THE SOUTHWESTERN NATURALIST 50(2):223–229 JUNE 2005

TRACK COUNT CALIBRATION TO ESTIMATE DENSITY OF

WHITE-TAILED DEER (ODOCOILEUS VIRGINIANUS) IN MEXICAN DRY
TROPICAL FOREST

SALVADOR MANDUJANO*

Departamento de Biodiversidad y Ecologı́a Animal, Instituto de Ecologı́a A. C., Km 2.5 Carretera Ant. a Coatepec No.
351 Congregación del Haya, Xalapa 91070, Veracruz, México
*Correspondent: mandujan@ecologia.edu.mx

ABSTRACT Track counting is a quick, easy, cost-effective technique to estimate the population
density of white-tailed deer (Odocoileus virginianus). By employing the double-count procedure, I
generated 2 models in which track counts were calibrated to estimate the density of deer in dry
tropical forest in Chamela on the Mexican Pacific Coast. For both models, I calibrated a track
index using density obtained from the line-transect method as a reference. The first model was
based on simple linear regression and the second on the strip-transect method. I discuss the
usefulness of these models for monitoring local populations as well as possible applications in
other regions.

RESUMEN Contar huellas es una técnica fácil, rápida, y poco costosa para estimar la densidad
poblacional del venado cola blanca (Odocoileus virginianus). Empleando el procedimiento del do-
ble conteo, generé 2 modelos en los que se calibra el conteo de huellas para estimar la densidad
de venado en el bosque tropical seco de Chamela, en la costa Pacı́fica de México. En ambos
modelos, calibré el conteo de huellas usando como referencia la densidad obtenida con el método
de conteo en transectos de lı́nea. El primer modelo se basó en una regresión lineal simple, y el
segundo en el método de transecto de franja. Discuto la utilidad de estos modelos para monitorear
poblaciones locales ası́ como la posibilidad de aplicaciones en otras regiones.

The double-sampling procedure has been
used to estimate the population density of
many taxa (Caughley, 1974; Pollock and Ken-
dall, 1987; Hone, 1988a). It has, for example,
been applied to crocodiles (Bayliss et al.,
1986), emu (Caughley and Grice, 1982), and
mammals as diverse as kangaroos (Coulson
and Raines, 1985; Barnes et al., 1986; Short
and Hone, 1988), weasels (Graham, 2002),
lynx (Andrén et al., 2002), feral pigs (Hone,
1988b), pronghorn (Firchow et al., 1990), and
deer (Floyd et al., 1979; Bobek et al., 1986; Es-
cos and Alados, 1988; White et al., 1989; Fuller,
1991; Potvin et al., 2002). Double sampling
consists of 2 phases (Eberhardt and Simmons,
1987). During the first phase, 2 independent
methods are applied simultaneously; the den-
sity estimate obtained with the first model is
considered more accurate but costlier to apply
in the field, whereas the second yields less re-
liable results but is easy and inexpensive. Dur-
ing the second phase, the estimate obtained
with the less precise method is fitted to that of
the more reliable method. If there are paired
data between an abundance index and density,
the index can be calibrated with a linear re-
gression fit (McCaffery, 1976; Eberhardt, 1978;
Hone, 1986; Parer and Price, 1987; Menkes et
al., 1990).
It is not easy to select a method of determin-
ing density of populations in tropical forests
that is accurate, easily applied in the field, and
inexpensive (Duckworth, 1998; Caro, 1999). In
the case of white-tailed deer (Odocoileus virgi-
nianus), methods used elsewhere ( Jeter, 1965;
Jenkins and Marchinton, 1969; Mooty, 1980)
are not appropriate, because they are based on
assumptions developed with information gen-
erated in temperate zones (Mandujano and
Gallina, 1995). Furthermore, it is difficult to
see deer in tropical forests due to limited visi-
bility and to frequent harassment by people.
Therefore, the track count might be the best
way to estimate density of deer in tropical for-
224 The Southwestern Naturalist
ests (Fritzen et al., 1995). Along dirt roads,
track counts correlate well with other indices
of population abundance (Mooty et al., 1984).
