Evaluation of Museum Collection Data for Use in

Biodiversity Assessment
W. F. PONDER,* G. A. CARTER, P. FLEMONS, AND R. R. CHAPMAN
Centre for Biodiversity and Conservation Research, Australian Museum, 6 College Street, Sydney,
NSW 2010, Australia

Abstract: Natural-history collections in museums contain data critical to decisions in biodiversity conservation. Collectively, these specimen-based data describe the distributions of known taxa in time and space. As
the most comprehensive, reliable source of knowledge for most described species, these records are potentially
available to answer a wide range of conservation and research questions. Nevertheless, these data have shortcomings, notably geographic gaps, resulting mainly from the ad hoc nature of collecting effort. This problem
has been frequently cited but rarely addressed in a systematic manner. We have developed a methodology to
evaluate museum collection data, in particular the reliability of distributional data for narrow-range taxa.
We included only those taxa for which there were an appropriate number of records, expert verification of
identifications, and acceptable locality accuracy. First, we compared the available data for the taxon of interest to the background data, comprised of records for those organisms likely to be captured by the same
methods or by the same collectors as the taxon of interest. The adequacyof background sampling effort was
assessed through calculation of statistics describing the separation, density, and clustering of points, and
through generation of a sampling density contour surface. Geographical information systems (GIS) technology was then used to model predicted distributions of species based on abiotic (e.g., climatic and geological)
data. The robustness of these predicted distributions can be tested iteratively or by bootstrapping. Together,
these methods provide an objective means to assess the likelihood of the distributions obtained from museum
collection records representing true distributions. Potentially, they could be used to evaluate any point data
to be collated in species maps, biodiversity assessment, or similar applications requiring distributional information.
Anlisis del Uso de Datos de Colecciones de Museo en la Evaluacin de la Biodiversidad
Resumen: Las colecciones de historia natural en museos contienen datos crticos para la toma de decisiones
en la conservacin de la biodiversidad. Colectivamente estos datos basados en especimenes describen las distribuciones de taxa conocidos en tiempo y espacio. Como las ms confiables fuentes integrales de informacin para la mayora de las especies descritas, estos registros son potencialmente accesibles como una
solucin de costo-utilidad a un amplio rango de preguntas de conservacin e investigacin. No obstante, estos datos tienen deficiencias, brechas geogrficas notables, resultando principalmente de la naturaleza ad
hoc de los esfuerzos de colecta. Este problema ha sido citado frecuentemente pero raramente ha sido abordado de una manera sistemtica. Desarrollamos una metodologa para evaluar datos de colecciones de museo, en particular la confiabilidad de los datos de distribucin de taxones de rango corto. Solo se incluyeron
aquellos datos para los que existi un nmero adecuado de registro, verificaciones expertas de identificaciones y una aceptable precisin de la localidad. Los datos disponibles para los taxones de inters fueron
comparados con datos previos compuestos de registros de aquellos organismos propensos a ser capturados
por los mismos mtodos y por los mismos colectores de los taxones de inters. La adecuacin de los esfuerzos de muestreos previos fue evaluada mediante clculos estadsticos que describieran la separacin, la densidad y el anidado de los puntos y la generacin de un muestreo de un contorno de superficie de la densidad.
Se emple tecnologa de Sistemas de informacin Geogrfica (GIS) para modelar las distribuciones predichas
de las especies basndose en datos abiticos (climticos y geolgicos). La contundencia de estas distribu*email winstonp@austmus.gov.au
Paper submitted November 4, 1999; revised manuscript accepted July 19, 2000.

648
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Ponder et al.

Evaluating Museum Data

649

ciones predichas puede ser evaluada iterativamente o por el mtodo de bootstrapping. En conjunto, estos
mtodos proveen medios objetivos para evaluar la posibilidad de que las distribuciones obtenidas de registros de colecciones de museo representen las distribuciones verdaderas. Esto podra ser usado para evaluar
cualquier dato a ser cotejado en mapas de especies, evaluaciones de biodiversidad, o aplicaciones similares
que requieren informacin de distribucin.

