficient, LCO
muskox during January 1988 were with the entire set of locations, which included compared those processed by LCO (Fig. 28). This animal was known,
cation estimates of a
processing can be useful because it allows the user to evaluate the performance of individual PTT's by seeing
the causes for normal processing failure (see Table 3) and indices of the signal strength as received by the satellite.
from midwinter
aerial tracking, to
be restricted
personal
to a small
home range
tion, 1988).
(P.
Reynolds,
ANWR,
communicawas
The apparent home range of
the animal
Alternate location estimates are provided by Argos in
LCO
Table 15.
processing, which can be used in place of the normally processed location in those rare instances when the two
Mean and median errors of locations calculated
have been reversed. (Reversals have occurred for less than 0. 1 % of locations received by our projects during the past
six
using normal processing (LQ 1,2, and 3) and Argos's LCO processing. All data are from third-generation
months.)
platform transmitter terminals (PTT's) at known locations on Kodiak Island, Alaska; PTT's were not de-
We
had few data
to assess the precision of the
new
animal-tracking service using PTT's at known locations. We obtained 85 location estimates processed under LCO
ployed on animals. were obtained.
No
locations classified LI 6 or LI 9
from PTT's used for testing during June 1 988. Of the total, 64 were classified as acceptable by Argos under their LCO
criteria,
LI category
and thus were displayed as
LQO
locations in the
normally received files. Mean and median errors were 5. 14 and 2.89 km, respectively. For comparison, mean and
median errors from 323 normally processed location estimates from the same PTT's during the same period were
1.03 and 0.61
Precision
km, respectively. was highest for LCO location estimates with
the highest LI indices (i.e., those barely failing the criteria for normal processing). For example, location estimates with LI indices 5 and 6 had mean and median errors only
slightly greater than
normally processed location
esti-
30
FISH
Table 16. Proportion of locations for each species obtained using Argos's location class zero (LCO) processing, and the proportion among those in each of the location indicator (LI) categories. (See Table 3 for description of LCO categories.) Percentages in LI 6 and 9 were zero for all projects. Sample sizes are for LCO locations only. Data were
collected from February through September 1988.
Species
and
1
31
Alaska,
facility,
we have
evaluated a
lonics, Inc., since September 1986. The transmission. from the satellites'
VHP
of two IBM-compatible computers: runs a satellite prediction program that shows where the
satellites are at all
LUT built by TeLUT receives data The LUT consists an XT model, which
satellite
which the
LUT received data from these
PTT's was
79%
(min.-max., 69-85%) of that for Argos. The mean number of locations estimated by the LUT was 50% (min.-max.,
24-72%) of that calculated by Argos. The LUT recorded a mean of 56% (min.-max., 41-66%) of the messages received by Argos for these 18 PTT's. Two important factors that contributed to the lower
quantity of data and locations provided by the LUT were signal interference and the lack of adequate Doppler data
times and characteristics of
overpasses; and an AT model, which operates the tracking antenna, receives and processes the VHP signal containing Argos messages, estimates PTT locations, and produces a report following the overpass.
LUT points a 4-m, two-beam yagi antenna toward is above the horizon. The VHP signal the satellite while
The
it
from reference platforms for some overpasses. Our LUT was located near a major communications facility and a
television station; signal reception
was blocked whenever
containing the
the strength
Argos data
is
received, decoded, and stored
the antenna pointed toward these sources of radio inter-
During each overpass, the LUT displays and quality of each incoming signal, the position of the antenna, and the identification of PTT's for
for later analysis.
ference. Primarily because of radio interference and signal
blockage by hills and buildings, the

locations for overpasses
LUT calculated only 6

satellite ele-
where the maximum
which messages are being received. Doppler data from are used to reference PTT's placed at known locations
determine the position of the satellite in its orbit more precisely. After the overpass, the LUT uses satellite and
was < 30, compared to 236 locations calculated by Argos during the same experiment. We had only a single
vation
reference
PTT
located in Alaska, and lack of sufficient
data from this

locations for
Doppler
data, received
from the reference PTT's during
the overpass, to estimate
PTT
locations.
prevented the LUT from calculating 729 of 2,265 overpasses (32%) during the December 1987 experiment. The lack of reference data
PTT
A LUT cannot provide as many

PTT
location estimates and
may not be a problem for LUT's
sensor messages as standard Argos processing can. For data to be received by a LUT, the satellite must view both
the
located in the contiguous United States or other locations where several reference
satellite. Tel-
platforms can be placed within view of the

onics, Inc.,
is
and
LUT
simultaneously. Argos uses tape re-
now
testing a
new LUT system with
more
corders on the satellite to store messages for playback to ground receiving stations; therefore, to receive data, only
the
expensive and sophisticated tracking antenna to reduce or

eliminate interference from other radio sources.
Precision
PTT
needs to be within view of the
satellite.
Further-
more, radio interference near the horizon from VHP sources, especially in metropolitan areas, may reduce the
The precision of
locations calculated
by the
LUT
was
number and
quality of messages received
ticularly those with omnidirectional antennas.

tionally, the software
by a LUT, parAddiis
determined in March 1988 using nine PTT's placed at known locations near Fairbanks. PTT's were placed
at
used to estimate locations
elevations of 152, 708, and 902
(three at each
propri-
among LUT manufacturers and between LUT's and Argos. Therefore, even when the received data
etary and differs
are identical, estimated locations
may
not be.
was used in the site); however, an elevation of location calculations to enable comparisons of location accuracy with standard Argos processing. Locations estimated by the LUT (n = 93) had a mean error of 12.3 km. (For comparison, the mean error for 354 locations estimated by Argos was 1 .4 km.) Fifty-six per-
Performance
Reliability
cent of the
location.
LUT
locations were within 5
km
of the true
When
overpasses with a
maximum
satellite ele-
During December 1987, we compared the number of messages received by our LUT from 18 PTT's deployed
in northern Alaska and Yukon Territory with number received using standard Argos processing. The caribou ranged 394-633 km from Fairbanks during the experiment (mean = 546 km). For each PTT, we calculated ( 1 ) the number of overpasses during which at least one message was received by the LUT or Argos, (2) the number of locations estimated, and (3) the total number of messages received. The mean number of overpasses for
vation exceeding 70 were excluded, the mean error of locations estimated by the LUT fell to 5.7 km, and the
on caribou
the
percentage of locations within 5

tion rose to
km of the true PTT loca-
68%. Argos
rarely calculates locations for
overpasses greater than 70
because of poor location
accuracy. Precision of locations dropped off markedly when PTT's were far from the nearest reference platform. The
transmitter at
Nome, Alaska, had been used as
a reference
platform for our
LUT and helped to calibrate locations in
32
FISH
northwestern Alaska. However,

transmitter
when
the position of this
mit 6 h/day
or, in the
1
was
deliberately treated as an
unknown,
the
h/day between
May
case of 13 third-generation PTT's, 12 and 30 September and 6 h every
was more than 800 km away. Under these circumstances, the median location error from a randomly chosen sample of 200 location estimates fell to 17 km, and the mean fell to 47.4 km. More than 18% of these locations were > 100 km from the true location, and only 2.5% were within 2 km.
nearest transmitter that could be used for reference
near Fairbanks
other day during the rest of the year. These duty cycles gave a theoretical battery life of 1 year. Five of 42 PTT's
deployed on caribou before October 1986 experienced message failure within 6 months of deployment. A mean
of 3.5 locations per day was obtained for caribou with second-generation PTT's. Caribou with third-generation
PTT's were located 8.0 times daily between

September.
Daily
the
May
and
Cost Comparison
The primary advantages of a LUT compared to standard Argos processing are ( 1 ) avoidance of the usual 3-5-h
delay for data processing; (2) greater processing flex-
movement
rates of radio-collared caribou
from
two herds
which
differ greatly in size (165,000
versus 16,000 caribou) and separation of seasonal were similar except during spring and fall migraranges
tions (Fancy et
al.
1989). In both herds,
movement rates in
reduced cost for some applications. The cost of a LUT is approximately $20,000-40,000. A system using S-band transmissions and a more sophisticated
ibility;
and
(3)
July exceeded those during migration. The annual distances traveled by caribou cows ranged to 5,055 km; these
were the longest movements recorded for any

animal.
terrestrial
tracking antenna costs about $65,000.
LUT users must still
pay Argos for use of the system, although the minimum rate is only 25% of the standard processing cost. Users
request archived data for a particular month will be charged for a full month of standard processing plus a
service fee.
In situations
for our caribou studies.

tively
We considered satellite telemetry a useful research tool We were satisfied with the rela-
who
low failure rate and high efficiency of data-gathering at these latitudes, and we considered that location precision was adequate for our objectives.
where reduced data quantity and location
accuracy are acceptable, a

studies that involve as
LUT can
be cost effective for
few as
five PTT's.
For example,
Polar Bear: Beaufort Sea

Polar bears occupy one of the most remote habitats in
the polar ice cap. The pack ice substrate is in constant motion, and there are no permanent fuel caches or logistics bases on its surface. Further, only the Cetacea and the world
assuming the processing charge assessed during was equivalent to the 1987, the purchase price of our annual processing cost for 10 PTT's transmitting daily.
AFWRC
LUT
Field Studies
Caribou: Northern Alaska and Yukon
Since 1985, the
some members of the Pinnipedia range more widely than polar bears. Satellite telemetry was used to gather data on polar bear movements and activities that would not otherwise be obtainable. Forty-four PTT's were deployed on polar bears in the Beaufort Sea beginning in spring 1985. Through June 1988, 10,547 locations and 128,038 activity
AFWRC and ADFG have used satellite

The informa-
telemetry to monitor the daily movements and activity of caribou of the Porcupine and Central Arctic herds in
and temperature data were recorded.