The track count is inexpensive and easy to ap-
ply on a large scale ( Jenkins and Marchinton,
1969) and can detect up to 20% of variations
in population size (Downing et al., 1965). Oth-
er researchers have correlated the number of
tracks with the amount of excrement (Mooty
et al., 1984), number of deer (Tyson, 1959;
Daniel and Frels, 1971; Bobek et al., 1986),
and paths made by deer (McCaffery, 1976).
The development of a density estimator using
track counts requires modeling the relation-
ship between the number of animals in the
area, spatial distribution, and the abundance
of tracks (Caughley, 1977). Tyson (1959) and
Daniel and Frels (1971) developed models that
convert track counts to population density.
In a tropical forest, Mandujano and Gallina
(1995) compared estimated densities of white-
tailed deer obtained with transect-line meth-
ods using the Fourier Model to those obtained
with track counts using the Tyson (1959) Mod-
el. The assumptions of the Tyson Model, which
was developed for temperate habitats where
deer move more widely than in tropical habi-
tats (Sánchez-Rojas et al., 1995), are not satis-
fied in tropical forest. Consequently, estimates
derived from the Tyson Model are biased as
much as 85% more than estimates based on
the Fourier Model (Mandujano and Gallina,
1995). Despite the drawback to the Tyson Mod-
el, it is an attractive method for estimating den-
sity due to the ease with which tracks, as op-
posed to the deer themselves, can be counted
in the study area. The purpose of this study is
to present 2 models for estimating density of
deer in tropical forests using the track index.
Both models were calibrated using the double-
sampling procedure. I considered the density
obtained through the line-transect method as
more accurate but also more costly in terms of
sampling effort, and track counts as less accu-
rate but easier to apply in the field. I also dis-
cuss the possible application of both models in
other regions.
METHODS Study Area This research was conduct-
ed at the Chamela Biology Station of the Universi-
dad Nacional Autónoma de México (UNAM), on
the coast of Jalisco, Mexico (198309N, 1058009W; Fig.
1). The Station covers 3,600 ha at an elevation of 30
vol. 50, no. 2
to 580 m. The mean annual temperature is 258C; the
warmest months are May through September (Bull-
ock, 1986). Mean annual rainfall for 1977 through
1997 was 748 mm (SD 5 119 mm). The rainy season
runs from July to October, when 80% of annual rain-
fall occurs. The dominant vegetation is dry tropical
forest, located on shallow-soiled slopes. The arboreal
stratum varies from 4 to 15 m and has a well-devel-
oped understory. Numerous species of trees and
shrubs lose leaves in the dry season (Lott et al.,
1987). Tropical semi-deciduous forest occurs in
deep soils of protected areas along major streams,
where the arboreal stratum ranges from 10 to 25 m.
Direct Deer Count Method From 5 to 8 permanent
transects were established on dirt roads (Fig. 1) and
varied from 6 to 11 km in length. I walked along the
roads (1 to 2 km/h) between 0700 and 1200 h and
1600 and 1900 h 2 or 3 times per month. All obser-
vations were grouped into 4 classes of perpendicular
distance: 0 to 10 m, 11 to 20 m, 21 to 30 m, and 31
to 40 m. I estimated monthly density (D, individu-
als/km2) as D 5 nf(0)/2L, where n is the number of
animals detected, f(0) is the probabilistic function
of density at a perpendicular distance of 0 m, and L
is total transect length (km). To estimate f(0) and
the standard error, I employed the DISTANCE Re-
lease 2 program (Thomas et al., 2003). I sampled
during the rainy seasons ( July to November) of 1989
and 1990 and during the dry seasons (December to
June) of 1990 and 1991.
Track Count Method I established 12 strip tran-
sects 500 3 0.90 m on dirt roads (Fig. 1). No tran-
sects were placed in heavy traffic or rocky areas, nor
where 2 roads were close together. One day before
the track count took place, I swept transects clean
of litter, removing loose dirt to erase old tracks and
to permit a clearer impression. After 20 to 24 h, I
observed the transects between 0700 and 1200 h,
repeating the procedure twice during each sampling
month. When I found tracks, I recorded the follow-
ing data: track length and width, distance or stride
between tracks, and the direction and number of
animals when .1 series of tracks were found near
each other. To calculate density, I used the Tyson
Model, modified to metric units: D 5 (n/((t 3 0.5)/
1.6))/(2.56), where D is deer density/km2, n the to-
tal number of tracks counted, and t the total number
of transects on the 500-m transect. I estimated pop-
ulation density for each sampling period. In the fol-
lowing 2 models, n/t was used as a track index (IT)
to derive a density estimate. Nevertheless, the meth-
od of estimating density varied with each model.
