Gottschalk - Et - Al 2005 Modelling Bird by RS and GIS

International Journal of Remote Sensing
Vol. 26, No. 12, 20 June 2005, 26312656
Review article
Thirty years of analysing and modelling avian habitat relationships using

satellite imagery data: a review
T. K. GOTTSCHALK*{, F. HUETTMANN{ and M. EHLERS

{Justus-Liebig-University Giessen, Research Centre for Bio Systems, Land Use and
Nutrition, Department of Animal Ecology, Heinrich-Buff-Ring 2632, 35398 Giessen,
Germany
{University of Alaska Fairbanks, EWAHALE Laboratory, Biology and Wildlife
Department, Institute of Arctic Biology, Alaska 99775-7000, USA
University of Vechta, Institute for Environmental Sciences, P.O. Box 1553, 49364
Vechta, Germany
(Received 25 July 2003; in final form 13 October 2004 )
The application of remote sensing and Geographic Information Systems (GIS)

technologies provides powerful tools when used to investigate wildlife and its
habitat for an analysis or modelling approach. In this context, birds have been of
great and progressive value as biological and environmental indicators. In order
to learn about the common approaches usedits methods, processing steps,
trends, advantages and challengesover 120 representative publications of the
last 30 years that made use of satellite images for avian applications have been
reviewed.
The reviewed studies have shown that GIS-based analyses of satellite and bird
data have been well established for efficient ecosystem descriptions and species
modelling within a large range of scales and habitats. In order to improve the
quality of inference and for comparative analyses, it is recommended that further
studies are documented in detail. Also, in order to verify and improve the
obtained results, additional ground data on the main structure of the vegetation
relevant to the bird species in question are usually necessary. Satellite-based
remote sensing applications in ornithology could be used increasingly for
assisting in habitat evaluation, habitat modelling and monitoring programmes
and in achieving overall wildlife conservation and management objectives
effectively. This is especially true for remote regions of the world that are difficult
to access where few habitat studies have been undertaken to date but whose study
is urgently needed.
1. Introduction
The conservation of biodiversity of the 21st century faces two main challenges:
habitat loss and species extinction (see, for example, IUCN (2000) and BirdLife
International (2000)). It is widely accepted that the provisioning of habitat, and
managing it for fauna and flora, is crucial for the conservation and protection of
biodiversity. Current data on the distribution, status and ecology of wildlife species
are generally not available for many countries and for many habitats of the world.
*Corresponding author. Email: thomasgottschalk@surfeu.de

International Journal of Remote Sensing
ISSN 0143-1161 print/ISSN 1366-5901 online # 2005 Taylor & Francis Group Ltd
http://www.tandf.co.uk/journals
DOI: 10.1080/01431160512331338041
2632 T. Gottschalk et al.
This is especially true for the tropics, which present a habitat that covers about 10%
of the world. However, other large knowledge gaps still exist for the worlds oceans
that cover even two-thirds of the Earths surface.
Given the vast number of ecosystems in the world still unknown to us,
conventional ground-based survey and mapping methods cannot always deliver the
necessary information in a timely and cost-effective fashion. Satellite images
generally present the sole source of habitat data for many regions of the world
important to wildlife. With satellite-based remote sensing technology, we may be
able to make real progress in understanding why more species occur in some places
than in others and in identifying the most critical places (Bibby et al. 1992), allowing
for effective habitat conservation and management.
While habitat conservation focuses on general biodiversity and wildlife, birds are
traditionally among the species that have received much of the attention and
investigations. Reasons for this bias are simple: birds are relatively easy to identify,
their taxonomy is established, they are ubiquitous and they are sensitive or vulnerable
to environmental changes (Steele et al. 1984, Morrison 1986, Baillie 1991, Furness et al.
1993). Birds are prominent representatives of wildlife and have been used effectively as
bio-indicators of the state of complex ecosystems; often, their research presents role-
models for general wildlife research. Several examples of effects on birds related to
human-induced change to their habitat, such as land management practises,
introduction of exotic plants and animals, and hunting were given in Morrison
(1986) and Furness and Greenwood (1993). Further, Brooks et al. (2001) presented an
example of how birds were used in a land conservation planning programme to
represent the majority of other terrestrial vertebrate diversity.
By reviewing 109 representative studies published in 122 publications of the last
30 years, we attempt to give a state-of-art overview for a wide range of approaches
to remote sensing techniques that involve birdhabitat linkages. We further suggest
directions for applications, studies and improvements.
