Effects of sample size on home range delineation for woodland

caribou (Rangifer tarandus caribou) in Gaff Topsails, NL.

Jonathan W. Strickland
Dept. Environment and Conservation, Wildlife Division. Corner Brook, NL
FALL. 2oo9.

Abstract: To study the effects of sample size on home range delineation for the Gaff Topsails woodland caribou population,
fixed band kernel estimation was used. Kernel H-bands were calculated using mean step length of caribou. This method is
preferred over classical statistical methods such as least squares cross validation because it is an objective method that is
directly linked to the biology of the animal and is not affected at high sample sizes used. When studied at the herd level, a
minimum sample size (# of collars) was determined to adequately delineate home range area for the Gaff Topsails woodland
caribou population. When evaluated during calving, a minimum sample size could not be determined rather the relationship
between number of collars and home range area appeared linear. Spatial separation of females during calving was high. The
effects of collar temporal resolution on home range delineation were also studied. As temporal resolution decreased (becomes
coarse), home range area decreases. Home range fragmentation performs according to a polynomial function as temporal
resolution is decreased. Minimum concave polygons (MCCP’s) were used to delineate home ranges and were compared to
other home range methods. MCCP’s adequately describe the overall extend of home ranges at all temporal resolutions used in
this study but lacks information about home range interiors.

Introduction:
To date, wildlife telemetry studies have always been limited by one or more factors. The most common
of these limiting factors is budget associated with salary, field equipment, data analysis, and animal
collar fees. The purpose of this study is to determine the effects of sample size on woodland caribou
(Rangifer tarandus caribou) home range delineation. Information gathered about this relationship may
give us information regarding the minimum amount of data required to answer specific questions about
woodland caribou of insular Newfoundland. Information gathered from this analysis is intended to aid
Research/Management personnel with the development of new telemetry studies and/or modification
of existing studies provided they are similar in nature to the woodland caribou program investigated
here. Subsequent analysis will attempt to determine the following;

1. The effect of sample size (number of collars) on home range delineation for females of a
particular population/herd.
2. The effect of sample size (number of collars) on home range delineation for females of a
particular population/herd for a particular season (i.e. during calving season).
3. The effect of collar temporal resolution on home range delineation for an individual female
caribou.

Methods:
Data Collection:
The Department of Environment and Conservation, Wildlife Division, Corner Brook, NL has been
collecting female woodland caribou location data using GPS and Satellite (SAT) collars throughout the
island of Newfoundland from 2005 to present. Due to time constraints and the limited scope of this
project, all analysis was carried out on a sub-sampled dataset consisting of data from 15 GPS collars
from the Gaff Topsails population collected during 2007. This area was selected in order to complement
previously published and unpublished work completed in the area. GPS collar data was collected using
Lotek GPS4400 collars at a location frequency of 1 location per 2 hours, with a spatial error < 5 meters
(LOTEK Wireless). Data preprocessing included removing any locations collected when the collar was not
on the animal along with any locations having a Dilution of Precision (DOP) value > 5 (Adrados et al,
2002).

Home range delineation for females of a particular population/herd:

To study the effect of sample size on the home range area for females of a particular population/herd of
woodland caribou, fixed kernel density estimation was used. Kernels were created using Hawth’s tools
Version 3.27 (Beyer, 2002-2006), an extension for ArcGIS 9.3, for a range of sample sizes from n=15
collars to n=1 collar. Other methods used to delineate animal home ranges were not used for the
following reasons; adaptive band kernel estimation is known to overestimate home-range size (Seaman
and Powell 1996, Seaman et al. 1999, Powell 2000), Harmonic mean estimation was disregarded due to
similar issues (Naef-Daenzer 1993, Worton 1995). Minimum Convex Polygons (MCP’s) were also avoided
due to a host of known problems with this method (Harris et al., 1990, Horner and Powell 1990, Mills
2006, Powell 1987, Powell 2000, Stahlecker and Smith 1993, van Winkle 1975) the most severe being
large overestimations of home range sizes and lack of information about home range interiors.
Although fixed kernel density estimation was selected as the most appropriate estimator of Home
ranges for this study, the choice of an appropriate “smoothing factor” (i.e. bandwidth or H-band) is
much more important than the choice of a kernel (Worton 1989). There should be little difference
between the estimates of home range produced by different kernel functions (i.e. bivariate normal vs.
quadratic), compared to the differences caused by the choice of a smoothing factor (Klemelä 2009,
Wand and Jones 1995). Although objective methods exist for selecting bandwidth based on statistical
properties of the data, there is currently no objective method to tie band width to biology or location
error (Powell, 2000) ), except that bandwidth should be greater than location error (Silverman 1986).
Objective statistical methods (i.e. least squares cross validation) often produce poor results in wildlife
studies, especially at high sample sizes (Hemson, 2005).
Here I present a new objective method of bandwidth selection calculated using the ‘Animal Movements’
tool from Hawth’s tools Version 3.27 (Beyer, 2002-2006). Distance values were calculated for individual
animal step lengths. Mean step length was then calculated using all step length values contained in a
given file. Using data collected at 1 location/2hrs this gives a representative home range size/shape
(figure 1) however it is evident that adjustments would need to be made to this method for data
collected at low temporal resolutions which would result in high mean step length values causing over-
smoothing of associated kernels.
 Figure 1: Home range delineation for Gaff Topsails woodland caribou population, using fixed band
Kernel density estimation.

While calculating fixed kernels, 100 percent volume contour products were also created and then
converted to polygons in order to calculate total home range area. Home range areas were then plotted
against sample size (number of collars) in an attempt to detect a plateau in the data trend indicating the
minimum sample size required to delineate the home range of a particular population\herd of woodland
caribou for the study area.

Home range delineation for females of a particular population/herd for a particular season:
To study the effect of sample size on female caribou population/herd home ranges for a particular
season, home ranges were studied for the calving period of woodland caribou in Newfoundland.
Previous calf collaring programs carried out by the Newfoundland Dept. Of Environment and
Conservation, Wildlife Division, has determined peak calving to occur on or around June 03. Since most
calves of North American caribou are born in a brief 2-week period (Bergerud, 1975), this date was
buffered by one week on either end of peak calving, resulting in a two week calving period of May 27 –
June 10. Fixed kernel density estimation was used to calculate kernel home ranges for this time period
using h-band methods previously described. Home range area determined by 100 percent volume
contours was plotted against sample size (# GPS collars) in an attempt to determine the minimum
sample size required to adequately delineate the home range of a particular population/herd during
calving. Minimum sample sizes can be calculated in the same manner for all other aggregation periods
of woodland caribou including Pre-calving, Post-calving, Fly season, fall shuffle, Pre-Rut, Rut, Post-Rut,
Fall Migration, Winter, and Spring Migration (Bergerud, 1971) or more broad scale time periods derived
from combining several of the periods aforementioned.
In order to better understand the results of this analysis, a calculation of percent area overlap of
individual caribou during calving (for Gaff Topsails, NL, spring calving 2007) was completed. Percent
overlap was calculated by delineating kernel home ranges for each individual caribou for the date range
and study area previously stated. The ‘Intersect’ tool found in ESRI ArcGIS 9.3 was then used to develop
an ‘overlap matrix’ of home ranges, which was then used to create a frequency histogram of percentage
overlap.