Introduction
The value of museum natural-history collections for biodiversity research has been recently emphasized (e.g.,
Hoagland 1989; Hawksworth & Mound 1991; Nielsen &
West 1994; Davis 1996 and references therein). These
collections are essentially huge databases that have accumulated over long periods and that can thus provide a
historical perspective to complement contemporary field
surveys. In contrast to other forms of data, they are regularly accessed by specialists, and the material they contain is available for validation in the future, making them
essential to the integrity of biological knowledge (Cotterill 1995, 1997).
Museum collections may be particularly useful as a
source of data on invertebrates, which comprise around
99% of animal life but have been largely ignored in most
conservation and management programs. Although the
need to consider invertebrates has gained considerable
recognition in the last few years (e.g., Disney 1986; Majer 1987; Kremen et al. 1993; Heywood 1995; New
1998; Ponder & Lunney 1999), it is often considered
difficult because of the relative lack of published data,
perceived time and cost involved in wide-ranging invertebrate surveys, and lack of appropriate taxonomic expertise. Museum collections hold detailed distributional
data for many groups of invertebrates, and it is important that these data be evaluated and, where adequate,
utilized in conservation planning.
Problems with the use of collection data include lack
of access or availability, as well as inherent shortcomings (especially geographic gaps) in the data themselves.
Often the data are unpublished or the collections are not
electronically databased, and collections are usually arranged taxonomically, necessitating inefficient manual
retrieval of details for large suites of taxa. Shortcomings
of the data include the ad hoc nature of the collections,
presence-only data, biased sampling, and large collecting
gaps in time and space, although the extent of such gaps
depends on both the area and group under study. For instance, although the British biota has been extremely
well sampled (e.g., Prendergast et al. 1993a), a study on
Amazonian taxa (Kress et al. 1998) found that more than
a quarter of the primary grid cells lacked data. Some
taxa, such as the small flies discussed by Bickel (1999),
require specialized collecting and are less likely to be
represented in invertebrate collections than more con-

spicuous groups such as butterflies, larger beetles, large

land snails, or freshwater crayfish.
Collection data have been used in a number of recent
papers to map species distributions or determine areas
of conservation importance (e.g., Prendergast et al. 1993a;
Visnen & Helivaara 1994; Vane-Wright et al. 1994;
Drinkrow & Cherry 1995; Skelton et al. 1995; Williams
et al. 1996; Kress et al. 1998; McCarthy 1998). But there
have been relatively few attempts to address the shortcomings of collection data. The problem of undersampling during inventories was addressed by Colwell and
Coddington (1994), who used several techniques designed to give a measure of survey completeness. Nelson
et al. (1990) showed that supposed centers of endemism
were strongly correlated with collecting intensity, whereas
Prendergast et al. (1993b) and Fagan and Kareiva (1997)
introduced methods to correct for sampling effort in determining the number of species in a region.
The mapping of collection records for individual taxa
may allow some checking of the spatial accuracy of the
data (e.g., the position of apparent outliers), but it gives
little indication as to whether the collection records accurately reflect the true distribution of the taxon. We developed a methodology to determine the reliability of
distributional data, with a particular focus on seemingly
narrow-range taxa. Species represented by one or a few
records may be widespread but rare (only occasionally
found, despite searching effort) or simply infrequently
collected through lack of effort. The problem is therefore to distinguish between these taxa and those that
can be reliably judged to be of very restricted distribution. Our methods were developed as part of the Australian Museums Narrow Range Endemics Program (NREP)
(Ponder 1999), the aim of which is to identify areas important for the conservation of narrow-range, nonmarine invertebrate taxa (i.e., endemicity hotspots; Myers
1988, 1990). The approach is flexible and could potentially be used to evaluate any distributional data obtained
from museum or herbarium collections or other sources.