Satellite telemetry
northern Alaska and northwestern Canada.

tion
is
provided information on maternity
used to assess potential effects of oil and gas development within the Arctic National Wildlife Refuge
(
AN WR) on caribou and to mitigate the effects of chang-
den entry and emergence dates of polar bears. Polar bears in dens maintain consistently warmer temperatures than those not in dens, sleep most of the time, and move very
little.
ing land use and resource
management practices. Between 1985 and 1987, more than 49,000 locations and 79,000 sets of sensor data (temperature and activity) were obtained for 34 adult female caribou using satellite telemetry. Caribou were captured on winter range when immobilizing drugs contained in a dart gun were fired from a helicopter. The 1 .5-1 .6-kg collar package included a conventional VHP transmitter that was used to relocate the caribou by aircraft. PTT's were programmed to trans-
We
PTT's
to provide clues to entrance
used activity and temperature sensors within and emergence times.
We documented dates of den entrances for 22 polar bears

using these data; however, because many PTT's failed to reach the end of their expected lifetime, we documented emergence times for only seven of these bears. Also,
PTT's often did not provide locations of denning those positions that were fixed during denning tended to be inaccurate. One den that was visited in 1988 was consis-
33
Alaska
Canada
\
Fig. 29. Movements of female polar bears in the Chukchi and Bering seas (A) and the
60O
Beaufort Sea (B),
May 1985-May
1988.
B.
tently positioned
by the
satellite 11.1
km
from
its
actual
location.
moved
of our existing knowledge of the seasonal and
this,
Similarly, during winter 1986, many collared bears to locations southwest of Point Barrow, Alaska
Much
areas where
we had
not seen
them
before. Contradicting
annual movements and distribution of Beaufort Sea bears

has been obtained using conventional telemetry in recent years. Conventional telemetry has shown that although polar bears are seasonal to general regions or activity
areas, these areas are extremely large,
however, were activity and temperature data transmitting from some of these "moving" bears, suggesting
that they
were
in maternity dens. Therefore,
we hypothe-
sometimes exceed-
sized that unusual currents that year in the southern Beaufort Sea had passively carried bears that had denned on the
ice.
ing 259,000
km 2
Satellite telemetry has
expanded our
This hypothesis was subsequently corroborated by
knowledge of the size of these activity areas (Fig. 29). Satellite telemetry has also, in some cases, provided details needed to determine the purpose of some of the
longest movements. For example, in previous years, some bears wearing VHP transmitters were radio-tracked in
aerial telemetry.
Because of the high costs of using aircraft with convenwe are limited to 5-6 survey flights each year. With satellite telemetry, we can obtain much more
tional telemetry,
northwesterly directions until they were beyond the range of survey aircraft or until they entered the waters of the
Soviet Union.
movement data, although it is on a smaller sample of bears. Future applications that may make use of such detailed data include studying the relations between
detailed
We suspected that those bears were moving

of the polar basin to den, because food
is
movements of
rently,
the sea ice
and those of polar bears. Cur-
to the stable ice
scare far offshore and foraging is therefore difficult. Activity and temperature sensor data received from satellite
however, the unreliability and relatively short life of PTT's limits our ability to conduct such a study. span Studies requiring frequent visual relocations of marked
individuals (e.g., predator-prey relations) may potentially be made more feasible by satellite telemetry because investigators can fly directly to the animals, rather than
confirmed our hypothesis that many of those bears traveling far offshore were seeking and entering maternity dens.
collars has
34
FISH
having to search large areas for them. However, greater reliability of PTT's deployed on polar bears is necessary
for these studies to be feasible.
and refurbished and placed on another cow muskox in July 1987. The second collar transmitted for 6 months
until the
another
Polar Bear: Chukchi Sea and Bering Sea

Satellite telemetry is
muskox was killed; it was then placed on cow in July 1987 without being refurbished.
being used in the Chukchi Sea and
the Bering
Sea
to define the seasonal
movement
patterns
Both collars have been functioning for almost one They were programmed to provide intensive sampling periods at 12- week intervals during which they
year.
used by polar bears. The bears occur seasonally in Alaskan waters but also spend time in waters under jurisdiction of the Soviet Union, where aerial surveys required by conventional telemetry are not permitted.
total area
and
transmitted 16 h/day for 5 days. During the remainder of the year, collars transmitted 6 h/day every third day. third satellite collar, with a duty cycle of 6 h/day every
other day,
was deployed on
cow muskox from October
Using satellite telemetry data, we have found that as the sea ice retreats from the Chukchi Sea, polar bears also
retreat into Soviet waters
1987-April 1988.
Preliminary analysis of movement data from one animal during 1986-87 and three animals during 1987-88 indicated that muskoxen have small home ranges and
and often spend summer
in the
in
vicinity of Wrangel Island. When the sea ice advances fall, polar bears again move into U.S. waters.
move
only short distances during the darkest, coldest

(Fig. 30).
months of winter
To
ters.
satisfy study objectives,
lar individual
female bears
we have attempted to recolwhen they return to U.S. waIn July 1987,
This effort has met with only partial success, because some PTT's have failed prematurely and because some
Muskox: Greenland
adult
bears have denned while in Soviet waters or territory and did not return to U.S. waters during the PTT's battery life.
In an attempt to extend battery life through a second
two third-generation PTT's were put onto male muskoxen in the Kap Kobenhavn area in Peary
Land, northern Greenland (82.5 N, 22.5 W), as part of a cooperative study with D.R. Klein, ACWRU. The collars
we have experimented with altering However, the success of this experiment is not duty cycles.
spring capture season,
yet determined.
were deployed
to
high latitudes for
provide data on muskoxen activity at comparison with data from muskoxen
The longevity of PTT's seems
to
be improving, as does
the potential of the saltwater switch for further interpreting the polar bear-sea ice relation. Satellite telemetry is currently the only methodology available for addressing several of the
populations at lower latitudes. The study area is a high arctic polar desert with most vegetation limited to sedgegrass and willow communities in scattered locations
where meltwater
the
is
available throughout
summer. As with
ANWR
major objectives of the western polar bear
made
study, logistical problems in winter data collection by other means impractical. Trials
muskox
project.
with captive muskoxen and various orientations of the PTT's mercury tip-switch were not able to accurately dif-
Muskox: Arctic Slope

Muskoxen were reintroduced to the coastal plain of the in 1969 and 1970. Muskoxen have been radiocollared since 1982 to document distribution and movements of the population (Reynolds 1987). The animals
ANWR
the simple mercury However, a tip-switch orientation was chosen tip-switch. that seemed capable of providing a measure of active-
ferentiate
muskoxen behaviors using
versus-inactive time for
for
muskoxen in northern Greenland comparison with data from other muskoxen

to transmit for a 6-h
display high fidelity to specific geographic areas and remain year round on the coastal plain. Data on distribution,
populations.
The two PTT's were programmed

period during each 5
1
h.
movements, and activity patterns of muskoxen in winter are needed to assess potential effects of petroleum develto obtain with conventional radiotelemetry
Then, the beginning of the 6-h
opment. Such information has been particularly difficult because of

tions
period was shifted 3 h later every two days, so that all hours of the day would eventually be sampled. The first transmissions from the two transmitters were received on
7 July 1987. One transmitter provided locations and temperature data at least through July 1988, but the activity sensor malfunctioned in mid-October. The second PIT
darkness, blowing snow, and other adverse weather condi-
on the arctic coastal plain during winter. In 1984, a first-generation satellite collar was deployed on a muskox in to test how the collar functioned
ANWR
(Reynolds 1989). In November 1986, two cow muskoxen were collared with third-generation satellite collars.
provided location and activity data for approximately 2 weeks, after which no transmissions were received for > 7 months. For 3 days beginning 15 March 1988 and sporadically since then
transmissions from this second PTT,
One
collar failed almost immediately but
was repaired
including short-term activity data, were again received.
35
Fig. 30.
Movement patterns of muskoxen on