Home range delineation for an individual female caribou:

To study the effects of collar temporal resolution on home range delineation for an individual female
caribou, one animal was randomly chosen from the Gaff Topsails dataset to be used in the analysis. 35
new datasets were then created from this file, each decreasing the temporal resolution from the original
1 location/2hours to a minimum of 1 location/100hours (approx. 1location/4days), where low (or
coarse) temporal resolution relates to longer time periods between successive locations. Temporal
resolution was decreased by deleting every second record for file 2, every third for file 3, etc. The
coarsest temporal resolution investigated (approximately 1 location/ 4 days) is significant as it is the
temporal resolution of a large number of SAT collars currently active in Newfoundland. Home ranges
were delineated for all 36 files using fixed kernel density estimation. 100 percent volume contour areas
were then calculated and plotted against temporal resolution in an attempt to detect a plateau in the
graph indicating the coarsest temporal resolution useful for adequate delineation of home range areas.
H-band was calculated as the average step length for the original file; however the same H-band was
used for all calculations in this particular analysis since an increase in spatiotemporal resolution causes
an increase in mean step length. Poor H-band selection causes a false relationship between home range
area and temporal resolution, where home range area increases as temporal resolution becomes
coarser, therefore some type of correction factor would need to be applied to H-band calculations in
order to compare home range area size at different spatiotemporal resolutions.
To better understand the results, a calculation of home range fragmentation was then completed to
determine the effect of collar temporal resolution on area fragmentation within the boundaries of an
individual’s home range. MCP’s were then used to study the effect of location frequency on home range
perimeter. Due to previously mentioned issues with MCP’s however, home ranges were also delineated
using a concave hull method (Minimum concave polygons, MCCP’s). ESRI’s ArcGIS model builder was
used here to create a tool capable of calculating MCCP’s from a set of input points, calculating the home
range area, and outputting the results (figure 2). To make best use of this tool, a scratch folder should be
created in the tools ‘Environments’ section to ensure all intermediate files can be stored in a pre-
defined directory other than the source file folder. Intermediate files may be used for future analysis,
data verification, or be deleted once the model is finished running.
Figure 2: model tool creation designed to delineate woodland caribou home ranges using a concave hull
calculation. Calculations in the red box are those completed using typeconvert 2.4.0 for ArcGIS.
Calculations in the blue box are completed using ArcToolbox. ‘P’ refers to a ‘model parameter’.

Results:

Home range delineation for females of a particular population/herd:

Upon investigation of the effects of sample size (number of collars) on home range area, it was
determined that as sample size decreases, home range area also decreases. A plateau in home range
area was detected at a sample size of approximately 15 collars, indicating a minimum of 15 GPS collared
individuals with a temporal resolution of 1 location/2 hours is required to adequately delineate the
home range of the Gaff Topsails population of woodland caribou (figure 3). Adequate home range
delineation may also be obtained by using multiple Satellite (SAT) collared individuals, or a mixture of
both GPS and SAT collars, however minimum sample sizes would need to be calculated according to
collar types used, the temporal resolution of the collar, and the area in which the collars are to be
deployed. Minimum sample sizes can be determined for all cases using the methods previously
described. Minimum temporal resolution requirements may exist however and will later be discussed in
greater detail.
Figure 3: Kernel home range area versus sample size (# collars) for woodland caribou in Gaff Topsails, NL, 2007.

Home range delineation for females of a particular population/herd for a particular season:
When studied on seasonal bases, in this case during calving (May 27-June 10), results indicate that as
sample size increases, home range area also increases (figure 4). A data plateau was not detected in this
analysis however, rather this relationship appears to be linear (y = 34.565x), meaning that although a
sample size of 15 collars was adequate to delineate the home range for this population of caribou, it is
not adequate to delineate the populations overall area utilized for calving (i.e. calving grounds).
Figure 4: Home range Area (km²) versus sample size (# GPS collars) for Gaff Topsails, NL, during calving (May 27 –
June 10, 2007).

Further visual inspection of points indicated there was some degree of spatial separation among females
during calving (figure 5) representing a possible explanation for the fact that a minimum sample size
does not appear to have been attained to determine population level ‘calving grounds’. Upon
completion of an overlap matrix of kernel home ranges for the area (Table 1), a mean percent overlap
was calculated to be 15.6% with a maximum percent overlap of 50.1%, indicating extremely low
amounts of space co-existence between females during calving (figure 6). With results showing a mean
calving area to be 66.08 km², it is evident that a more accurate analysis would likely determine this value
to be an over-estimation of co-existence.
Figure 5: Female woodland caribou GPS telemetry locations, Gaff Topsails, NL, during calving (may 27 – June 10,
2007).