Model I This model is based on the equation ob-
tained from the simple linear regression model, us-
ing the track index as an independent variable and
the density obtained with the line-transect method
as the dependent variable. I paired data obtained
from the 4 sampling periods. The linear-regression
June 2005 Mandujano—Track count calibration estimate 225

FIG. 1 Location of study area, detailed map of roads used as transects for direct counting, and approx-
imate location of 500-m 3 0.90-m transects for track counting (the thick black line on road maps) for white-
tailed deer (Odocoileus virginianus) in Jalisco, Mexico. From http://www.ibiologia.unam.mx/ebchamela/
HIST2.html.
226 The Southwestern Naturalist
slope was tested using an analysis of variance to as-
certain the functional relationship between track in-
dex and density. I also estimated confidence inter-
vals for the regression coefficient.
Model II Finding fresh deer tracks on the road
implies that, at a particular time, I had the oppor-
tunity to see the animal leaving them. Such a sight-
ing could have been from a perpendicular distance
of 0 to 40 m. This strip includes the maximum width
at which I saw deer on line transects. Therefore, for
this model, assuming that I determine the average
number of times that an individual crossed the road
over 24 h, the track index can be transformed into
population density. This is the model proposed:
1 t 2132
n 2
D5 3 100
1 2
1,000 3 2 3 W
10,000
where n/t is the track index (IT), 2 to obtain the
number of tracks/kilometer, and 3 the number of
times that, on average, a deer crosses the road (Dan-
iel and Frels, 1971). The denominator is the surface
of the strip transect (2Lw, expressed in hectares),
where W is transect width. Thus, Equation 1 can be
resolved as follows:
333.5
D 5 IT 3
W el II resulted in significant differences between
To estimate density using the track index with this
model, it is necessary to define the appropriate W.
To do this, I substituted various widths (10, 20, 30,
and 40 m) into Equation 2 and used a one-way anal-
ysis of variance to compare the densities obtained.
To determine the appropriate width, I used paired
t-Student tests to compare the estimates obtained us-
ing each width against that obtained with the line-
transect method. The width at which there was no
statistical difference between Model II and the line
transect was substituted as the calibration factor in
Equation 2.
Finally, I applied a one-way analysis of variance to
determine differences in the density estimates ob-
tained from the 4 models (transect line, Tyson, Mod-
el I, and Model II) and used the Student-Newman-
Keuls Test for comparison of means (Zar, 1985).
RESULTS I counted 226 tracks on 82 km
(164 transects) and 177 deer on 418 km of line
transects. The track index was 1.32 6 0.25 (SD)
tracks/500 m, while the direct index was 0.43
6 0.07 deer/km. Both indices increased si-
multaneously, although the correlation was not
significant (r2 5 0.87, df 5 3, P 5 0.07). On
the contrary, there was a correlation between
track index (IT) and the density estimated with
vol. 50, no. 2
FIG. 2 Relationship between track index (track
number/500 m) and population density (deer/km2)
of white-tailed deer (Odocoileus virginianus) in Jalis-
co, Mexico, obtained with the line-transect method.
Each point on the graph represents a season (rainy
or dry) during the 2 years of the study period. Dot-
(1) ted lines represent confidence intervals to 95%.
line transects (Fig. 2; r2 5 0.90, F 5 18.2, df 5
1, 2, P 5 0.05). Thus, the regression equation
for Model I that predicted density on the basis
of track index was:
D 5 6.0 1 4.5 3 IT (3)
and the standard error of the slope was 1.41 (t
5 4.2, n 5 4, P 5 0.05).