Methods
We evaluated the robustness of distributions mapped
from collection data in two stages. In step 1 we compared the available data for each species with the background data, an indication of background sampling ef-

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fort or sampling intensity. In step 2 we used geographic

information system (GIS) technology for predictive species modeling.
Data Selection
The data had to meet a number of criteria to be included
in the analysis. Taxonomic groups (genera, families, or
higher groups) were acceptable providing that they (1)
had been sampled at an appropriate intensity in the area
of interest and (2) contained one or more taxa of interest or were suitable as part of a background group for
a particular taxon of interest.
Taxon of interest (TI) refers to the particular species-group taxon (usually a species) whose distribution
is being analyzed. The choice of TI depends primarily on
the aim of the analysis: in the NREP, for example, the TIs
are all narrow-range taxa. The number of records (samples from a particular spot on a particular occasion)
available for each TI is also important, however. The estimation of distributions of taxa known only from a
small number of collection events can be equivocal, especially if the taxa are spread over a relatively large area
or if background sampling is sparse. We used a minimum of three records, but this could be increased to improve reliability.
The protocols we used for the inclusion of individual
records were as follows: (1) the same expert verifies
the identity of all records accepted for a particular TI;
unnamed taxa (e.g., morphospecies) were also acceptable and were coded; and (2) the locality accuracy is appropriate to the scale of the particular question being
addressed. In the NREP, all records are assigned a six-category accuracy code (10 m, 10100 m, 100 m1 km,
110 km, 10100 km, 100 km) and can be filtered accordingly. Records can also be sorted by date, so that,
for example, records older than a defined date can be
excluded if the reliability of the data is poor or if significant changes are known to have occurred as a result of
habitat modification. Similarly, records could be filtered
to take into account particular biological attributes of
the TI, such as host plant or season.
Background Groups
The background group for any particular TI is defined as
those organisms likely to be targeted by the same collector or caught by the same collecting methods as the TI.
Because records of background-group taxa are essentially used as a surrogate for searching or sampling effort
for the TI, the choice of appropriate taxa for inclusion in
the background group is important. They are not fixed
and may be readily redefined to test the effects of adding
or subtracting taxa or using filters to exclude inappropriate data (e.g., collections made during seasons in which
the TI does not occur). The background group is not

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necessarily strictly a taxonomic group because it is intended to reflect the likely biases of collectors and collecting methods, and it may contain several taxonomic
groups or parts thereof. For example, whereas some entomologists target mainly the group in which they have
an interest (e.g., butterflies), others may collect all those
insects that happen to be caught by the particular sampling methods employed (e.g., pitfall traps or foliage fogging). By contrast, land snails, which we used as an example, are generally collected as a group, either by eye
(larger species) or from litter samples (smaller species).
Because not all collectors employ both methods, we divided land snails into two independent background
groups (based on size, not phylogenetic relationship) for
analysis.
Background Sampling Analysis
Our basic premise is that if the background sampling effort is adequate, then the mapped distribution of a particular taxon may be presumed to accurately reflect its
true distribution. Survey sites where background sampling has taken place but the TI is absent can be used as
pseudo-absences (in lieu of true absence data) to define the distribution of the taxon. In contrast, if there is
little or no background sampling effort for an area, it is
unlikely that the distribution of the TI as described by
the available records will be accurate.
We used two approaches to assess the robustness of a
TIs distribution in terms of its background sampling effort, the first statistical and the second based on density
surfaces. We illustrate these approaches with a dataset
consisting of actual background data for land snails from
northern New South Wales, overlain with hypothetical
TI distributions based on commonly observed distributional patterns for narrow-range taxa.
SPATIAL STATISTICS

A set of statistics was generated that described the spatial distribution of geographic points for each set of data
(i.e., each TI and its background group). The program
used for calculating these statistics is available from the
second author.
These statistics describe the spatial distribution of
records and do not directly indicate the robustness of
the background sampling data. They can be used to indicate problems with the data, however, such as excessive
clustering, highly scattered distributions, or the presence of outliers (erroneously located collection sites)
problems that might require the data to be examined
further or discarded. This can be particularly useful in instances where there are a large number of taxa to be
screened, precluding individual visual examination.
Within-group statistics describe the spatial distribution and relationships between points in each dataset.

Ponder et al.

