David R.B.

Stockwell

Niche Modelling

CRC PRESS
Boca Raton London New York Washington, D.C.
Chapter 1
Chapter

A bias is generally understood as a prejudice in the sense for having a predilec-

tion to one particular point of view or ideological perspective. Statistical bias
is a quantifiable tendency of a statistical estimator to over or underestimate
the quantity that is being estimated.
In formal terms, if θ̂ is an estimator of a parameter θ, then θ̂ is biased if
the expected value of θ, E(θ̂), is not equal to θ.
Bias is a concern when any process is supposed to be objective and fair.
Similarly statistical bias of an estimator should be of concern, as that bias
can influence results and undermine the objectivity of conclusions.
For example, systematic errors increase the likelihood of assigning statisti-
cal significance to chance events (Type I error). Given an hypothesis

H0 : θ̂ = θ

and the alternative

Ha : θ̂ 6= θ

then bias would tend to increase the chance of rejecting H0 when H0 is

true, or committing a Type I error.
Bias is more insidious than uncertainty introduced by small samples. For
example, errors in estimation of a mean value are generally symmetric, that
is, equally as likely to be above or below the actual value. Thus while a small
sample introduces uncertainty, it does not necessarily bias the result.
The detection of bias in modeling should be a primary concern. While in-
creasing the sample size can reduce uncertainty in symmetric errors, averaging
is not a solution to asymmetric errors. The variation in symmetric sampling
errors reduces our power to detect statistical patterns in data (Type II error).
Type II errors only lead us to reserve judgment, but Type I errors cause us
to draw incorrect conclusions. In most cases, drawing incorrect conclusions
should be a greater concern than failing to find significance.

3
4 Niche Modelling

1.1 Range shift

We look at potential bias in the range-shift methodology. This methodology
examines effects of potential shifts in the ranges of species that may be brought
on by changes in the environment.
In order to better understand the response of species to environmental
change, and potentially predict responses, an ecological niche model is devel-
oped based on present day environmental variables and then reapplied using
the different variables, either from the past or future. This often leads to a
‘shift’ in the predicted distribution of the species. Shift-modeling is the name
for comparing predicted ranges of species in the past and future with the
present.

1.1.1 Example: climate change

The main applications of shift-modeling to date have been the reconstruc-
tion of past distributions of species under paleo-climatic conditions, and the
possible effects climate change on biological communities.
Climate is always changing. Since the last major extinction at the start
of the Pleistocene ice-age at 700kyr BP, the earth’s climate system has oscil-
lated between glacial and interglacial states approximately every 200,000 years
(PJR+ 99). Temperatures in Antarctica varied by about 10◦ C from peak to
trough, and transitions of at least 2◦ C occur on average every 10,000 years
(AMS+ 97; AMN+ 03; MGLM+ 04).
Local climate changes can be even more abrupt. The Younger Dryas event
at about 12kyr BP changed local temperatures 7◦ C in 1000 years, and events
at 8.2kyr BP cooled Europe even more abruptly. In response to these changes,
the fossil record shows changes to faunal composition and richness with a few
recent extinctions attributable to recent climate changes (JW99). There is
some observational evidence that ranges of some fauna and flora have changed
in response to the increase of 0.6◦ C in the last century and will continue to
change in response to climate changes (PY03; RPH+ 03; TWC+ 04).

1.2 Range-shift Model

Here we develop a simple geometric model to estimate the asymmetric error
in simple range-shift niche models. The particular statistic of concern is the
change in the potential range area available to species. Note the range-shift
methodology does not explicitly represent other factors such as:
 Chapter 5

• dispersal ability,

• rate of change, or

• threshold effects.

Variation in dispersal ability of individual species in response to change

is modeled by two geometric combinations of ranges: new ranges represent-
ing free dispersal and the intersection of new and old areas representing no
dispersal. These alternatives are intended to ‘bracket’ the possible range of
behaviors.
Simple shift models incorporate rate of change in temperature only by way
of dispersal and no-dispersal scenarios. They contain no history effects that
could incorporate a threshold effect, and potential novel combinations of cli-
mate.
As bias is the tendency for a statistic to over or underestimate the true
value, bias is quantified by showing a particular statistic does not estimate the
correct value, by comparing the true value with the estimated value. Deviation
in the expected value of the new or intersection area due to errors quantifies
the bias.
A theoretical model including errors in the parameters allows calculation
of both the true area, and the change in area resulting from errors in those
parameters. Range area predictions are regarded as biased if the observed
value of range area Ae does not equal the expected value Ae . Thus area can
be treated as a statistical parameter like accuracy (SP02a).
Consider a shape such as a circle with width, or radius r. Shifting the shape
laterally by s produces areas:

• old AO ,

• new AN ,

• an intersection area AI , and

• a union AU .

These areas can be calculated for squares and circles using the usual for-
mulas. In this case the area of the square is AO = (2r)2 . The formula for
intersection areas of a circle for shift s when radius r remains a constant value
of 1 is:

AI = 2r2 cos − 1(s2 /2sr)

The equation for intersection of a square is somewhat simpler:

AI = 2r(2r − s)
6 Niche Modelling

O N

FIGURE 1.1: Theoretical model of shift in species distribution from change

in climate. Dashed circle marked O is old range, solid circle marked N is new
range and I is intersection area.