The substitution of different widths in Mod-
(2)
density estimates (F 5 34.4, df 5 3, 15, P 5
0.0001). With an increase in transect width, es-
timated density decreased (Table 1, SNK test,
P , 0.05). Comparison of these estimates and
the density obtained with line transect showed
that when transect widths of 10 and 20 m were
used, significant differences resulted (t 5 27.5,
df 5 6, P 5 0.0003, and t 5 24.6, df 5 6, P 5
0.004, respectively); in contrast, use of 30-m
and 40-m widths produced no differences (t 5
21.8, df 5 6, P 5 0.12, and t 5 0.75, df 5 6, P
5 0.48, respectively). In consequence, when a
strip-transect width of 40 m was employed, the
density estimate was similar to that obtained
with the line-transect method. Thus, by substi-
tuting W 5 40 m in Equation 2, the expression
is reduced as follows:
D 5 8.338 3 IT (4)
Finally, the density obtained with the Tyson
Model produced the lowest estimate of all
models (Table 1, F 5 48.4, df 5 6, 27, P 5
0.00001).
DISCUSSION With the track-count method,
6.4 times more deer tracks were recorded per
June 2005 Mandujano—Track count calibration estimate 227

TABLE 1—Estimations of population density (D, deer/km2) of white-tailed deer (Odocoileus virginianus) in
dry tropical forest in Chamela, Mexico, obtained with the line-transect method (DTL), the track-count meth-
od using the Tyson Model (DTyson), Model I (DModel I) using linear regression, and Model II (DModel II) based
on the strip-transect method with varying widths (w). P values on the last line indicate the comparison of
density as estimated with each model against that estimated with the line-transect method.

Methods
DModel II*
Season/year DTL DTyson DModel I w 5 10 m w 5 20 m w 5 30 m w 5 40 m
Rainy/1989 11.3 1.4 11.9 40.9 20.5 13.6 10.2
Dry/1990 13.3 2.0 13.1 55.6 27.8 18.5 13.9
Rainy/1990 10.6 1.3 10.4 35.6 17.8 11.9 8.9
Dry/1991 12.5 1.7 12.6 44.1 22.1 14.7 11.0
Mean 12.0 1.6 12.0 44.1 22.1 14.7 11.0
SE 0.6 0.2 0.6 4.2 2.1 1.4 1.1
P ,0.0001 0.99 0.0003 0.004 0.12 0.48
* In Model II, different values of the transect width (w) were substituted in Equation 2.

km than the number of animals observed di-
rectly; moreover, the sampling effort was 5
times lower with track counting. As reported
by others (such as Daniel and Frels, 1971),
track counts had the advantages of minimally
disturbing the population and of obtaining a
relatively large sample size. Furthermore, the
number of tracks was correlated with the esti-
mated density obtained with the line-transect
method, as previously reported by Tyson
(1959), Daniel and Frels (1971), McCaffery
(1976), Mooty et al. (1984), and Bobek et al.
(1986). In addition to supplying data on the
number of animals, the track-count method
permits the detection of differences in track
characteristics that can indicate age group
(fawn or adult) and probably gender (Mc-
Cullough, 1965). Tracks can be monitored
during the rainy season, when soil conditions
yield clearer tracks. For these reasons, the
track count is an appealing and easily applied
method for estimating density of deer in dry
tropical forest.
The main limitation of the track-count meth-
od was that, in this habitat, dirt roads were not
located randomly. Human activity can cause
disturbance that keeps deer away from roads,
or, on the contrary, deer can be attracted to
roads as foraging sites during certain times of
year (Harlow and Downing, 1967; Sage et al.,
1983; Mooty et al., 1984). Ideally, transects
should be established randomly for track
counting, but in tropical forest, this is neither
simple nor economical due to abrupt topo-
graphic changes and dense vegetation. If hu-
mans cause little disturbance to deer, another
option might be to establish a certain transect
length, divide roads into equal segments, and
select randomly certain segments as transects,
as I did in this study. During the rainy season,
deer can be seen foraging alongside roads in
this region. However, preliminary data on deer
equipped with radiotransmitters (Sánchez-Ro-
jas et al., 1997) suggested that activities are not
near roads during all seasons. For this reason,
using roads as transects for track counts should
be limited as much as possible; it is only advis-
able to use forest paths when there is evidence
that doing so does not disturb deer.