They include the maximum, minimum, median, and

mean separation (and standard deviations) of each point
from every other point in the group, and each point
from its nearest neighbor, as well as (1) the area occupied by the points; (2) the density of points; (3) the
mean nearest-neighbor separation for a uniform distribution of the same number of points in the same-size area;
(4) the ratio of the actual mean nearest-neighbor separation to that of a uniform distribution; and (5) the number of points separated by more than the mean nearestneighbor distance.
These statistics are calculated for the TI, for the entire
background group, and for those background-group
records that fall within a certain radius of the TI records
(a so-called buffered background group). The buffer
the size of which can be chosen by the user depending
on the question at handis used because, in many analyses, the total background-group area is inappropriate to
the question at hand. The buffer also serves to minimize
the effect of variation in sampling intensity in the background group by standardizing the scale of analysis. This
is necessary because sampling variation (clustering) increases with area, partly because sampling often occurs
opportunistically and on a localized scale, and partly because distributions are usually not random.
Between-group statistics were generated to describe
the distribution of each TIs points relative to those of its
background group. These include the maximum, minimum, median, and mean separation of each TI point
from every point in the background group, and of the
nearest background-group neighbor of each TI point.
Statistics were calculated with both the full backgroundgroup data and the buffered background data, as for the
within-group statistics.
Following analysis of these statistics, a TI may be excluded from further analysis, or tagged for checking if it
contains outliers, if the data are considered excessively
clustered, or if there appears to be inadequate background sampling based on the densities of points and
distances between TI and background-group records.
The thresholds for each of these parameters must be decided on a project or group basis, with consideration of
the data sampling regime.
DENSITY SURFACES

The next step involved the production of a contouredbackground sampling-density surface based on the background data for each TI. Nelson et al. (1990) used an
analogous contouring method to show the density of angiosperm collections in the Amazon based on records
per grid square. The density surface was created by removing the TIs records from the background group and
using the Arcview (Environmental Systems Research Institute, Redlands, California) density function to model
a surface for which the values are projected numbers of

Evaluating Museum Data

651

collection records per square kilometer. These values

are calculated by assigning a density to each grid-cell,
taking into account other records within a user-defined
distance (in our case 50 km). The density value is analogous to survey effort for the TIs background group.
The removal of the TIs records prior to creation of
the density surface is designed to generate a surface that
is independent of sampling efforts for the TI. The rationale for this is that inclusion of the TI records results in
misleading values for background sampling effort for
some taxa, such as those that have been heavily collected at a particular location but have few background
records in the vicinity. In the case of distinctly allopatric
taxa, however, the TI records may need to be included
because their removal will leave a hole in the sampling-density surface.
A single figure reflecting the background sampling intensity for each TI can then be calculated by averaging
the value of the density surface at each TI point. This
calculation provides an objective value for background
sampling that can be used to make modeling decisions.
For instance, TIs can be included or excluded from further analyses based on whether the background sampling density for their known range meets a user-defined
minimum level.

Predictive Modeling
Background data and the records for one or more TIs
can be overlaid with various environmental parameters
that may explain sampling gaps, such as extensive agricultural land, urban development, or a significant change
in climate, geomorphology or, geology. Simple correlations such as these can be undertaken relatively easily by
overlaying transparencies or using GIS technology for
computer mapping. More sophisticated analysis requires
computer modeling, as outlined in this section.
Computer modeling enables testing of the observed
distributions obtained from museum-collection data against
predicted distributions modeled from abiotic (e.g., climatic and geological) data. Modeling has been used to
predict the distributions of species (e.g., Nix 1986; Reid
1998; Regional Forest Agreement Steering Committee
1998), which can then be mapped by GIS. Species-distribution models can be designed for an optimum trade-off
between accuracy (e.g., small grid cells) and available
computational ability. The modeling algorithm we used
is based on the BIOCLIM approach (Nix 1986), which
uses climatic variables and GIS technology to map predicted distributions of TIs.
VARIABLES