Table 1: Overlap matrix showing area of overlap (km²) between home range areas of female woodland caribou
during calving, Gaff Topsails, May 27 – June 10 2007
Figure 6: Frequency histogram of percent overlap in home ranges of individual female caribou in Gaff Topsails, NL
during calving (May 27 – June 10).

Home range delineation for an individual female caribou:

Kernel home range area decreased as the temporal resolution of a GPS collar was coarsened to the point
at which it simulated a SAT collar, (figure 7). As kernel home range area decreased, home range
fragmentation increased to a maximum at 1 location/ 69.89 hours (approximately 1 location/ 3 days)
and then decreased according to the polynomial function y = -0.008x² + 1.15x +6.21, R²= 0.894, SE=
4.098 (figure 7). Home range areas were also calculated for all files using minimum concave polygons
(MCCP’s) (figure 8). Results showed that home ranges based on MCCP’s were comparable to those
created using kernel methods but were not affected by a reduction in the temporal resolution of caribou
locations. Unlike kernels however, MCCP’s did not contain information about habitat usage and
avoidance within a given range (figure 9). Using MCCP’s, mean home range size for an individual caribou
in Gaff Topsails, NL in 2007 = 1517 km².
Figure 7: Effects of spatiotemporal scale on home range fragmentation and size, for woodland caribou, Gaff
Topsails, NL. 2007.

A B C

Figure 8: Home range delineation for an individual woodland caribou, located in Gaff Topsails, NL. Using A: fixed
band kernel estimation, B: Minimum Convex Polygon (MCP), and C: Concave Hull.
Figure 9: Effects of temporal resolution of GPS telemetry collars on woodland caribou home range size using
MCP’s, concave hulls (MCCP’s), and kernels. Gaff Topsails, NL. 2007.

Discussion:

Wildlife telemetry studies have become common and relatively easy to conduct over the last number of
years. Since few projects have unlimited resources however, important considerations must be made
such as the effects of sample size on information gathered using telemetry data. Here we investigate 15
GPS radio collars from the Gaff Topsails woodland caribou population to analyze the effects of sample
size on home range delineation of a particular population/herd, or for a particular population/herd
during a particular season (i.e. calving). The effect of temporal resolution (of a telemetry collar) on home
range delineation was also studied.

Home range delineation for females of a particular population/herd:

Using fixed kernel density estimation, it was determined that as sample size decreases, home range area
also decreases. A data plateau was observed at n=15 collars, meaning approximately 15 GPS telemetry
collars, with a temporal resolution of 1 location per 2 hours is the minimum sample size required for
Gaff Topsails, NL to adequately delineate home range area for 2007. Minimum sample size depends on
the symmetry, compactness, and degree of spatial overlap in an area and therefore varies.
Considerations must also be given to the possibility of programming collars to have alternative temporal
scales, to use other collar types (i.e. satellite collars), or to use a combination of multiple collar types
(i.e. SAT and GPS collars currently being used throughout most of the island of Newfoundland).
Minimum sample size requirements would vary for all cases mentioned and would have to be re-
calculated on an individual study/area bases.
Fixed kernel bandwidth was calculated in each case by determining the mean step length for caribou
included in that file, meaning the mean distance an animal travels between two consecutive data
recordings. This method was determined to be optimum for this dataset due to very large sample sizes
and the undying need for an objective method of determination that relates to the biology of the animal
rather than classical statistics (Powell, 2000). Kernel bandwidths calculated in this manner ensure a
‘tight kernel’ which can sufficiently detect holes in the animal’s home range. These holes should be
considered extremely important biologically, and should be investigated to see what animals are not
using these regions.