Both models derived in this study (Equa-
tions 3 and 4) served to monitor populations
of white-tailed deer in the tropical forest of
Chamela. Extrapolation to other sites should,
however, be considered with caution. To ob-
tain the regression equation for other regions
with Model I, it would be necessary first to pair
track data with a density estimate obtained
with the line-transect method, and then to cal-
ibrate the track count using a robust regression
model. Model II (Equation 2) is more widely
applicable to other tropical dry forests and dif-
ferent habitat types because of the use of a
strip transect with a defined width. The density
estimate will depend on an adequate defini-
tion of transect width; width can be deter-
mined using the farthest perpendicular dis-
tance at which deer have been observed at the
site in question. Thus, by substituting this
228 The Southwestern Naturalist
width into Equation 2, the track count can be
calibrated to estimate density.
With the double-sampling procedure, 2
equations were obtained that resulted from
calibrating track counts on the basis of the
density obtained with the line-transect method.
Because the density obtained with the line-
transect method is not free from possible bias
(Mandujano and Gallina, 1995), the applica-
tion of any of the models proposed in this
study should be done cautiously and not ex-
trapolated to other regions indiscriminately. It
would be interesting to calibrate track counts
or fecal group counts, a method widely used
in Mexico to estimate the density of deer
(Ezcurra and Gallina, 1981), with estimates of
density obtained with more direct methods,
such as the identification of individuals
through DNA (e.g., Kohn et al., 1999; Taberlet
and Luikart, 1999). One alternative might be
to recalibrate the Tyson Model with accurate
data on home range size and daily distance
traveled by deer in tropical zones (Fritzen et
al., 1995). It is important to develop and im-
prove methods that accurately estimate popu-
lation size in tropical regions where deer com-
monly are hunted. The double-sampling pro-
cedure is appropriate; using 2 methods allows
calibration for extensive monitoring of animal
populations.
I thank L. E. Martinez-Romero, who aided me in
sampling, and J. M. Aranda for discussions about
ideas. I also thank S. Gallina and E. Naranjo for sug-
gestions about the manuscript. The Station of Biol-
ogy ‘‘Chamela’’ of the UNAM granted me all the
facilities to carry out the study. The National Coun-
cil of Science and Technology supported the project
financially.
LITERATURE CITED
ANDRÉN, H., J. D. C. LINNELL, O. LIBERG, P. AHLQVIST,
R. ANDERSEN, A. DANELL, R. FRANZÉN, T. KVAM, J.
ODDEN, AND P. SEGERSTRÖM. 2002. Estimating to-
tal lynx Lynx lynx population size from census of
family groups. Wildlife Biology 8:299–306.
BARNES, A., G. J. E. HILL, AND G. R. WILSON. 1986.
Correcting for incomplete sighting in aerial sur-
veys of kangaroos. Australian Wildlife Research
13:339–348.
BAYLISS, P., G. J. W. WEBB, P. WHITEHEAD, K. DEMPSEY,
AND A. SMITH. 1986. Estimating the abundance of
saltwater crocodiles, Crocodylus porosus Schneider,
in trial wetlands of the Northern Territory: a
mark-recapture experiment to correct spotlight
vol. 50, no. 2
counts to absolute numbers, and the calibration
of helicopter and spotlight counts. Australian
Wildlife Research 13:309–320.
BOBEK, B., K. PERZANOWSKI, AND J. ZIELIÑSKI. 1986.
Red deer population census in mountains: test-
ing of an alternative method. Acta Theriologica
31:423–431.
BULLOCK, S. H. 1986. Climate of Chamela, Jalisco,
and trends in the South Coastal Region of Mex-
ico. Archives Meteorological Geographic Biocli-
mate 36:297–316.
CARO, T. M. 1999. Conservation monitoring: esti-
mating mammal densities in woodland habitats.
Animal Conservation 2:305–315.
CAUGHLEY, G. 1974. Bias in aerial survey. Journal of
Wildlife Management 38:921–933.
CAUGHLEY, G. 1977. Analysis of vertebrate popula-
tions. John Wiley, London, United Kingdom.