Using the formulation of the square leads to simpler calculations. Fig-

ure 1.2 shows the intersection areas for squares and circles at different shifts
for squares (solid line) and for circles (dashed line). The intersection area of
a square is only slightly greater than the intersection area of the circle. We
will consider only the square in order to simplify calculations.
We now incorporate three error terms into the equations:

• error in the shift term se ,

• error in width term re , and

• error in the proportional change in area AN /AO or pe .

The new area is given by:

AN = (2rre )2 pe

The intersection area is given by:

AI = 2rre (2rre − sse )pe

 4 Chapter 7

r=1
3

r=0.9
2

r=0.8
1
0

0.0 0.5 1.0 1.5 2.0

Shift s

FIGURE 1.2: The change in the areas of intersection of a square and circle
for different shifts (s) and widths (r).
8 Niche Modelling

●
s

FIGURE 1.3: Combined effect of shift and width error.

1.3 Forms of bias

The model quantifies the reduction in intersection area simply as a function
of the geometric considerations. A number of consequences are immediately
apparent:

• With no shift or width error, shift alone will not reduce ranges.

• For a given shift, species with smaller ranges will have greater reduction
in intersection areas that species with larger ranges.

• Due to edge effects, proportional areal reduction depends on the size of

shifts relative to the size of the study region.

• Apparent new ranges are considerably reduced by width and shift errors
that reduce new widths and increase apparent shifts.

We now look at motivations for size and direction of the individual error
terms.
 Chapter 9

1.3.1 Width r and width error

Generally it is assumed that width as determined by the model of occur-
rences is a good indication of the actual range of the species. However model
determined width may be subject to errors. Researchers have noted the prob-
lem of recovering accurate response curves from incomplete geographic ranges
(AM96; TBAL04)). Because the samples of observations are incomplete,
all empirical models underestimate width of the potential niche, or ‘overfit’
(SP02b). Thus there will inevitably be areas of potential habitat outside the
known observations.
The consequence of overfitting is prediction of a smaller area than would
otherwise be predicted. Overfitting has been shown to be a potent source of
bias in climate envelope methods due to the truncation of the range of vari-
ables as discussed previously, underestimates of areas by up to 15% (BHP05).
Other studies have shown similar errors in range due to sample size. Where
overfitting is defined as the difference between accuracy of a model on a train-
ing set versus a test set, errors range from 30% for 10 data points to around
5% at 100 points when datasets are sufficiently large (SP02b). Studies indi-
cate overfitting also varies by method. In one study of a large number of bird
species, overfitting of species with twenty observations was approximately 10%
with GARP and 15% for logistic regression (SP02b). Figure 1.3 illustrates the
combined effect of shifting the square range (dotted) and width error (solid).

1.3.2 Shift s and shift error

A major source of shift error is differential response of species to climate
change. It is generally assumed that the only two possible responses to cli-
mate change are migration to new ranges or contraction of the range to the
intersection of the old and the new area.
If, however, some species maintain old ranges or expand to form a union
of old and new ranges, average areal change is overestimated. This could
come about if climate sensitivity was orthogonal to dispersiveness. Some
physiological studies show climate based ecological niche models exaggerate
the effects of climate, ignoring complex responses of species to other factors
(Ll96; SCF+ 01; Loe03). Differential (non-climatically driven) behavior in the
leading and trailing edges of ranges is supported by observational evidence
(PY03; RPH+ 03; TTR+ 04). For example, many temperate trees can survive
outside their current range as sub-dominants even though dominated by faster
growing sub-tropicals (Loe03).

1.3.3 Proportional pe
Map boundaries are a potential source of width bias in range-shift modeling.
As climate patterns shift, a certain percentage of distributions will be moved
out of study area, reducing their apparent size. Compensation cannot occur
10 Niche Modelling

●
s

FIGURE 1.4: Combined effect of shift and shift error

 Chapter 11

●
s

FIGURE 1.5: Combined effect of shift, shift error, width error and pro-
portional error.

elsewhere in the study area because: (1) the ranges of species entirely outside
the old area are not represented, and (2) truncated distributions are fit to
species at edges. This is an artifactual effect introduced because species’
ranges cover the landscape beyond the limited study area.

1.4 Quantifying bias

Figure 1.5 shows the effect of new range from all errors. We illustrate the
effects of these errors in the parameters by substituting the following values
into the theoretical shift model.

• unit radius r = 1,

• radius error re = 0.85,

• shift s = 0.8,
12 Niche Modelling

• shift error se = 1.33 and

• proportional change error pe = 0.85

Substitution of these parameters into the theoretical model gives the fol-
lowing results.

• With no bias errors the new area is an unchanged 4 and the intersection
area is 2.4.
• All errors combined reduce new area to 2.46 and intersection area to
0.96.
• The theoretical model suggests errors from biases reduce the size of new
areas to 62% and the size of intersections areas to 40% of their expected
values.

1.5 Summary
The theoretical range-shift model estimates bias in the range-shift method.
The bias results in exaggerated estimates of reduction in range, and potentially
exaggeration of risk to species from climate change. Strong and unambiguous
conclusions are not justified when there is significant biases, and can be a
considerable source of errors.
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