Refuge
1986-1987. Data cour-
the Arctic National Wildlife
(ANWR), Alaska,
tesy of
P.
Reynolds.
Arctic
National
Wildlife
Refuge
Brown Bear: Western Brooks Range

Brown
bears in the western Brooks
home range estimation techniques are known to be sample

size-dependent (Anderson 1982; Swihart and Slade 1985b). The use of satellite telemetry enabled Reynolds to
obtain
Range have been
ADFG since 1977 (Reynolds and Hechtel 1980). This long-term study has made it possible to observe interactions among bears with known
studied by H. Reynolds of
more than 100
locations for each bear during this
3-month period which is substantially more than would have otherwise been possible.
family histories. Despite the wealth of information on this bear population, frequent relocations during a single sea-
Brown Bear: Kodiak Island

Kodiak Island's brown bear population has been the focus of numerous studies. Investigations on the southern part of the island have (1) revealed factors that influence
habitat use
aerial
son have never been obtained, mostly due to logistic and budgetary constraints. In July 1987, objectives of equipping three adult females with radio collars included deter-
mining the minimum number of relocations needed to adequately describe home ranges and assessing the degree of spatial and temporal overlap among females, two of
which were a mother and her adult daughter. All three were fitted with PTT's transmitting for 3 h twice daily. The
by brown bears, (2) assessed the efficiency of and ground inventories along salmon spawningstreams, and (3) determined reasons for the use of particular
streams (Barnes 1985).
PTT's were programmed bears were in dens.

the three bears during
sible.
to
suspend operation
when
the
in
Two
radio collars were deployed during summer 1987 an attempt to refine previous information on the timing
to transmit 8 h/day during
The increased number of seasonal locations for each of 1 987 made some new analyses pos-
of movements between salmon streams. Both PTT's were
programmed
summer and
fall,
Overlap among home ranges of the three bears are shown using modified minimum area polygons (Harvey
once every 4 days during denning, then to resume transmitting 8 h/day in spring. Previous studies (Barnes 1985) had shown that individual bears often moved from stream
to stream to feed
and Barbour 1965; Fig. 31). Only one location from any group occurring within the 3-h duty cycle was used for
home range
the three
estimation.
The magnitude of overlap among
on different runs of spawning salmon. However, the timing of these movements was unknown
tions of the
must be interpreted while considering the limitahome range estimation method. Overlap
because inclement weather frequently made it unsafe to locate bears with aircraft. Collars were not intended to
assist in habitat-use studies
among home ranges estimated here is likely overestimated because location error is not considered; however, most
because of concern about the
precision of locations on Kodiak Island, where the terrain
36
FISH
possibly even more) as the 2 1 previous locations and the
subsequent 9 added together. Most positions during frequent relocations were in broad valleys or in open areas near Karluk Lake and its outlet. The two positions south of
the lake suggested this bear
in precipitous terrain
may have been spending time with a resulting loss in number of locations. This would imply that the number of relocations
of bears in different habitat types on Kodiak Island is not an accurate reflection of the relative amount of time spent
in each.
Dall Sheep: Brooks Range

Objectives of a Dall sheep study in Alaska's Brooks
Range conducted by M. Hansen of the University of Alaska included determining the accuracy of satellite locations for animals inhabiting mountainous terrain and deter-
mining seasonal movements and home range of an adult male Dall sheep. A PTT was placed on a adult ram in
October 1986, and care was taken
tightly to prevent
its
to secure the collar very
chafing
several
hampering the animal's movements or neck. Detailed observations were made for
after attachment to
weeks
determine whether the
20
Kilometers
collar adversely affected the animal's behavior or health.
Data were received from the
PTT
until
it
was removed
from the animal

Fig. 31.
in
October 1987. Movements and home
Home ranges of three brown bears in the western Brooks
Range, Alaska, as estimated by a modified minimum area method (Harvey and Harbour 1965). Data courtesy of H. Reynolds, Alaska Department of Fish and Game.
range were analyzed by selecting only one location each day of transmission. Additional information on activity
and migration was provided by the 24-h
activity index.
The instrumented ram did not seem

affected by carrying the
to other
to
be adversely
PTT and acted in a manner similar
characterized by steep topography and habitat types may change within relatively short linear distances.
is
The ram
ters
rams carrying conventional VHP transmitters. participated fully in the rut and was one of two
were dominant
in all social
large individuals that
encoun-
Although each collar was deployed on an adult female at

about the same time, the quantity of data from the two collars varied sharply. From 16 July until the end of September, the first
observed.
He remained
with a group of rams through
the remainder of the year

individual.
and was consistently a dominant
PTT provided 61
site to
locations. This bear
moved
from her initial capture

again.
satellite
another drainage, then back
The timing of these movements as indicated by the data was verified by conventional radio-tracking. Thus, satellite telemetry seemed to successfully indicate the timing of movements among salmon streams. The second PTT provided only 32 locations from its deployment on 1 8 July until the end of September. This bear
also
Although no data were available on his previous movement patterns, he followed what was generally believed to be the predominant movement pattern for sheep in the area of the Brooks Range (Fig. 33) and was
consistently found in areas occupied by other rams.
the
When
one year, some sliding of the collar along the neck was noted, with resultant matting and
after
PTT was removed
abrasion of hair along the dorsal surface. However, these affects appeared to be no different from those resulting
moved from her area of capture, but location frequen-
from
of
lighter
VHP transmitters.
on rams
However, the importance
cy dropped dramatically in August. Because of poor weather for flying, her exact location during this time
could not be verified. However, biologists might interpret the small number (2) of locations south of Karluk Lake
(Fig. 32) as
the rut
damage during was reemphasized with this heavier package. The 24-h activity index seemed to be generally correfitting collars
tightly to avoid
lated with periods of foraging
and migration, although not

in the
an unimportant foray, when
in fact, these
locations represented roughly the
same amount of time (or
unambiguously November and late June
so. Distinct
peaks
index during late
up
to nine times the levels seen
37
Grayback Mountain
10
Fig. 32. Locations of an adult female
20
brown bear on Kodiak Island during summer 1987. A. 18 July-3 August. B. 13 August-17 August. C. 5 September-29 September. Data courtesy of V. Barnes, U.S. Fish and Wildlife Service.
38
FISH
Kilometers
winter and spring 1986-87 and summer and Fig. 33. Locations of a Dall sheep ram in the Brooks Range, Alaska, during showing movement between the two seasonal ranges. Data courtesy of M. Hansen, University of Alaska.
fall
1987,
39
during late summer and winter coincided with the peak of the rut and spring migration, respectively (Fig. 34).
With further ground verification, the 24-h index was concluded to be useful as a reflection of general activity levels
for
10 Q
male Dall sheep.
Elk: Yellowstone National Park

Elk study in Yellowstone National Park by D. Vales focused on the social behavior and feeding strategies of
adult males, especially during winter. Vales placed PTT's on a yearling male and a 10-year-old male in early Sep-
collar
tember, just as the rut was beginning. The yearling elk's was fitted loosely, to allow further growth. The
fitted tightly,
J
Fig. 34.
M A M
ASOND
mature elk's collar was

that
it
with the expectation
would loosen
after the rut.
Annual changes in the 24-h activity index for a Dall sheep ram in the Brooks Range, Alaska. Data courtesy of M. Hansen, University of Alaska.
In addition to the information on general movements desired by park managers, this study sought quantitative data on behaviors during all times of day during winter and
further calibration of the short-term activity index for elk
(see Short-term Activity Index). Therefore, a duty cycle of
8
co V)
03
40 30
20
10
Locations
No
locations
6 h of transmission every 50 h was chosen so that as many times of the day as possible were sampled within each
Q-
4-week interval. This duty cycle deliberately spread information throughout the 24-h day but resulted in fewer locations. As expected, little information was gathered during those periods of low overpass frequency (Fig. 35). As with the Kodiak Island bear study, substantial variation in the performance of the two PTT's was noted. From
O
.0
(D
E
=3
deployment in mid-September 1987 through January 1 988, one PTT yielded 1 1 8 locations while the other yielded only 49. However, unlike the Kodiak Island study, there was no obvious relation between PTT performance and
topographic features or habitat selection (D. Vales, personal communication). Hours of transmission of the two
I
18
20 22
10
12
14
Hour of Day (UT)

Fig. 35. Daily pattern of data acquisition for two elk in Yellowstone National Park, September 1987-January 1988. Both
PTT's were
identical, as
were
relative
performances with
during
PTT's had duty cycles of 6 h on 44 h off, allowing sampling all hours of the day within each 25-day cycle. Data
regard to time of day. The two collars cant difference in signal strength (P
showed no
signifi-
courtesy of D. Vales, University of Idaho.
0.093, n = 27); the transmitter that produced fewer locations however, and poorer precision did have consistently lower signal
strength. Additionally, the motion detector in one of the two PTT's malfunctioned during winter. During visual observations of the instrumented animal, it was seen to walk and feed while the 60-s activity index continued to
assumed an average elevation was used, it yielded movement patterns much closer to those known from ground telemetry and visual locations (Fig. 36B).
Mule Deer: Southeastern Idaho