CAUGHLEY, G., AND D. GRICE. 1982. A correction fac-
tor for counting emus from the air, and its ap-
plication to counts in western Australia. Austra-
lian Wildlife Research 9:253–259.
COULSON, G. M., AND J. A. RAINES. 1985. Methods for
small-scale surveys of gray kangaroo populations.
Australian Wildlife Research 12:119–125.
DANIEL, W. S., AND D. B. FRELS. 1971. A track-count
method for censusing white-tailed deer. Texas
Parks and Wildlife Department, La Porte.
DOWNING, R. L., W. H. MOORE, AND J. KIGHT. 1965.
Comparison of deer census techniques applied to
a known population in a Georgia enclosure.
Southeastern Game and Fish Communicating
Meeting: 26–30.
DUCKWORTH, J. W. 1998. The difficulty of estimating
population densities of nocturnal forest mam-
mals from transect count of animals. Journal of
Zoology 246:443–486.
EBERHARDT, L. L. 1978. Transect methods for popu-
lation studies. Journal of Wildlife Management
42:1–31.
EBERHARDT, L. L., AND M. A. SIMMONS. 1987. Calibra-
tion population indices by doubling sampling.
Journal of Wildlife Management 51:665–675.
ESCOS, J., AND C. L. ALADOS. 1988. Estimating moun-
tain ungulate density in Sierras de Cazorla y Se-
gura. Mammalia 52:425–428.
EZCURRA, E., AND S. GALLINA. 1981. Biology and pop-
ulation dynamics of white-tailed deer in north-
western Mexico. In: P. F. Ffolliot and S. Gallina,
editors. Deer biology, habitat requirements, and
management in western North America: a bina-
tional Mexico-United States Man and Biosphere
(MAB) Program Investigation. Instituto de Ecol-
ogı́a, A.C. México. Pages 77–108.
FIRCHOW, K. M., M. R. VAUGHAN, AND W. R. MYTTON.
1990. Comparison of aerial survey techniques for
pronghorns. Wildlife Society Bulletin 18:18–23.
FLOYD, T. J., L. D. MECH, AND M. E. NELSON. 1979.
An improved method of censusing deer in decid-
June 2005 Mandujano—Track count calibration estimate
uous-coniferous forest. Journal of Wildlife Man-
agement 43:258–261.
FRITZEN, D. E., R. F. LABISKY, D. E. EASTON, AND J. C.
KILGO. 1995. Nocturnal movements of white-
tailed deer: implications for refinement of track-
count surveys. Wildlife Society Bulletin 23:187–
193.
FULLER, T. K. 1991. Do pellet counts index white-
tailed deer numbers and population changes?
Journal of Wildlife Management 55:393–396.
GRAHAM, I. M. 2002. Estimating weasel Mustela nivalis
abundance from tunnel tracking indices at fluc-
tuating field vole Microtus agrestis density. Wildlife
Biology 8:279–288.
HARLOW, R. F., AND R. L. DOWNING. 1967. Evaluating
the deer track census method used in the South-
east. Proceedings of the 21st Annual Conference
of the Southeast Association of Game and Fish:
39–41.
HONE, J. 1986. Accuracy of the multiple regression
method for estimating population density in strip
transects. Australian Wildlife Research 13:121–
126.
HONE, J. 1988a. A test of the accuracy of line and
strip transect estimators in aerial survey. Austra-
lian Wildlife Research 15:493–497.
HONE, J. 1988b. Evaluation of methods for ground
survey of feral pigs and their sign. Acta Theriol-
ogica 33:451–465.
JENKINS, J. H., AND R. L. MARCHINTON. 1969. Prob-
lems in censusing the white-tailed deer. Proceed-
ings of the Symposium on White-tailed Deer in
Southern Forest Habitat: 115–118.
JETER, L. K. 1965. Census methods. In: R. F. Harlow
and F. K. Jones, editors. The white-tailed deer in
Florida. Florida Game and Fresh Water Fish
Commission, Technical Bulletin Number 9, Tal-
lahassee. Pages 186–192.
KOHN, M. H., E. C. YORK, D. A. KAMRADT, G. SAUVA-
JOT, AND R. K. WAYNE. 1999. Estimating popula-
tion size by genotyping faeces. Proceedings of the
Royal Society of London B 266:657–663.