2. Methods
To identify papers that focus on bird and satellite remote sensing we used the terms
satellite, Landsat, SPOT, AVHRR, IRS, Quickbird, Ikonos or Orbview in
combination with the words avian or bird in the ISI Web of Science, which searches
for these words in the abstract, title and keywords of papers published mainly during the
last 10 years. However, most papers were found according to the snowball system:
papers focusing on birds and satellite remote sensing were searched for additional
references of the same topics. Our search was limited to studies published since the 1970s,
the beginning of the public use of satellite images, up to August 2004. Although this
method might have missed publications we assume that we have covered a representa-
tive part of the spectrum on birds in conjunction with satellite remote sensing topics.
3. Literature review
3.1 On the general use of satellite images and birds
Since 1972, when satellite images became widely available for public use, extensive
research has been conducted in which satellite images and animals have been
investigated, increasingly supported by Geographic Information Systems (GIS) (see,
for example Martinko 1981, Saxon 1983, Ormsby and Lunetta 1987, Pearce 1991,
Hansen et al. 2001, Kerr et al. 2001).
Avian habitat relationships 2633
Table 1. Classification scheme of avian habitat studies.
Analysing possibilities Possible output Map type

1. Identifying Maps of distribution Binary scaled
2. Characterizing Maps of habitat use Ordinal scaled
3. Monitoring Maps of change Binary/ordinal scaled
4. Predictive modelling Maps of potential distribution Binary/ordinal scaled
The objective of most of these studies was to find and to describe relationships
between habitat categorization, based on spectral reflectance pattern and geo-
referenced bird species records. Herr and Queen (1993) distinguish three different
ways of applying GIS in order to assess wildlife habitat. This classification is used in
table 1 and a fourth category of predictive modelling was added. The table was
supplemented by possible map outputs. It is worthwhile emphasizing that many of
the mentioned analysing possibilities are interrelated and often can be combined.
For instance, in order to model bird distributions the main characteristics of its
habitat need to be identified and mapped first, at least on a minimum level. In order
to monitor habitats effectively they have to be identified.
According to the classification scheme of table 1, habitats of the species were
identified in 50 out of 109 analysed studies. Seventy-seven studies dealt with
characterizing and classifying habitats, whereas modelling approaches to predict
species distribution or abundance were found in 48 studies. Monitoring of bird
species habitats was the objective in only 10 studies.
We found that most of the 109 analysed studies were performed in North America
(58 in USA, eight in Canada), three were conducted in Central America (two in
Costa Rica, one northern Central America), two in South America (Argentina,
Ecuador), 22 were carried out in Europe (nine in Great Britain, five in Spain, two in
Norway and Sweden and one in Austria, Belgium, France and Italy), eight in Africa
(Botswana, Senegal, Uganda, Tanzania, two in East Africa and two Crozet
Archipelago) and just four in Asia (China, India, Russia, Turkey), four in Australia
and two in Antarctica (table 2). Only 10 published studies that used satellite remote
sensing techniques have been conducted in tropical regions. In the reviewed studies
there is a huge variation in habitats and sizes of the study areas with the smallest
covering 13 km2 in Kansas, USA (Palmeirim 1988) and the largest with
5 393 343 km2 in Australia (Roshier et al. 2001) or the whole United States with
about 8 000 000 km2 (OConnor et al. 1996). Other large-scale studies comprise three
African countries (Wallin et al. 1992, Johnson et al. 1998) or the north-west Atlantic
(Huettmann and Diamond 2001). The median of the size of the study areas was
1580 km2 (n575). As computer technology and worldwide data availability has
developed considerably there is a steady increase in the size of the study areas during
the 30 year period publication were analysed for that review.
Except for nine studies (Priede 1983, Haney and McGillivary 1985a, Briggs et al.
1987, Haney 1989b, Jouventin et al. 1994, Guinet et al. 1995, Rodhouse et al. 1996,
Huettmann and Diamond 2001, Yen et al. 2004), the use of remote sensing imagery
for bird habitat investigations deals with terrestrial habitats. Most studies do not
provide an exact percentage of the different types of habitat that are investigated.
Therefore, we can report here only our own estimations. While remote sensing
applications of open habitats such as agricultural land, heath, pasture, grass-, wet-
and moorland were predominant, forests as the main habitats were listed in more
than a third of the studies.
Table 2. Location and area of the studies reviewed.