As
part of a larger study,
show only zero values. Movements of both elk were relatively restricted during winter, when they remained primarily in the Gardiner and
game
biologist C.
Brown
deployed four PTT's on adult female mule deer just before the hunting season in 1987. The primary objectives were
Mammoth
areas of the park (Fig. 36A). Because of the
high elevation of this area (1,640-2,300

level), locations calculated
above sea
assuming sea
level displayed
movement data during the hunting season and afterwards to judge whether cover use and behavior just patterns differed between those times. Two deer were
to obtain
fitted
considerable longitudinal error.
When
a correction that
with
.6-kg second-generation PTT's, and
two were
40
FISH
three of the four deer remained fairly sedentary, but one
made
major roads
a southerly movement that involved crossing a few in the area.
Moose: South-central Alaska

Mammoth NJ.
%
>J
In a study of
-.._
moose near
Wasilla, Alaska, biologist R.
Modaferri used conventional radiotelemetry. However, even when the weather allowed him to obtain locations, he
only gathered spot information about the activity patterns of these moose. An attempt was made, using Argos, to obtain detailed information on feeding and resting patterns
and
to
determine whether specific habitats were
used.
Two adult female moose were captured in December
1987 and fitted with second-generation PTT's. Both PTT's were programmed to transmit for 1 8 h every 3 days.
This resulted in up to 13 locations being obtained during each transmission period, followed by 54 h without
locations.
Locations within each of the 18-h periods were generally within 2-3 km of each other (Fig. 37). Because location errors are expected to be of approximately this
magni-
tude (Fig. 37, inset),

true
it
would be
difficult to discriminate
movements from "movements" caused merely by
telemetry error.
Activity patterns during winter were an additional focus of this moose study. The 24-h index was significantly correlated (Spearman rs = 0.543, P < 0.02) with the dis-
tance traveled between days of PTT transmission (estimated by calculating the minimum distance between the single best location from each 1 8-h transmission period;
Fig. 38).
Analyses of the short-term activity index have
not been completed.
Wolf: Northwestern Alaska

In April 1987, a 1.2-kg third-generation PTT with C-size lithium batteries was deployed on a male wolf in
Fig. 36.
during
old;
Movements of two bull elk in Yellowstone National Park fall 1987. Solid lines represent movements of a 12-year dashed lines represent movements of a yearling. A. Locaby assuming PTT's were
at
northwestern Alaska as part of a cooperative study between W. Ballard of ADFG and the AFWRC. This prototype PTT was used with VHP transmitters in a study to obtain daily movement data for wolf packs on the winter
tions calculated
sea level. B.
Locations adjusted by assuming a mean elevation for the study area. Data courtesy of D. Vales, University of Idaho.
range of the Western Arctic caribou herd. A primary obfor censusjective of the study was to develop procedures ing wolves on caribou winter range.
size and low temperawinter in the study area, the PTT tures (<-40 C) during was expected to transmit for only 6 months on a duty cycle
fitted
with 1.2-kg third-generation PTT's. The third-
Because of the smaller battery
generation PTT's were
programmed
to transmit 18 h/day,
beginning just before the hunting season. To prolong battery life, the transmission schedule changed to 6 h every 3
of 6 h on-42 h
(Fig.
days immediately following the hunting season. The duty cycle worked as planned and provided intensive coverage during the hunting season. During that time,
off. However, the PTT provided locations and sensor data until the wolf was shot by a 39)
hunter in late February 1988; it continued to transmit until June 1988. Data were received from 876 satellite over-
41
Captive
Moose
A.
Wild Moose
B.
Lake
o
Kilometers
o
Kilometers
Fig. 37. Movements of an adult female moose in south-central Alaska (A) during three nonconsecutive 1 8-h periods during winter 1988. Apparent "movements" of a nearly sedentary moose (B) were generated by randomly selecting from among successive locations of a captive moose within a 5-ha pen. Data courtesy of R. Modaferri, Alaska Department of Fish and Game.
passes between 1 April 1987 and 28 February 1988. Adequate data for calculating the wolf's location were obtained from
tween April 1987 and February 1988 was 2,618 km. Activity data provided by this prototype wolf PTT was
of
little
512 of these overpasses. Ballard obtained an
value. There
was no
significant correlation
(/
average of 3. 1 4.7 (standard deviation; SD) locations per day; at least one location was received on 92% of the 167 days the transmitter was active. The remaining 364 overpasses provided sensor data (e.g., canister temperature and short- and long-term indices of the wolf's activity) but no
location.
0.33, 21, 0.14) between mean distance traveled 2-week intervals and the mean long-term activity during
n =
P=
index. In contrast to
rest
work with other
species, periods of
and activity could not be discerned from the short-
The minimum
distance traveled by the wolf be-
term activity counts. The mercury switch within the canister was oriented parallel to the wolf's spine and to the
5,000
0.1
.c
4,000
0.08
^ 1 ^ $
Mte
Fig. 38. Relation between the 24-h activity index and rate of movement for an adult female
moose in south-central Alaska. Data courtesy

of R. Modaferri, Alaska Department of Fish
^ ~ <
"o
3,000
2,000
1,000
C 0.06
and Game.
.c
42
FISH
Kobuk Valley
National Park
Fig. 39.
Movements of an adult male wolf in northwestern Alaska April 1987-February

1988. Data courtesy of W. Ballard, Alaska Department of Fish and Game.
Kilometers
so
Selqwik
National Wildlife Refuge
bottom of the canister. In our studies of captive wolves, the canister rested against the wolf's chest, and the mercury
switch was activated by even slight body movements,
including breathing motions as the wolf rested. We attribute the seeming inability to detect activity patterns in the
lard
seem
and Fancy 1989). Larger estimates of home range to be the result of greater numbers of relocations,
detection of unusual movements, and
more
consistent
wolf
to
improper orientation of the mercury switch, and

that future researchers orient the anterior
1-6
coverage than that provided by conventional methods; this can be only partly explained by errors associated with locations determined by satellite. Consistent and frequent
relocation of wolves using satellites provides data sets for
we recommend
canister.
end of the switch +2

less sensitive to slight
still
relative to the
at the anterior
bottom of the
Switches elevated
end should be
evaluating wolf movements and home range that are superior to those provided by conventional methods, particularly in
body motions such
as breathing but
remote areas.
be activated by body movements during activity. Calibration studies to determine the best switch orientation
for
wolves need
(Ballard et
to
be conducted using captive wolves.
Walrus: Bristol Bay, Alaska

The status of the Pacific walrus population has been assessed with aerial surveys (Estes and Gilbert 1978), but most surveys include biases that can be difficult to quantify.
Six wolves are currently being tracked using the Argos
DCLS
that
1990). Preliminary analyses suggest ranges estimated from satellite-determined locations are 75% larger than those from relocations ob-
al.
home
In particular, walruses cannot be observed while div-
tained by conventional methods (fixed-wing aircraft; Bal-
ing but can be observed relatively easily
when hauled
out
43
on
ice or at traditional terrestrial sites.
Because of the vast
number of dives > 10 m deep; and the deepest dive during

These PTT's have generally worked well for 1-2 months but have failed thereafter. Reasons for the relatively short life spans are currently
the 24-h sampling period.
expanses and unpredictable weather off Alaska's western coast, quantifying patterns in diving and haul-out behaviors using traditional
VHP telemetry
and fixed-wing
air-
craft
ters
would be prohibitively expensive. Satellite transmitwere used to develop a method of quantifying these
being investigated.
behaviors, and thus of improving the reliability of subse-
quent aerial surveys. The behavior and habitats used by marine
mammals
Application and Sampling Considerations