LOTT, E. J., S. H. BULLOCK, AND J. SOLIS-MAGALLANES.
1987. Floristic diversity and structure of upland
and arroyo forests in coastal Jalisco. Biotropica
19:228–235.
MANDUJANO, S., AND S. GALLINA. 1995. Comparison
of deer censusing methods in tropical dry forest.
Wildlife Society Bulletin 23:180–186.
MCCAFFERY, K. R. 1976. Deer trail counts as an index
to population and habitat use. Journal Wildlife
Management 40:308–316.
MCCULLOUGH, D. R. 1965. Sex characteristics of
black-tailed deer hooves. Journal of Wildlife Man-
agement 29:210–212.
MENKES, G. E., JR., D. E. BIGGINS, AND S. H. ANDER-
SON. 1990. Visual counts as an index of white-
tailed prairie dog density. Wildlife Society Bulle-
tin 18:290–296.
229
MOOTY, J. J. 1980. Monitoring deer populations in
the northern forested areas of the Midwest. In:
R. L. Hire and S. Nehls, editors. White-tailed deer
population management in the north central
States. Proceedings of the 1979 Symposium of
the Northern Central Section Wildlife Society.
Pages 13–22.
MOOTY, J. J., P. D. KARNS, AND H. HEISYEG. 1984. The
relationship between white-tailed deer track
counts and pellet-group surveys. Journal of Wild-
life Management 48:275–279.
PARER, I., AND W. J PRICE. 1987. Sight counts as an
index of the number of rabbits, Oryctolagus cunic-
ulus. Australian Wildlife Research 14:569–570.
POLLOCK, K. H., AND W. L. KENDALL. 1987. Visibility
bias in aerial surveys: a review of estimation pro-
cedures. Journal of Wildlife Management 51:
502–510.
POTVIN, F., L. BRETON, AND L.-P. RIVEST. 2002. La
technique du double inventaire aèrien pour me-
surer l’abondance des popultions de cerf de vir-
ginie: ses fundaments scientifiques. Sociètè de la
Fauna et des Parcs du Québec, Canada.
SAGE, R. W., W. C. TIERSON, G. F. MATTFELD, AND D.
F. BEHREND. 1983. White-tailed deer visibility and
behavior along forest roads. Journal of Wildlife
Management 47:940–953.
SANCHEZ-ROJAS, G., S. GALLINA, AND S. MANDUJANO.
1997. Area de actividad y uso del habitat de dos
venados cola blanca (Odocoileus virginianus) en
un bosque tropical de la costa de Jalisco, México.
Acta Zoologica Mexicana (n.s.) 72:39–54.
SHORT, J., AND J. HONE. 1988. Calibrating aerial sur-
veys of kangaroos by comparison with drive
counts. Australian Wildlife Research 15:277–284.
TABERLET, P., AND G. LUIKART. 1999. Non-invasive ge-
netic sampling and individual identification. Bi-
ological Journal of the Linnean Society 68:41–55.
THOMAS, L., J. L. LAAKE, S. STRINDBERG, F. F. C.
MARQUES, S. T. BUCKLAND, D. L. BORCHERS, D. R.
ANDERSON, K. P. BURNHAM, S. L. HEDLEY, J. H.
POLLARD, AND J. R. B. BISHOP. 2003. Distance 4.1
Release 2. Research Unit for Wildlife Population
Assessment, University of St. Andrews, United
Kingdom. http://www.ruwpa.st-and.ac.uk/
distance/.
TYSON, E. L. 1959. A deer drive vs. track census.
Transnational North America Wildlife Confer-
ence 24:457–464.
WHITE, G. C., R. M. BARTMANN, L. H. CARPENTER, AND
R. A. GARROTT. 1989. Evaluation of aerial line
transects for estimating mule deer densities. Jour-
nal of Wildlife Management 53:625–635.
ZAR, J. H. 1984. Biostatistical analysis, second edi-
tion. Prentice Hall, Inc., Englewood Cliffs, New
Jersey.
Submitted 4 June 2003. Accepted 23 October 2004.
Associate Editor was Cheri A. Jones.