Home range delineation for females of a particular population/herd for a particular season:
A study of the effects of sample size on home range delineation during calving (May 27 – June 10)
indicated that as sample size increased, home range area also increased in a linear fashion (i.e. no data
plateau). This implies that there is no way to measure home range area during calving using a sample of
animals (i.e. every female would have to be collared). The more likely explanation for these results is
that we have not reached a data plateau with the sample size we have even though a plateau may exist
somewhere between 15 < n < total population of females.

An investigation of the degree of spatial overlap between individual females at this time further explains
our results. With a mean percent overlap of 15.6% and maximum of 50.1%, we understand that females
are not aggregating in traditional ‘calving grounds’ but are rather separated into individual calving sites.
This behavioral shift is further supported by a potential underestimation of spatial overlap mainly
caused by the use of a generalized two week calving period (May 27 – June 10). Since individuals calf at
different times throughout this time frame, possible pre and post calving migration locations may be
included in the analysis inflating home range area. Although all collars are on female caribou at this
time, it is also possible locations may be acquired from a barren doe (female without calf), causing
further bias of results.
Larger sample sizes may be required to adequately delineate home ranges for specific periods (seasons)
if spatial separation of individuals is high and should be considered in other projects.

Home range delineation for an individual female caribou:

Upon studying the effects of the temporal resolution of a collar on home range delineation, it was
determined that as temporal resolution decreases (i.e. becomes coarse), kernel home range area
decreases, and kernel home range fragmentation increases to a temporal resolution of 1 location/3 days
and then decreases according to a polynomial function. A data plateau was not detected in this analysis
to be able to comment on the minimum temporal resolution required to adequately delineate home
ranges for woodland caribou in Gaff Topsails, NL using kernel methods. Results indicate a temporal scale
< 1 location/2hours may be adequate and should be tested in a field setting in order to confirm. By
calculating the degree of kernel home range fragmentation at varying temporal resolutions, it was
observed that home range interiors are affected more by temporal resolution than the overall home
range extent (defined by the perimeter) for woodland caribou. Since MCCP’s give information about the
overall extent of a home range, without incorporating large areas, devoid of fixes, as can occur at the
extremities of convex polygons (Harris et al., 1990), MCCP’s were calculated for all files and compared
with other home range methods. MCCP’s may be extremely useful for home range delineation when
temporal resolution is low, giving low sample size of points, when internal home range composition is
not important (i.e. select conservation applications), or when used in some combination with kernel
home ranges at large sample sizes (i.e. large number of locations).

Conclusion:
Home range delineation is greatly affected by sample size. The minimum sample size required to
adequately delineate home range areas for Gaff Topsails, NL, 2007 is approximately 15 GPS collars with
a temporal scale of 1 location/2hours. To delineate home ranges during calving (i.e. calving grounds), 15
collars is inadequate due to a low amount of spatial overlap between individuals. Home range
delineation is also affected by the temporal resolution of a collar. As temporal resolution decreases
(becomes coarse), home range area decreases. A minimum temporal resolution requirement was not
found in this analysis, and may be more frequent than 1 location/2hours. Home range fragmentation
increases to a temporal resolution of 1 location/3 days of and then decreases according to a polynomial
function. Minimum concave polygons are useful for delineating home ranges at low temporal
resolutions when information about home range interior is not required.
It is recommended that minimum data requirements proposed in this study should not be used to limit
projects that have the potential for more data collection. Larger datasets may leave possibilities open
for future research requiring larger sample sizes. Proposed sample sizes should be considered when an
individual/group wishes to answer specific research questions with the lowest possible budget, or would
like to prioritize research objectives in order to diversify research goals or optimize results.
It is recommended that the information gathered in this report be used as guidelines for a much larger
project considering all woodland caribou telemetry data gathered in Newfoundland from 2005 to
present. Further questions may also be answered including the minimum amount of time a collar must
remain on an individual to get its true home range size (i.e. number of years), The effects of using
multiple collar types together (i.e. GPS and SAT) on home range delineation, and the variation in data
requirements amongst island caribou populations based on population compactness, symmetry, and
spatial overlap of individuals.

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