Climatic variables for species-distribution modeling are

obtained from environmental surfaces generated by the

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ESOCLIM program (Nix 1986; Busby 1991). Basic meteorological surfaces (e.g., temperature and precipitation)
are used to derive a range of environmental surfaces
including variability and seasonality in rainfall, temperature, and radiationthat are more likely to reflect biological dependencies or critical limiting factors for the
species. Other important variables, which can be extrapolated from a digital elevation model (DEM), include topographic indices such as ruggedness, slope, and aspect. Other useful variables include soil fertility, depth,
and moisture, geology, and vegetation. These surfaces
are used in turn to define environmental profiles for TIs
based on available records. Many of these variables
particularly rainfall and temperature and their seasonalityhave already been widely used to predict species
distributions (e.g., Nix 1986; Austin & Meyers 1996; Ferrier & Watson 1997; Peters & Thackway 1998; Reid et al.
2001).
MODELING DISTRIBUTIONS OF TAXA

We modeled predicted distributions of TIs from collection-site data and environmental variables using an Arcview script. The script read the value of each bioclimatic
variable at each site, sorted and categorized them, and
ranked the values for each variable in ascending order to
create an environmental profile for the TI. Proportions
of the profile (confidence intervals) were used to exclude outlying values, depending on the level of confidence in the spatial accuracy of sites. Outliers may give
a misleading environmental profile and affect the geographical distribution predicted by the model. Commonly,
the 595% confidence interval is used to eliminate the
influence of outliers (e.g., Nix 1986; Reid et al. 2001), although this is an arbitrary setting. These intervals were
applied to the environmental surfaces to create a map of
the modeled or predicted TI distribution.
Distributions were further refined by adding other GIS
layers, such as geology and vegetation. For instance, a TI
may be associated with specific substrate or vegetation
types which, if known, allows elimination of predicted
distributions that fall outside these types. Predicted distributions should be verified by appropriate experts and,
where feasible, by field collecting.

Ponder et al.

& Watson 1997) delivers a measurement of the robustness of both the model and the raw data, ensuring that
the model can provide consistent results given changes
in the site information. Bootstrapping involves quantification of the response of the model to subsets of the
original data, the measurement being the percent overlap between models at successive levels of subsampling
(e.g., 100%, 95%, 90%, etc.), averaged over a number of
random samples at each level. Low levels of agreement
between the subset models indicate the presence of outliers or of poor sampling coverage.

Results
The distributions of the five hypothetical narrow-range
land snailsTIs a, b, c, d, and ewere analyzed based
on the same background group (actual land-snail data).
The relationship among these species and the background data reflect patterns commonly found in nature
(Fig. 1).
Background Sampling
The TIs a and d appear to have better levels of background sampling than either b or c, even though c has
more records than d (Fig.1). The TI e occurs in both
well-sampled and poorly sampled areas. Analysis of this
example, through calculation of spatial statistics and
generation of a background sampling-density surface, is
described below.

ROBUSTNESS OF MODELS

We assessed the robustness of the TIs distribution

model by evaluating the model either on an iterative basis or by bootstrapping. In iterative assessment, potential outliers are examined after each run, allowing their
identification and location to be checked against the
original collection data. Modeling can indicate incorrectly identified or databased records, which makes it
extremely useful for museum-collection data.
Bootstrapping (sampling without replacement; Ferrier

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Figure 1. Distributions of five hypothetical taxa of interest (TI, ae), with the background records (crosses)
based on actual data for land snails in northern New
South Wales.

Ponder et al.

Table 1.

Evaluating Museum Data

Sample of within-group statistics describing the spatial distribution and relationships between points in each data set.*

TI

Data set

No.
records

TI
50-km buffer
BG
TI
50-km buffer
BG
TI
50-m buffer
BG
TI
50-km buffer
BG
TI
50-km buffer
BG

8
89
317
3
34
322
6
68
319
3
64
322
3
148
322

b
c
d
e

653

MeanWithinGp
(km)

MaxWithinGp
(km)

MeanNearest
(km)

MaxNearest
(km)

Area
(km2)

Density
(km2)

UniformNearest
(km)

Actual/
Uniform
(%)

No. 
MeanNearest

5.6811
47.1323
99.6857
2.8112
50.8413
99.1236
6.0263
50.0836
99.1385
21.392
42.1471
98.5816
63.2289
75.8079
99.0628