In addition to field considerations, the quantity and
present special problems for satellite telemetry. Because salt water does not allow transmission of radio signals, a
saltwater switch
the animal
was used
to turn the transmitter
on when
saltwater
quality of data received using satellite telemetry require
was above
the water's surface.
The
consideration of data processing methods and analytical
switch also functioned as a sensor, quantifying the amount of time the animal spent out of the water during a sampling
period.
procedures.
Data Processing
For projects with few satellite collars deployed or simple objectives, it may be possible to analyze data without
computers. However, for most applications, the quantity and complexity of data necessitate computer processing.
August 1987, a prototype FIT designed for walrus was attached to a male walrus on Round Island near
In
Togiak, Alaska.
1987;
it
The PTT functioned
until
December
provided information on animal location, temperature, duration of the last dive recorded, average time
spent below the surface during the past 24 h, and number of dives during the past 24 h. Movements of this walrus
Computers allow for rapid storing, sorting, summarizing, mapping, and analyzing of data. Some tasks are impossible without the aid of a
computer: algorithms required to
from coastal areas
into Bristol
Bay and back during
fall
predict the time and location of satellite passes are too
1987 are shown in Fig. 40.
complex
to
be formulated without a computer. Also, com-
PTT's subsequently deployed on walruses have had the

temperature sensor replaced with a pressure sensor. In addition to animal location and the amount of time spent out of water, these units were designed to yield the amount
of time spent at depths of 0-4, 4-10, and >10 m; the
puting distances between locations and areas formed by polygons are tasks that cannot realistically be attempted
without computers. Many of our analyses used a geographic information system (GIS) to store, select, and map
locational data.
Fig. 40. Locations of an adult male walrus tracked by satellite in Bristol Bay, Alaska,
fall
1987. A. 14 August. B. 26 August.
C. 16 September. D. 18 October. E. 22 October. F. 14 November. Data courtesy of S. Hills, U. S. Fish and Wildlife Service.
44
FISH
Our work with satellite telemetry data required frequent

development of programs designed to assist data analysis (Fancy et al. 1988). We developed systems that were used
automatically to achieve specific objectives
jects using large quantities of satellite data have
locations falling within the specified
window and with

it
specified criteria
chose the best location offered
then
first
found the next cluster of locations, beginning with the
other pro-
developed
observation not in the previous cluster. Beginning in April 1987, criteria for choosing caribou locations were the
location with the highest LQ index (3 > 2 > 1 ) and, in case of a tie, the location calculated from the greatest number of
similar systems. Merrick and

series of
Mate (1985) developed a
programs
for dealing with satellite data for ceta-
ceans. Also the Wildlife- Wildlands Institute in Missoula,
Montana, has developed a series of programs to reformat Argos data and produce files that can be manipulated on a
messages. Other criteria that might be used include choosing locations estimated by the best satellite overpass with
elevation closest to the
microcomputer by dB ASE
III+.
estimated
variation.
optimum (see Fig. 8) or those when the PTT displayed minimum temperature
summary
The system we developed had three components: a data stage, in which Argos data were summarized
from an overpass in a single stage, in which smaller files were
into files with all information
sulting display of animal
Choice of a time window substantially altered the removements. Figure 41 portrays
record; a differentiation
the
movements of an adult female caribou from the Porcu-
created consisting only of information from overpasses fixing a location; and a formatting stage, in which these smaller files were converted to GIS -compatible formats
for presentation either as location points or vectors be-
pine herd as she traveled from her wintering area toward her eventual calving site. The general movement pattern remained unchanged, regardless of which location fre-
tween successive locations. In
this last stage,
summary
quency was used, but short-term movements were progressively less evident as shorter time windows were used.
statistics were also computed. We also adapted a NASA program for predicting the times and characteristics of satellite orbits. Predictions were used to determine optimum duty cycles for transmitters and to synchronize direct
Fancy et al. (1989) used a 1-h window to assess movement

patterns of Alaskan caribou.
Even
after selecting
among
locations within a window,

to
the locations occasionally
seemed
be biologically un-
observations of an animal with satellite overpasses to evaluate activity sensors or location accuracy.
reasonable.
The program
incorporated algorithms that flagged a location whenever the animal's calculated rate of move-
We
calculated satellite azimuth, elevation, and range at all times during each overpass. Description of an earlier version of these programs
is
provided by Fancy
et al. (1988).
ment exceeded a specified tolerance, which was unique to each species. When successive locations were closely spaced in time, we found this method helpful.
Independence of Successive Locations Independence of successive observations
Sampling Concerns
Selecting Locations
is critical
for
some statistical analyses of animal movements, but independence can be violated when observations are closely
error
is
As with conventional telemetry,

in location estimates
always present
obtained from
satellite telemetry.
clustered in time, as often occurs with satellite telemetry data. Schoener (1981) devised a procedure to assess the
clear
example was when the two satellites passed over an animal within 10-15 min of each other: animals some-
independence assumption. Swihart and Slade (1985a) derived a test of significance for deviations from the expected value of Schoener 's ratio and, by doing so, devel-
times appeared to
mals' rates of
make
ments" from one location
spectacularly quick "moveto another. Estimates of ani-
oped a method to determine whether a given data set meets

the independence assumption.
movement would have become inflated if these apparent movements (many of which were attributable to telemetry error) were included in analyses. review here two suggested algorithms for choosing among
They
also suggested that a

still
data set failing to achieve independence could
be used
We
by systematically excluding observations (thereby increasing the elapsed time between successive observations) until the resulting series satisfied the
criterion.
competing locations
in
such situations. Both create an
independence
objective set of rules to govern selection of data for analysis, although neither solves the problem of error.
We examined some monthly series of satellite-obtained

locations from different species, calculating Schoener 's ratio and Swihart and Slade's critical value each time.
The algorithm we used allowed us to specify a time window during which only one location was to be selected
for inclusion with the resulting data. This
window was
Most data
sets
we examined
failed the test of indepen-
varied, depending on the objectives of the analysis and the PTT's duty cycle. The algorithm identified the cluster of
dence, although considerable variation among species and seasons was noted. For example, the movements of a
45
A.
Beaufort
2i
Brown Bear
Alaska
20
2
30
40
50
60
Moose
B.
Beaufort
&
J
CD
1.5
seo
-C
O O
0.5
Alaska
CO
10
20
30
40
50
60
2i
Mule Deer
1.5
0.51
10
20
30
40
50
60
Hours between Successive Locations

Fig. 42. Relation between Schoener's ratio (solid line) and its critical value (dashed line) and the time interval between
successive locations. Statistical independence is achieved when the ratio exceeds its critical value (Swihart and Slade
1985b). A. Brown bear on the arctic coast, July 1987. B. Moose in south-central Alaska, January 1988 (data courtesy of R. Modaferri, Alaska Department of Fish and Game). C. Mule
Fig. 41.
site in
1987 from
Movements of a Porcupine herd caribou during June its wintering area in Yukon Territory to its calving
northern Alaska. A. All data are plotted. B. Only the best location estimates within each 1-h "window" are plotted.
deer in southeastern Idaho, October 1987 (data courtesy of C. Brown, Idaho Department of Fish and Game).
C. Only the best location estimates from each 24-h

are plotted.
"window"
when
were considered.
the arctic slope in Alaska during July were highly autocorrelated when all data were considered (Fig. 42A). These data only met the independence criterion
locations taken at approximately two-day intervals If the data set were to be considered
brown bear on
in this way, total sample size during the month would be reduced from 61 locations to 10. Similar analysis of
the
movements of a moose
in
Alaska during January
46
FISH
resulted in very different conclusions (Fig. 42B). Here,
based on three situations where
satellite
telemetry was
dependence among
successive locations seemed to
be only a minor problem. However, locations from a mule deer in Idaho were far from achieving independence,
used as an alternative to conventional telemetry (Table 17). Satellite collars, each including a VHP transmitter,
were assumed
even when locations were restricted to one every two days (Fig. 42C).
to cost $3,300 each and were to be replaced 3-4 times during a 5-year study, given a transmitter life of 12-18 months. Each VHP transmitter cost $330 and had
Time of Sampling
Many analyses require that

we used were
the
locations be a
ple of all the true locations of an animal.
random samMost duty cycles

at
an assumed life of 3 years, needing replacement only once. Then, a second transmitter was purchased for each animal to replace the used collar when the animal was recaptured.
The Argos processing
regular
that
is,
transmissions occurred
same time during each transmission-day. Thus, sampling was more nearly systematic than random. Systematic sampling can sometimes be substituted for random
sampling with
arise
tern.
fee was assumed to be $8.22/day per transmitter, or $3,000 per transmitter-year. Labor costs were not included in these examples.
These examples suggested that satellite telemetry is most cost effective in situations where air charter costs are
high and a large area must be searched to relocate all radiocollared animals, as with the Porcupine caribou herd (Fig.
43). In this example,
when
adverse effect, although problems can systematic sampling matches an existing patlittle
Such a
situation
may occur when sampling
locations
at regular intervals, especially
with duty cycles having a
we assumed that caribou could be anywhere within the herd's range during each tracking
24-h period or integer multiple thereof.