ID No. Reference Study area (study size, location, country)

1 Katibah and Graves (1978) 422.4 km2, Mendocino County, north coast of
California, USA
2 Cannon et al. (1982) 41 km2, western Oklahoma, USA
3 Lyon (1983) 109 km2, Sauvie Island, Oregon, USA
4 Cary et al. (1983) 6340 km2, north-west Missouri, USA
5 Priede (1983) California, USA
Briggs et al. (1984)
6 Palmeirim (1985) 52 km2, north-east Kansas, USA
7 Haney and McGillivary (1985a) Southern South Atlantic, USA
Haney and McGillivary (1985b)
Haney (1985)
Haney (1986)
Haney (1989a)
8 Schwaller et al. (1984) Ross Island, Antarctic
Schwaller et al. (1986)
Schwaller et al. (1989)
9 Haney (1989b) Gulf of Alaska and South Atlantic Bight, USA
10 Young et al. (1987) Washington, USA
11 Briggs et al. (1987) Entire Californian coast, USA
12 Hodgson et al. (1987) 1519 km2, Birdsville, south-east Georgia, USA
13 Shaw and Atkinson (1988) 54 695 km2, central Texas, USA
14 Hodgson et al. (1988) 1519 km2, Birdsville, south-east Georgia, USA
15 Palmeirim (1988) 13 km2, Kansas, USA
16 Miller and Conroy (1990) 25 km2, Eleuthera Island, Bahamas, USA
17 Sader et al. (1991) 570 km2, Sarapique region, north-east Costa Rica
18 Lauga and Joachim (1992) 2327 km2, north-west of Toulouse, south-west
France
19 Stoms et al. (1992) 25 000 km2, southern California, USA
20 Wallin et al. (1992) Somalia, Kenya, Tanzania
21 Aspinall and Veitch (1993) 1500 km2, north-east Scotland
22 Herr and Queen (1993) North-west Minnesota, USA
23 Homer et al. (1993) 2548 km2, Utah, USA
24 Griffiths et al. (1993) 28861 km2 squares in southern Upland/northern
Pennines, UK
25 Griffiths et al. (1993) 15261 km2, Grampian region, UK
26 Green and Griffiths (1994) 600 km2, Breckland, UK
27 Gustafson et al. (1994) 3620 100 ha, west-central and north-central
Indiana, USA
28 Andries et al. (1994) 288 km2, Hageland, Belgium
29 Jouventin et al. (1994) Possesion Island, Crozet Archipelago, south-west
Indian Ocean, France
30 Roseberry et al. (1994) 1127 km2, Hamilton County, Illinois, USA
31 Guinet et al. (1995) Ile aux Cochons, south-west Indian Ocean,
France
32 Knick and Rotenberry (1995) 2000 km2, Prey National Conservation Area,
south-western Idaho, USA
33 Warkentin et al. (1995) 732 km2, Selva Lacandona, Mexico
34 Avery and Haines-Young (1990) 79 sites, 1850 km2, Flow Country, northern
Scotland, UK
Lavers and Haines-Young
(1996a)
Lavers et al. (1996)
Lavers and Haines-Young (1996b)
Lavers and Haines-Young (1997)
35 Yates et al. (1996) Wash and Essex coast, east UK
Table 2. (Continued).