12.2536
119.7919
235.9132
3.7971
99.3243
235.9132
13.7797
109.0267
235.9132
30.9614
103.4356
235.9132
80.6653
176.5659
235.9132

2.3796
0.1525
0.1048
1.8759
1.2992
0.1047
2.0977
0.2572
0.1054
15.0772
0.2604
0.1053
53.5686
0.1093
0.1047

4.4577
0.2927
1.7805
2.7100
4.5999
1.7805
8.8457
0.9089
1.7805
18.1605
2.6361
1.7805
57.3369
0.2927
1.7805

42
8634
34824
3
7502
34824
32
8510
34824
109
7392
34824
1724
19275
34824

0.1905
0.0103
0.0091
1.000
0.0045
0.0092
0.1875
0.008
0.0092
0.0275
0.0087
0.0092
0.0017
0.0077
0.0092

2.2913
9.8494
10.4812
1.000
14.8542
10.3995
2.3094
11.1869
10.4483
6.0277
10.7471
10.3995
23.9722
11.4121
10.3995

103.8521
1.5481
1.0002
187.5905
8.7461
1.0065
90.832
2.2988
1.0087
250.1314
2.4234
1.0124
223.4613
0.9577
1.0065

3
34
3
1
5
3
1
10
3
1
7
3
1
7
3

*Statistics calculated for each taxon of interest (TI, ae), its buffered background group (BG, all points within 50 km of the TI records), and
full BG. MeanWithinGp and MaxWithinGp, average and maximum distances between points in the group; MeanNearest and MaxNearest, average and maximum distances to the nearest neighbor of each point; UniformNearest, nearest-neighbor separation that would occur if the points
were distributed uniformly; Actual/Uniform, ratio of the actual nearest-neighbor separation (MeanNearest) to that of a uniform distribution
(UniformNearest); No.  MeanNearest, number of points separated by more than the mean nearest neighbor distance.

SPATIAL STATISTICS

Subsets of the within-group and between-group statistics were calculated for each TI and its background
group (Tables 1 & 2 respectively).
Within-group statistics can indicate the presence of outliers or clustering in the distribution of a TI or its background group. A small number of records (50%) with
more than the average nearest-neighbor separation (no. 
MeanNearest [average distance to nearest neighbor of
each point]) suggests the presence of one or more outliers. Close to half (3/8) of the records of TI a have a
greater than average nearest-neighbor separation, suggesting a fairly uniform distribution, whereas only one-sixth
of the records of c do so, indicating an outlier (Table 1).
For TIs with few records (e.g., b, d, and e), comparison of

MaxNearest (maximum distance to nearest neighbor of

each point) to MeanNearest may help flag outliers.
Clustering in a TI may indicate either outliers or a disjunct distribution. The degree of uniformity or clustering of records may be shown by the ratio of the actual
nearest-neighbor separation to that of a uniform distribution (Actual/Uniform statistic), with values of around
100 generally indicating a more uniform distribution and
lesser values indicating greater clustering. The high values for TIs b, d, and e are artefacts (each has only three
records), but TI c has a value of 90.8, indicating some
clustering. The value for TI a (103.9), however, suggests
that these points are distributed more uniformly.
For a TI, high density and low separation of points
(MeanNearest or the average [MeanWithinGp] and maxi-

Table 2. Sample of between-group statistics describing the distribution of each taxon of interests (TI) points relative to those of its
background group (BG).*
TI

Background data

TI records

BG records

MeanBtwnGp

SD(MeanBtwnGp)

MinBtwnGp

50-km buffer
BG
50-km buffer
BG
50-km buffer
BG
50-km buffer
BG
50-km buffer
BG

8
8
3
3
6
6
3
3
3
3

89
317
34
322
68
319
64
322
148
322

34.8486
80.5213
37.9954
77.0229
42.107
88.153
33.4761
105.961
64.7971
80.0879

19.8957
40.104
10.1238
21.592
9.8964
34.0319
18.4945
48.7315
31.0023
32.3276

0.912
0.912
7.4944
7.4944
14.594
14.594
1.6987
1.6987
1.905
1.905

b
c
d
e

*Comparison of each TI (ae) with its buffered and full BG: MeanBtwnGp, average distance between TI and BG points; SD(MeanBtwnGp), standard deviation of this; MinBtwnGp, minimum distance between a BG and a TI point.