display circadian rhythms that
may
Many species coincide with such
sampling periods (Swihart and Slade 1985a), potentially

biasing the data.
flight. Therefore, the entire range required searching regardless of the number of radio collars deployed, and the per-animal cost to relocate caribou when 50 collars were
deployed was
also
20%
of the cost to relocate 10 collars.
We
accuracy was comparable to or better than that obtained using satellite telemetry and that
that location
assumed
Cost Comparisons
some situations, satellite telemetry may be the only means of acquiring data necessary to meet study objecIn
tives. In
each radio-collared caribou therefore had

visually.
to
be located
Given these assumptions,

if
satellite
telemetry was
three
cost effective
(n
study objectives required more than
= -
10) caribou or 13 (n
= 50)
locations per year. If daily
most cases conventional telemetry may also be used, and the costs of the two approaches may be a factor in determining which is best for a particular study. Unforit is
locations were needed, costs using

(n
10) or 10 (n
VHP telemetry were 43 times higher than those using 50)
satellite telemetry.
tunately,
son between
satellite
not possible to make a single cost compariand conventional telemetry, because
Radio-tracking costs in the Kodiak Island brown bear example (Fig. 44) were only 5% of those in the first
costs can vary greatly
between different study areas for animal capture, air charter, and other factors. Each researcher must determine the costs for their own study.
example because of the smaller study area, and lower air charter
ters
size
and location of the
costs. Satellite transmit-
The following hypothetical
cost comparisons were
in
were programmed to transmit only one day each week winter while the bears were in their dens, and therefore
Table 17. Cost comparison between satellite and

Transmitter
VHP
telemetry systems for hypothetical 5 -year studies of 3 species.
47
n=io n=50
10
20
30
VHP telemetry.
40
50 100
300
Locations/Year
Fig. 43. Cost-benefit analysis of satellite versus conventional
Porcupine caribou herd example.
transmitter life
example,
than 62 (n
satellite
was increased to at telemetry was
least
8 months. In this
if
cost effective
more
reduced, and 315 locations each year were required before satellite telemetry became cost effective.
needed to meet study objectives.
10 collars deployed) locations per year were If 50 collars were de-
Directions for Future Research

Our work involved various
applications of satellite te-
ployed, satellite telemetry would only be cost effective if more than one location per day were required. The cost to
obtain daily locations using telemetry was three times that of using satellite telemetry for the 10 collar
VHP
lemetry in wildlife research. Many of these were not possible just two or three years ago. We expect that continued
example; costs were similar
if 50 collars were deployed. The third example compared costs for a study of mule deer movements in Idaho (Fig. 45). Clover traps (Clover
work on both
refine the
is
list
the technical and analytical aspects will

satellite
of applications for which
telemetry
appropriate.
1956) were used to capture deer for the first time, but recaptures required the use of a helicopter and net gun.
Radio-tracking costs were again low compared to the Porcupine caribou herd example. If 10 collars were deployed,
satellite
researchers could consider applications for sateltelemetry if the precision and accuracy of location estimates were improved. Many have expressed doubts
lite
More
telemetry
was cost
effective
when
at least
42
about using satellite data for analyzing habitat use on as fine a scale as is desired, primarily because of imprecision
of locations. Improved precision in the future might
locations per year locations using
were needed. The cost

telemetry
to obtain daily
that of
come
VHP
was four times
from improvements
that
in the
PTT itself, from the algorithms
using satellite telemetry. In the 50 collar example, the peranimal cost to relocate deer using VHP telemetry was
used to calculate locations, or in analysis routines in a GIS can correct exactly for elevational bias.
48
FISH
n=lo n=50
o
i/
10
30
VHP telemetry.
40
50 100
300
Locations/Year
Kodiak brown bear example.
We have documented a useful role for the motion sensor

currently in use
An
additional limitation
is
battery
life.
Most PTT's we
generating 24h and 60-s activity indices. However, both indices had limitations, even on those species experiencing the most
that
is,
on Telonics PTT's
deployed had a one-year life expectancy. Some bear collars had two years expected life spans, but none have yet
lasted
success. For caribou,
not successfully calibrated indices with behaviors observed in the wild. For reliable
estimates of activity budgets, this research Further development of the motion sensor
sary to calibrate either index.
It
we have
individuals over
for
more than one year. Many studies monitor the same numerous years, requiring yearly capture replacing PTT's. Each time an animal is handled it is
to risk of injury or death; also, research
is still
needed.
exposed
budgets
may
be neces-
are strained.
may
yet be possible to
refine the ability of the 60-s index to discriminate
among
Some limitations of the present system may be overcome with more sophisticated analytical treatment. Examples include corrections for elevations (other than those assumed by Argos) by way of a sophisticated GIS, and the
activity types (e.g., standing still versus lying still, grazing versus browsing), but this will require further development of the sensor itself. Issues such as whether multiple
sensors in different configurations would produce improvements, or whether sensors might be placed in remote locations on the animal, may be fruitful areas for further
development of correction factors for autocorrelated data that would allow use of complete data sets without violating important statistical assumptions. The exploration of time-series approaches toward wildlife data has been sug-
work. Other sensors with potential applications in wildlife research may include devices for measuring battery voltage, atmospheric pressure
elevation), heart rate, and
(i.e.,
tula
gested by some statisticians (Dunn and Gipson 1977; Panand Pollock 1985). These techniques should be ex-
determining the animal's
plored by biologists and statisticians confronted with

these problems.
body temperature.
As methods
are developed that incorpo-
49
n=lO n=50
'
'
'
'
'
'
'
'
i/iii
10
20
30
VHP telemetry.
40
50 100
300
Locations/Year
Idaho mule deer example.
rate telemetry error into statistical analyses of location, concerns about using imprecise data for habitat analysis may be partially alleviated.
location
is
the only alternative, satellite telemetry can
substantially reduce the risk of flying during the hazardous
Other areas for future development no doubt exist. This document has been prepared not only to report on the
current state of the art but to encourage others to consider the potentials of the technology with an eye toward
conditions frequently encountered in wildlife work. The appropriateness of satellite telemetry depends on study objectives. Advantages of satellite telemetry are
notable in cases where objectives require intensive data on individual animals, where movement information is decially at night or during inclement weather.
improvement.
Summary and
Satellite telemetry
Conclusions
where animals move long distances, espeAdvantages are minimized where objectives require modest amounts
sired daily, or
of data on
many
individuals or where animals either move
can circumvent many of the defi-
only slightly or are otherwise easily tracked from the ground or air. The lack of accuracy and precision of locations obtained
ciencies encountered with conventional telemetry. Factors such as hazardous weather, darkness, international boundaries,
ellite
from the current system
limits
its
appli-
cability for habitat selection studies to those in
which
and extensive animal movements do not hinder
sat-
telemetry systems. In addition to location information, sensors within satellite-compatible transmitters can
coarse-grained definitions of habitat types are used. Using these techniques, we have greatly increased our ability to monitor northern species, such as caribou, polar
bears,
monitor aspects of an animal's environment and behavior.
For some applications,

initial costs, is
high more cost effective than conventional tel-
satellite telemetry, despite
emetry. Perhaps most importantly, in areas where aerial
and muskoxen. New applications await other researchers. Despite limitations, satellite telemetry has unique potential as an operational tool for wildlife researchers.
50
FISH
Acknowledgments
This study was supported by administrative funds from the Pittman-Robertson program and by the U.S. Fish and
Wildlife Service and cooperating agencies.
We thank J.
Russell and S. Tomkiewicz, Telonics, Inc.,
for their efforts in technical matters; D. Clark, L. Morakis,
and A. Shaw of Service Argos helped us
to understand
and
We
thank R.
use the Argos system more effectively. Editorial improvements were suggested by M. Fuller, C. Halvorson, S.
Cameron, D. Reed, W. Regelin, and K. Whitten of Alaska Department of Fish and Game (ADFG); G. Elison, M.
Masteller, F. Mauer, G. Muehlenhardt, and G. Weiler of
Tomkiewicz, and an anonymous reviewer.
Arctic National Wildlife Refuge
(ANWR);
C. Gardiner of
References
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Alaska Fish and Wildlife Research Center
(AFWRC); M.
Anderson, D.
Hansen of the University of Akrka; W. Nixon and D. Russell of the Canadian Wildlife Service; and R. Hayes
and C. Smits of the Yukon Department of Renewable Resources for their assistance with field operations. C.
Curby, F. D'Erchia, J. Greslin, M. Hansen, M. Koschak, C.
Metzler, D. Reed, R. Slothower, and
and location system. Service Argos, Toulouse, France.
36pp.
Argos. 1984. Location and data collection satellite system user's guide. Service Argos, Toulouse, France. 36 pp.
Ballard,
Venable provided computer programming and data analysis. We thank M. Hansen, S. Hartz, M. McClure, S. Morstad, T. Paragi, and
J.
W.
B.,
and
S.
G. Fancy. 1989.
Satellite radio-
U. Petersen for working with captive animals. D. Guthrie and R. White of the University of Alaska, B. Davitt and C.
tracking of wolves.
IUCN/SSC Wolf Specialist Group,
University of Alaska, Fairbanks. 17 pp.