36 OConnor et al. (1996) 8000 000 km2, entire USA
37 Rodhouse et al. (1996) South Georgia, UK
38 Neave et al. (1996) 7581 km2, 165 field sites, south-east Australia
39 Austin et al. (1996) 2000 km2, Argyll, Scotland, UK
40 Gratto-Trevor (1996) 7650 km2, Mackenzie Delta, Northwest
Territories, Canada
41 Jrgensen and Nhr (1996) 35 000 km2, Ferlo region, northern Senegal
Nhr and Jrgensen (1997)
42 Donovan et al. (1997) Illinois, Indiana, Missouri, USA
43 White et al. (1997) 1580 km2, Monroe County, Pennsylvania, USA
44 Morrison (1997) 9 948 km2, Prince Charles Island, Foxe Basin,
Northwest Territories, Canada
45 Mack et al. (1997) 151 sample woodlands (200 m2 to 0.3 km2), area
of 75 km645 km, eastern UK
46 Tucker et al. (1997) 2903 km2, Catchment of River Tyne, northern
UK
47 Verlinden and Masogo (1997) Southern Kalahari, Botswana
48 Hepinstall and Sader (1997) 85 000 km2, Maine, USA
49 Thibault et al. (1998) Northern Quebec, Lacs des Loups Marins area,
Canada
50 Johnson et al. (1998) East Africa
51 Berry et al. (1998) Boulder, Colorado, USA
52 Roseberry and Sudkamp (1998) .145 900 km2, Illinois, USA
53 Fuller et al. (1998) 2497 km2, six study sites, Sango Bay, Uganda
54 Dettmers and Bart (1999) 560.9 km2 and 3260.3 km2, Wayne National
Forest, south-eastern Ohio, USA
55 Debinski et al. (1999) 324 km2, Greater Yellowstone Ecosystem, USA
56 Debinski et al. (2000) Gallatin National Forest, USA
57 Burke and Nol (2000) Ontario, Canada
58 Curnutt et al. (2000) 10 000 km2, Florida, USA
59 Miller and Cale (2000) 1680 km2, Kellerberrin district, Western Australia
60 Vander Haegen et al. (2000) 78 study plots, Columbia Basin, eastern
Washington, USA
61 Grillmayer (2000) Five study plots in Burgenland, Lower Austria
62 Howell et al. (2000) 10 study sites (3.4301.25 km2) in Missouri, USA
63 Haire et al. (2000) Boulder, Colorado, USA
64 Fauth et al. (2000) Northern Indiana, USA
65 Mortberg and Wallentinus (2000) 2500 km2, Sweden
66 Pino et al. (2000) 480 km2, Catalonia, Spain
67 Cully and Winter (2000) 14.2 km2 Saline County, Kansas, USA
68 Jones et al. (2000) Mid-Atlantic region, USA
69 Jones et al. (2001) Mid-Atlantic region, USA
70 Klaus et al. (2001) 1200 km2, province Gansu, China
71 Bajema and Lima (2001) 249.8 km2, 19 study sites, south-western Indiana,
USA
72 Pearce et al. (2001) North-east New South Wales, Australia
73 Daily et al. (2001) 21 study sites (ca 1.1 km2) within a 707 km2 circle,
San Vito, southern Costa Rica
74 Saveraid et al. (2001) 5060.01 km2, Greater Yellowstone Ecosystem,
USA
75 Osborne et al. (2001) ca 1266132 km, Madrid province, Spain
76 Roshier et al. (2001) 5393 343 km2, inland Australia
77 Huettmann and Diamond (2001) North Atlantic, Canada
78 Groom and Grubb (2002) 48 km river in Franklin, Madison and Union
Counties, Ohio, USA
Table 2. (Continued).

79 Shriner et al. (2002) Great Smoky Mountains National Park, USA
80 Shapiro et al. (2002) 861.6 km2, Fort Hood, Texas, USA
81 Pearson and Simons (2002) 3125 km2, Gulf coast of Texas, Louisana and
Mississippi, USA
82 Hepinstall et al. (2002) Maine, USA
83 Hunsaker et al. (2002) 606 km2, southern Sierra Nevada, California,
USA
84 Bani et al. (2002) 2500 km2, Lombardy, Italy
85 Pearman (2002) 2360.5 ha plots within 10 000 km2, Jatun Sacha
Preserve, Upper Amazon Basin, Ecuador
86 Reese et al. (2002) Dalarna county, Sweden
87 Suarez-Seone et al. (2002) 493 486 km2, peninsular Spain
88 Pearlstine et al. (2002) 176 187.56 km2, Florida, USA
89 Jepsen et al. (2002) 62 700 km2, Svalbard archipelago, Spitsbergen,
Norway
90 Debinski et al. (2002) Grand Teton National Park, Bridger-Teton
National Forest in Wyoming and Gallatin
National Forest in Montana, USA
91 Borboroglu et al. (2002) 340 km coastline and 45 islands in northern San
Jorge Gulf, Argentina
92 Meyer et al. (2002) Southern Oregon and northern California, USA
93 May et al. (2002) 2538 km2, 98 study plots, western South Dakota,
USA
94 Gottschalk (2003) 1100 km2, Serengeti Plains, Tanzania
95 Kastdalen et al. (2003) 2000 km2, Forollhogna National Park, Norway
96 Jenkins et al. (2003a) 500 km2, Everglades of South Florida, USA
Jenkins et al. (2003b)
97 Pidgeon et al. (2003) 661.08 km2 plots within 2406.76 km2, Fort Bliss
Military Reserve, northern Chihuahuan Desert
in south-central New Mexico, USA
98 Fearer and Stauffer (2003) 103.4 km2, Clinch Mountain Wildlife Management
Area, south-western Virginia, USA
99 Hennings and Edge (2003) 956.48 km2, 54 riparian sites, north-west Oregon,
USA
100 H-Acevedo and Currie (2003) 37 000 km2, North and Central America
101 Mayer and Cameron (2003) 644.53 km2, Ohio, USA
102 Jeganathan et al. (2004) Caddapah district, Andhra Pradesh, India
103 Bustamante and Seoane (2004) 37 700 km2, Andalusia, southern Spain
104 Yen et al. (2004). British Columbia, Canada
105 Venier et al. (2004) 800 000 km2, Great Lake Basin, Canada and USA
106 Hawkins (2004) 48 400 km2, Eastern North America
107 Berberoglu et al. (2004) Cukurova delta, Turkey
108 Seoane et al. (2004) 9 800 km2, western Andalusia, southern Spain
109 Fujita et al. (2004) 20 km bands along 21882679 km long migration
routes, Russia, China
3.2 Methodologies used in reviewed studies

The detailed methodologies of the studies depended mostly on bird census
techniques, remote sensing data, the use of additional environmental data and
statistical analyses. However, the typical studies using satellite-based remote sensing
and bird data follow a common pattern (see figure 1).