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mum [MaxWithinGp] distances between points in the

group) indicate a well-sampled taxon, whereas low density and high separation suggest a more widely distributed rare taxon. The TI e falls into the latter category,
with density-to-separation ratios an order of magnitude
higher than those of the other TIs. Thus, it is probable
that e has not been sampled adequately over its range
and that the available data do not accurately reflect its
true distribution.
The density of buffered background points is highest
for TI a (0.0103/km2) and lowest for b (0.0045/km2)
(Table 1). Higher confidence in background sampling is
given by a high density and low separation (MeanWithinGp) of buffered background points. The buffered
backgrounds for TIs c, d, and a were better than those
of e and, particularly, b.
Based on the between-group statistics TIs a and d had
the lowest values for average distance (MeanBtwnGp)
and minimum distance (MinBtwnGp) between TI and
the background group, indicating that the background
records were closer to the records for these TIs (Table
2). The TI c and, to a lesser extent, b have higher values,
suggesting insufficient sampling in the immediate vicinity and low confidence in the accuracy of their distributions. The TI e has a high standard deviation, indicating
variability in the distances between its own and background-group records.
A comparison of the separation between TI and background-group points (e.g., MeanBtwnGp) with the density of buffered background-group points (Fig. 2) provides the best indication of confidence in the adequacy
of background sampling, and hence robustness of a TIs
mapped distribution. The TIs a and d have the highest
densities of buffered background points and the lowest
mean separation of these points from their own records,
reflecting the higher sampling intensities in the areas
surrounding these TIs.
The TIs a and d have the most reliable levels of background sampling (Fig. 2). The TI c has a possible outlier
(no.  MeanNearest  1/6), but the distances between its
records and those of its background group (Fig. 2) indicate a better level of sampling than that of TI b, which has
the poorest background sampling of any taxon. The level
of sampling around TI e is highly variable and is probably
inadequate to accurately determine its distribution.
DENSITY SURFACES

An example of the density of background sampling intensity can be shown with TI a removed (TIs be are included as part of the background) (Fig. 3). The single-figure values for background sampling intensity calculated
for each TI were a  0.027586, b  0.001623, c 
0.000477, d  0.012117, and e  0.010391. These values were calculated by averaging the value of the TIs
density surface (Fig.2) at each TI point.

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Ponder et al.

Figure 2. A plot of the mean distances between taxon

of interest (TI) and buffered background-group (BG)
points against the density of sampling (i.e., number of
data points per km2) for the buffered BG.

The TI a has the highest mean value for background

sampling (0.0276); TI d (0.0121) had the second-highest. Both of these are two orders of magnitude or more
higher than either b (0.0016) or c (0.0005), even though
c has the second-greatest number of records. Thus, the
levels of background sampling for b and c appear to be
too low to substantiate the distribution given by the
available points. The intermediate value obtained for e
(0.0103) reflects the fact that its records occur in both
well- and poorly sampled areas and emphasizes the need
to consider variation in sampling across a TIs distribution. These figures support the conclusion gained from
visual inspection of the data, but supply a quantitative
measure of background sampling intensity.

Predictive Modeling
After checking and (if appropriate) removing outliers,
testing of the 100% model at different bootstrapping levels has shown that repeated random samples of the
records can lead to high agreement in subset models
even with samples as small as 50% of the original points.
Using the methods described, for a well-sampled endangered species of land snail (Meridolum corneovirens;
Pfeiffer 1851) in Western Sydney (for distribution map;
see Little 1999), we found 98% agreement in the model
at the 50% level, and 81% with only 25% of the original
records. This indicates the robustness of the modeled
distribution, even when only one-quarter of the available
records are used.

Ponder et al.

Figure 3. Contour surface of background sampling

for taxon of interest (TI) a. Circle shows the 50-km radius (buffered background) around TI a.