Ballard,
Robbins of Washington State University, and C. Schwartz of ADFG provided assistance and access to captive animals for the
Enzweiler,
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work. B. Blevins, C. Curby,
J.
W. B., L. A. Ayres, S. G. Fancy, D. J. Reed, K. E. Roney, and M. A. Spindler. 1990. Demography and movements of wolves in relation to the western Arctic
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valuable ideas and constructive criticisms.
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New
York. 37 1 pp.
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Beaty, D. W., S.
M. Tomkiewicz,
and
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Carter. 1987.
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studies
Wildl. Manage. 52:11-13. Boertje, R. D. 1981. Nutritional ecology of the Denali
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M.
S. thesis, University
of Alaska, Fair-
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Keating, Glacier National Park; J. Squires, University of Wyoming; K. Aune and R. Mace, Montana Department of
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M.
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Courrouyan, P. 1987. CML 86. Argos Newsl. 32:11. carCraighead, D. J., and J. J. Craighead. 1987. Tracking ibou using satellite telemetry. Natl. Geogr. Res. 3:462479.
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serves special mention.
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Estes, A., and J. R. Gilbert. 1978. Evaluation of an aerial survey of Pacific walruses (Odobenus rosmarus diverJ.
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J.
Fish. Res.
Board Can. 35:1 130-1 140.
65 pp.
Pank, L.
F., W. L. Regelin, D. Beaty, and J. A. Curatolo. 1985. Performance of a prototype satellite tracking system for caribou. Pages 97-1 1 8 in R. W. Weeks and F. M.
Fancy,
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S. G., L. F.
Garner, S.
Pank, D. C. Douglas, C. H. Curby, G. W. C. Amstrup, and W. L. Regelin. 1988. Satel-
telemetry: a
new tool for wildlife research and man-
agement. U.S. Fish Wildl. Serv., Resour. Publ. 172. Fancy, S. G., L. F. Pank, K. R. Whitten, and W. L. Regelin.
1989. Seasonal
as determined
Long, eds. Proceedings of the 5th international conference wildlife biotelemetry. Laramie, Wyo.
Pank, L.
1987.
F., S.
movements of caribou
by
satellite.
in arctic
Alaska
Can.
J.
Zool. 67:644650.
G. Fancy, M. C. Hansen, and W. L. Regelin. Remote sensing of animal location and activity
biologists 13. In press.
French,
1986. Environmental housings for animal PTT's. Argos Newsl. 26:7-9.

J.
via satellite. In Proceedings of the international con-
gress
game
Fuller, R., N. Levanon, T. E. Strikwerda, W. S. Seegar, J. Wall, H. D. Black, F. P. Ward, P. W. Howey, and J.
M.
Pantula, S. G.,
and K. H. Pollock. 1985. Nested analysis of variance with autocorrelated errors. Biometrics and J. Hechtel. 1980. Big game investiga-
Partelow. 1984. Feasibility of a bird-borne transmitter for tracking via satellite. Pages 1-6 in Proceedings of
41:909-920.
Reynolds, H.
V.,
symposium on biotelemetry, 6-12 1984, Dubrovnik, Yugoslavia. May Garner, G. W, S. C. Amstrup, D. C. Douglas, and C. L. Gardiner. 1989. Performance and utility of satellite tethe 8th international
tions. Structure, status, reproductive biology,
move-
ments, distribution
and habitat
utilization of a grizzly
lemetry during field studies of free-ranging polar bears in Alaska. Pages 67-76 in C. J. Amlaner, Jr., ed. Bio-
bear population. Fed. Aid Wildl. Rest. Proj. W-17-11, Job Prog. Rep., Alaska Dep. Fish Game, Juneau. 66 pp. Reynolds, P. E. 1987. Movements and activity patterns of
a satellite-collared
life
symposium on biotelemetry. University of Arkansas Press,

Fayetteville.
telemetry proceedings, 10th international
muskox in the Arctic National WildRefuge, Alaska, 1984-85. Pages 632-648 in G. W.
Garner and
P. E. Reynolds, eds. 1985 update report of the baseline study of the fish, wildlife, and their habi-
Gillingham, M. P., and F. L. Bunnell. 1985. Reliability of motion-sensitive radio collars for estimating activity of
black-tailed deer.
U.S. Fish Wildl. Serv., Anchorage, Alaska. Reynolds, P. E. 1989. An experimental satellite collar for
tats.
Harvey, M.
J.,
J. Wildl. Manage. 49:951-958. and R. W. Barbour. 1965. Home range of
muskoxen. Can.
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Zool. 67:1 122-1 124.
Microtus ochrogaster as determined by a modified min-
Roby, D. D. 1978. Behavioral patterns of barren-ground caribou of the Central Arctic herd adjacent to the TransAlaska pipeline. M.S. Fairbanks. 200 pp.
Schoener, T. W. 1981.
thesis, University of Alaska,
imum
area method.
J.
Mammal. 46:398-402.
Johnsen, H. K., A. Rognmo, K. J. Nilssen, and A. S. Blix. 1985. Seasonal changes in the relative importance of different avenues of heat loss in resting and running
reindeer.
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empirically based estimate of
home
range. Theor. Popul. Biol. 20:281-325.

P.
Acta Physiol. Scand. 123:73-79.
Strikwerda, T. E., H. D. Black, N. Levanon, and
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Le Traon, P. Y. 1987. Argos location accuracies. Pages 259-280 in Service Argos, Inc., International users conference and exhibit, 15-17 September 1987, Greenbelt,
Howey. 1985. The bird-borne

kins
APL
Johns Hop(Appl. Phys. Lab.) Tech. Dig. 6:60-67. Johns

transmitter.
Hopkins University, Baltimore, Md.

Strikwerda, T. E.,
Md.
Mate, B. R. 1987. Development of satellite-linked methods of large cetacean tagging and tracking in OCS lease areas. Unpubl. final report. OCS Study 87-0038, Minerals Management Service. Alaska Outer Continental Shelf Program Office, Anchorage. Mate, B. R., D. Beaty, C. Hoisington, R. Kutz, and M. L. Mate. 1983. Satellite monitoring of humpback whale
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and H. D. Black. 1986. Bird-borne
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and location program. Johns Hopkins APL (Appl. Phys. Lab.) Tech. Dig. 7:203-208. Swihart, R. K., and N. A. Slade. 1985a. Testing for independence of observations in animal movements. Ecol-
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diving behavior and movements. Unpublished progress report to Minerals Management Service. Alaska Outer
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Wildl. Manage. 49:1019-1025.
size. J.
52
FISH
GLOSSARY
An index derived from a motionActivity Index device on a PTT that, when calibrated to known sensing
can be used to estimate activity patterns of free-ranging animals wearing PTT's
activity,
Independence (of Spatial Data)

tistically
Locations are
at
sta-
independent when locations not a function of locations at time x
time x
are
LCO
Location class zero; a processing option offered
by Argos that will estimate PTT location from as few as two messages. Specially designed for the animaltracking community, this service also provides additional data on location quality when normal processing
fails,
ACWRU
Unit
Alaska Cooperative Wildlife Research

Alaska Department of Fish and Game Alaska Fish and Wildlife Research Center
Arctic National Wildlife Refuge
ADFG
AFWRC ANWR
Argos
system
LQ
and provides the alternate location Index Location quality index; ranging from
to
The name of
that collects
3, used by Argos to guide users regarding the probable precision of a location estimate. LQ1 is the least pre-
the organization operating the
cise;
and processes data received by
LQ3
the
most precise
satellite-
LUT
from
Local user terminal; a computerized
polar orbiting Tiros-N satellites Argos data collection and location sysArgos DCLS
tracking system
that receives real-time transmissions
tem (see Argos)
Azimuth
A horizontal direction expressed in degrees clockwise from an adopted reference direcmeasured

tion, usually true north
satellites above the horizon and processes Argos data contained with the transmitted signal; also known
as direct readout stations
Message
The
signal sent
by a
PTT
to the satellite,
Doppler shift a signal on
Perceived variation in the frequency of the electromagnetic spectrum caused by

the source or the receiver
pattern of active-inactive
consisting of identification code, sensor data, etc. An orbit that passes directly over both Polar Orbit
movement of either
Duty Cycle
Efficiency
PTT
Programmed
geographic poles Platform transmitter terminal; any transmitter used to send messages to Argos instruments on a
satellite
transmission periods for a PTT The rate over time at which location esti-
UHF
PTT's
in this
mates and sensor data from a

investigator
PTT
are received
by the
to transmit to the satellite
Ultra-high frequency; frequency band used by is with(401.650
MHz
Elevation (PTT) meters) of the

location
is
The elevation above sea
level (in
PTT
(or animal wearing
it)
when
its
UT VHP
VS WR
large
band) Universal Time; Greenwich
Mean Time
in
Very high frequency; frequency band used
estimated by Argos The elevation above the horiElevation (Satellite)
conventional radiotelemetry
zon
(in degrees)
of the
satellite as
it
makes
its
closest
Voltage standing wave ratio; reduction of effective radiated power through an antenna caused by a
approach to the
PTT
mass being close to the antenna.
VS WR can result
by
GIS
Geographic information system; computer software used to analyze and display spatially oriented data
in loss of data because transmissions are not received
the satellite.
list
of current Fish and Wildlife Technical Reports follows.
1.
Effects of
Weather on Breeding Ducks
in
North Dakota, by Merrill C.
Hammond
and Douglas H. Johnson. 1984.