1. Most of the studies involve satellite image classification using pre-defined
ecological classes believed to be important. The importance of this
classification scheme was either derived from expert knowledge, from the
literature, or habitat class separability according to its specific reflectance and
the characteristics of the remote sensing system. The inherent assumption is
that the produced map has ecological relevance, and predictive power to the
wildlife species of interest.
2. Bird data were collected, roughly for the same time period when the imagery
was taken.
3. Additional data were used, such as terrestrially mapped vegetation structures,
soil maps or aerial photographs.
4. The GIS analyses involved a logical or arithmetic map overlay operation of
the satellite imagery with the bird data and other environmental data, and
were followed by statistical analyses of relationships between the bird data
and occurring classes in the satellite imagery and the other environmental
data, respectively.
5. In some studies, these relationships were then modelled by predicting the
likelihood of occurrence in an area with unknown bird data but known
habitat data derived from a satellite.
3.2.1 Satellite imagery. The most frequently used satellite sensor was Landsat (see
table 3). While more than half of the studies were based on Landsat Thematic
Mapper (TM) images, Landsat Multi-Spectral Scanner (MSS) scenes were used in
11 studies, sometimes along with Landsat TM scenes. Analyses of the higher spatial
Figure 1. Flow chart of the methodology typically utilized in studies using satellite-based
remote sensing and bird data.
resolution French SPOT satellite were performed nine times (Miller and Conroy
1990, Andries et al. 1994, Green and Griffiths 1994, Guinet et al. 1995, Thibault et al.
1998, Debinski et al. 2000, 2002, Klaus et al. 2001, Saveraid et al. 2001) and the
newer Indian Remote Sensing (IRS) two times (Debinski et al. 2002, Kastdalen et al.
2003). The Advanced Very High Resolution Radiometer (AVHRR) from the
National Oceanic and Atmospheric Administration (NOAA) was applied in 22
studies; six times it was used together with other satellite sensors. These coarse-
resolution imageries were often used in large-scale studies covering entire or large
parts of countries (Wallin et al. 1992, Suarez-Seoane et al. 2002, Venier et al. 2004),
continents (OConnor et al. 1996, Johnson et al. 1998, Roshier et al. 2001, H-
Acevedo and Currie 2003) or the ocean (Haney and McGillivary 1985a, Jouventin
et al. 1994, Huettmann and Diamond 2001). With the help of NOAA AVHRR
imagery often the actual or integrated normalized difference vegetation index
(NDVI) was calculated to produce maps of primary production or to detect green
biomass (Wallin et al. 1992, Andries et al. 1994, Hepinstall and Sader 1997, Verlinden
and Masogo 1997, Osborne et al. 2001, Saveraid et al. 2001), in order to detect habitat
change (Sader et al. 1991, Debinski et al. 2000), to map wetland distribution (Roshier
et al. 2001) or to explain the distribution of centres of bird endemism (Johnson et al.
1998). The NOAA AVHRR Climate Data Set incorporating values of atmospheric
temperatures and sea surface temperatures in the Atlantic and the Pacific, were used
for instance by Haney (1989b), Jouventin et al. (1994), Rodhouse et al. (1996),
Huettmann and Diamond (2001), Meyer et al. (2002) and Yen et al. (2004) to
analyse seabird habitats or to model their distribution.