Discussion
Museum and herbarium data have played an important
role in the biodiversity assessment and conservation of
many vertebrate groups and vascular plants, and it is emphasized in international programs such as the Global
Taxonomic Initiative (American Museum of Natural History 1999). For invertebrates, information resides mainly
in museum collections. More effective utilization of
these data will be enhanced by development of more
rigorous approaches.
Our methods differ from those of previous studies,
which have focused mainly on the mapping of grid-based
data, because we have assessed the robustness of distributions of taxa based on point data from museum collections. Somewhat analogous methods for describing
patterns in distribution density (Rossi et al. 1992), originally developed for mining, have been adapted for ecological purposes from geostatistics (Royle et al. 1980).
They differ in that they rely on standardized sample
spacing (lag distance) and thus on grid-based surveys in
which sampling effort and spatial arrangement are regular.
Our methods are not dependent on area and can be
tailored for use at different scales and in different environments, depending on data availability. The use of
background sampling analysis, combined with predictive modeling, provides a means of independently testing the robustness of the known distribution of a particular taxon. Both approaches identify areas that can be
field-tested: the first indicates gaps in the sampling and
the second indicates areas with similar environmental
profiles where the TI might be expected to occur. Field
testing of these latter areas should be prioritized accord-

Evaluating Museum Data

655

ing to low or absent background sampling or the presence of suitable or original habitat (e.g., forest cover).
Other potential applications of museum-collection data
for which our methods could be used include comparison of current with past distributions, assessment of distributions relative to current or proposed land-use options, and assessment of the relationship of distributions
to various environmental variables. In addition, the application of environmental models to museum distributional data could be extended to analyze potential changes
in species distributions caused by changes in temperature, rainfall, or vegetation.
Limitations of these methods include the fact that there
is reduced confidence in predictive power with small
numbers of records and when the data are highly clustered. Another is that the background-sampling method
will not be able to verify the distributions of isolated
allopatric taxa surrounded by large areas of habitat unsuitable for members of their background group(s). This
problem can be partially overcome through use of appropriate environmental overlays, in an a posteriori fashion, that exclude the unsuitable areas. The fundamental
premise of the background-sampling analysis (that if the
intensity of background sampling is adequate, then
the mapped distribution probably reflects the true distribution) involves some fairly major biological assumptionsthat seasonality, habitat fidelity, and annual population variation are understood. These assumptions cannot
always be made, especially for insects and other taxa
with high seasonality and host specificity. This problem
can be partly overcome through application of appropriate filters to test particular biological assumptions, such
as seasonality and host-plant preference. In the same way,
data can also be filtered to include or exclude particular
sampling methods.
Other limitations of these methods include the need
for (1) accurate species-level identifications for at least
the TI (accurate identification of other taxa in the background group is not essential); (2) accuracy of the specimen-locality data; (3) appropriate choice of taxa for
background groups, requiring a reasonable knowledge
of collectors, collection methods, and the biology of
component groups; and (4) availability of environmental
layers for the modeling assessment.
These taxonomic and data requirements may lead to
some difficulties in areas for which detailed environmental data, or the appropriate taxonomic expertise, are unavailable. Nevertheless, even if the outcomes may be
less reliable, the methodology can still indicate sampling
gaps and give indications of the reliability of distributions by means of the statistical procedures and/or density surfaces.
It is important that methodologies for evaluating museum data continue to be developed and enhanced but,
equally, that their value be more widely recognized and
that they be made more accessible through databasing

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Evaluating Museum Data

and improved data quality. The move toward online access to museum-collection data should eventually result
in a global biodiversity facility of immense importance.
The provision of adequate resources is essential, and taxonomic expertise and ongoing field work will be indispensable to improving and expanding these data.

Acknowledgments
We thank J. Kelly for developing the program to provide
the statistics. Many people, in particular the collection
managers at the Australian Museum, have contributed to
the Narrow Range Endemics Program within which the
methods outlined in this paper were developed. D. Faith
commented on an early draft of the manuscript, and C.
Reid made useful comments on a semifinal draft. We
also thank L. Wilkie for statistical comment.

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