F. Hill
17pp.
2.
Lethal Dietary Toxicities of Environmental Contaminants and Pesticides to Coturnix, by El wood B. Camardese. 1986. 147 pp.
Effects of Vegetation Manipulation Kantrud. 1986. 15 pp.
and Michael
3.
on Breeding Waterfowl
in Prairie
Wetlands
A Literature Review, by Harold A.
4.
XYLOG: A Computer Program for Field Processing Locations of Radio-tagged Wildlife, by Wendell E. Dodge and
Alan
J.
Steiner. 1986.
22 pp.
5.
6.
Response of Lake Trout and Rainbow Trout to Dietary Cellulose, by H. A. Poston. 1986. 6 pp. DDE, DDT + Dieldrin: Residues in American Kestrels and Relations to Reproduction, by Stanley N. Wiemeyer, Richard D. Porter, Gary L. Hensler, and John R. Maestrelli. 1986. 33 pp.
Bird Behavior and Mortality in Relation to Power Lines in Prairie Habitats, by Craig A. Faanes. 1987. 24 pp. Limnological and Fishery Studies on Lake Sharpe, a Main-stem Missouri River Reservoir, 1964-1975, by Fred C.
June, Lance G.
7.
8.
9.
Trends
in
Beckman, Joseph H. Elrod, Gerald K. O'Bryan, and David A. Vogel. 1987. 83 pp. Spawning Populations of Pacific Anadromous Salmonids, by Gregory W. Konkel and John D. Mclntrye.
and Bioavailability of Mercury
in
1987. 25 pp.
10. Factors Affecting the Mobilization, Transport,
Reservoirs of the Upper Missouri
11.
12.
River Basin, by Glenn R. Phillips, Patricia A. Medvick, Donald R. Skaar, and Denise E. Knight. 1987. 64 pp. Waterfowl Status Report, 1980, by Albert N. Novara, James F. Voelzer, and Arthur R. Brazda. 1987. 93 pp. Waterfowl Status Report, 1981, by Albert N. Novara and James F. Voelzer. 1987. 99 pp.
13. Migration, Harvest,
and Population Characteristics of Mourning Doves Banded in the Western Management Unit, 1964-1977, by Roy E. Tomlinson, David D. Dolton, Henry M. Reeves, James D. Nichols, and Laurence A.
McKibben. 1988. 101 pp.

14. 15.
Waterfowl
Bird
Damage to Ripening Grain: An Overview, by C. Edward Knittle and Richard D. Porter. 1988. 17 pp. Damage to Sunflower in North Dakota, South Dakota, and Minnesota, 1979-1981, by Roger L. Hothem,
16. Influence
Richard W. DeHaven, and Steven D. Fairaizl. 1988. 1 1 pp. of Environmental Factors on Blackbird Damage to Sunflower, by David L. Otis and Catherine M. Kilburn.
of a Simulation Model to Decisions in Mallard Management, by Lewis
1988. llpp.
17. Applications
M. Cowardin, Douglas H.
18.
Johnson, Terry L. Shaffer, and Donald W. Sparling. 1988. 28 pp. Chemical Characteristics of Prairie Lakes in South-central North Dakota
Fish and Wildlife, by George A. Swanson,
Their Potential for Influencing Use by
Thomas
C. Winter, Vyto A. Adomaitis, and James

for Restoration,
W. LaBaugh. 1988.
19.
44pp. American Wildcelery (Vallisneria americana): Ecological Considerations and William L. Green. 1988. 24 pp.
by Carl E. Korschgen
20.
Temporal and Geographic Estimates of Survival and Recovery Rates for the Mallard, 1950 through 1985, by Diane S. Chu and Jay B. Hestbeck. 1989. 57 pp.
21 Migration of Radio-marked
.
Whooping Cranes from the Aransas-Wood Buffalo Population: Patterns of Habitat Use, Behavior, and Survival, by Marshall A. Howe. 1989. 33 pp. 22. Electrofishing, Power Related Phenomenon, by A. Lawrence Kolz and James B. Reynolds. 1989. 24 pp.
23.
A Computer Program for Sample Size Computations for Banding Studies, by Kenneth R. Wilson, James D. Nichols,
and James E. Hines. 1989. 19 pp. Program CONTRAST A General Program for the Analysis of Several Survival or Recovery Rate Estimates, by James E. Hines and John R. Sauer. 1989. 7 pp.
24.
25. Techniques for Shipboard Surveys of Marine Birds, by Patrick J. Gould and Douglas J. Forsell. 1989. 22 pp. 26. Nesting Ecology of Golden Eagles and Other Raptors in Southeastern Montana and Northern Wyoming, by Robert
L. Phillips,
27. Distribution
Anne H. Wheeler, Nick C.
Forrester,
J.
and Abundance of Golden Eagles and Other Raptors
Michael Lockhart, and Terrence P. McEneaney. 1990. 13 pp. in Campbell and Converse Counties, Wyoming, by
Robert L. Phillips and Alan E. Beske. 1990. 31 pp. 28. Genetic Differentiation of Walleye Stocks in Lake St. Clair and
29. Dicofol (Kelthane) as an environmental contaminant:
A review, by Donald R.
Western Lake Erie, by Thomas N. Todd. 1990. 19 pp. Clark, Jr. 1990. 37 pp.
TAKE PRIDE
in
U.S.
FISH
America
AND WILDLIFE SERVICE
DEPARTMENT OF THE INTERIOR
As the Nation's principal conservation agency, the Department of the Interior has responsibility for most of our nationally owned public lands and natural resources. This includes fostering the wisest use of our land and water resources, protecting our fish and wildlife, preserving the environmental and
and historical places, and providing for the enjoyment of life outdoor recreation. The Department assesses our energy and mineral resources and works to through assure that their development is in the best interests of all our people. The Department also has a major responsibility for American Indian reservation communities and for people who live in island territocultural values of our national parks
ries
under U.S. administration.

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Tracking Wildlife by Satellite: Current Systems and Performance

UNITED STATES DEPARTMENT OF THE INTERIOR FISH AND WILDLIFE SERVICE

Fish and Wildlife Technical Report

Technical Information Service (NTIS), 5285 Port Royal Road, Springfield,

Bird Rescue Research Center

San Francisco, California 2006

W. Garner, Steven C. Amstrup, Thomas R.

McCabe, and Larry

UNITED STATES DEPARTMENT OF THE INTERIOR FISH AND WILDLIFE SERVICE

Gathering Wildlife Data

Efficiency Precision and Accuracy of Locations

Long-term Activity Index

Overview and System Description Performance

Caribou: Northern Alaska and Yukon

Bear: Western Brooks Bear: Kodiak Island

Dall Sheep: Brooks Range Elk: Yellowstone National Park

Mule Deer: Southeastern Idaho

Data Processing Sampling Concerns

Summary and Conclusions

Current Systems and Performance

Missoula, Montana 59812

Anchorage, Alaska 99503

Anchorage, Alaska 99503

Anchorage, Alaska 99503

AND WILDLIFE TECHNICAL REPORT 30

of locations were <1,732

true location. Argos's

location class zero

Long-term and short-term indices of animal

calibrated to specific activity categories (e.g., lying, feeding, walking).

and problems with lack of independence among locations are discussed.

mule deer (Odocoileus hemionus)

Alaska and elk (Cervus elaphus) and in the Rocky Mountain

To provide the reader a more complete understanding of

that operates in the very

Argos system and

its applications to tracking animals, included an updated and shortened overview of

Argos system as presented by Fancy

ment, particularly for species that

Our experience has primarily been with

malian herbivores and carnivores, and our conclusions are

often constrained by the high cost of locating the animal,

reliable transmitters small

satellites to obtain data

on free-ranging animals is expandtechnology and improved equipment are

Alaska Fish and Wildlife Research Center

use by prospective users of the technology.

dian Wildlife Service,

Yukon Department of Renewable

Arizona), have participated

development of this of our studies on the re-

summarize our experiences using the and behavioral data on

ters that distribute results to users

TRACKING WILDLIFE BY SATELLITE

and receiver. The frequency

ceived by instruments on the

higher than the

away from the transmitter (Fig.

When the received and

near-polar, sun-synchronous orbit of the Tiros-N