A radar ERS image was applied by Thibault et al. (1998) to investigate ice-cover
on a river. The ice-cover was used to determine open water areas, which are the
habitat of the endangered harlequin duck Histrionicus histrionicus.
Detailed information on the processing infrastructure such as operating system,
software and hardware as well as the spectral bands used for image classification
were often lacking. Forty-nine out of the 109 reviewed studies did not provide
information about geo-coding of the remote sensing imagery. However, in 34 studies
images were geometrically corrected.
In terms of thematic resolution, the number of habitat classes derived from
remote sensing analyses ranged from 2 to 71 with a mean number of approximately
9.3 habitat (or vegetation/land use) classes (n568).
Only 24 of the reviewed studies provided details on the overall accuracy of the
classification process, which varied between 60% and 99% (mean value of about
85%). In 59 studies accuracy assessments were not performed or at least not
mentioned. Some studies utilized an existing land cover classification scheme, such
as, for example, the Centre for Ecology and Hydrology Landsat cover map of Great
Britain, which was used by Mack et al. (1997) and Dettmers and Bart (1999) or the
CORINE land-cover map of Europe, used by Seoane et al. (2004). When existing
land-use maps were used usually no details of the classification process or the
accuracy of the used map were mentioned. Further, 14 studies used raw data,
without classifying the image or calculating, for instance, the NDVI.
In terms of temporal scale, for many of the investigated studies the bird census
period does not fit with the exact time when the satellite image was taken. The
difference between time of satellite image acquisition and bird census period spans
up to 78 years (mean difference of 3.5 years). In only 42 out of 81 studies which
provided details this difference was less than 1 year.
Most analyses were not solely based on satellite imagery but used also additional
data and covariates. In 30% of the studies aerial photographsmainly colour infrared
pictureswere included. In the majority of these studies they were used to verify image
classification. Only a few studies used actual terrestrial estimations of per cent cover as
additional vegetation type information. In 20 of the 109 analysed studies, a digital
elevation model (DEM) was used. With the help of DEM data, slope, aspect or land
ruggedness could be calculated and then used for further birdhabitat investigations.
Other additional data sources were information on soils, nitrogen deposition, water
depths, climate and different vegetation measurements such as vegetation height or
cover value of a specific vegetation type, and man-made features such as roads and
buildings. Those terrestrial sampled data were used in 36% of all studies.
In several instances spatial texture measures, i.e. patchiness of a specific habitat,
spatial configuration, fragmentation, area and boundary length of landscape
elements and vegetation types were calculated. For example, Fauth et al. (2000),
Haire et al. (2000), Saveraid et al. (2001) and Meyer et al. (2002) used
FRAGSTATS, a spatial pattern analysis program for quantifying landscape
structure (McGarigal and Marks 1994). Hepinstall and Sader (1997) successfully
applied spatial texture measures to determine species preferences. Also, distances
between bird records and, for example, agricultural land, coast, shelf edge, roads or
buildings were calculated. These additional data were used for further refinements of
the birds habitat analysis or its model.
3.2.2 Bird data. In order to link the habitat information with birds, and depending
on the habitat or species being targeted, three main bird census techniques were used
(see table 4). Mostly, simple counts of sightings of bird species or their nests were
conducted. This was carried out in systematic and non-systematic ways, and
sometimes with the help of tools such as aircraft (Briggs et al. 1984, Hodgson et al.
1987, 1988, Haney 1989b), helicopter (Wallin et al. 1992, Herr and Queen 1993,
Morrison 1997, Jenkins et al. 2003a), boat (Briggs et al. 1984, Haney and
McGillivary 1985a, Briggs et al. 1987, Haney 1989b, Huettmann and Diamond
2001, Yen et al. 2004), canoe (Groom and Grubb 2002), radio telemetry (Young
et al. 1987, Hunsaker et al. 2002, Fearer and Stauffer 2003) or satellite telemetry
(Guinet et al. 1995, Jouventin et al. 1994, Rodhouse et al. 1996). A direct link
between bird breeding sites and satellite imagery was applied by Guinet et al. (1995)
and Schwaller et al. (1984, 1986, 1989). They identified nesting rookeries of penguins
on the ground by their spectral reflectance using Landsat TM and SPOT imagery.
Transect counts and Point-Stop Counts (PSC) were performed in 21 and 29
studies, respectively. Furthermore, bird data derived from the literature was used
in 32 studies. Twenty-two studies used presence/absence data and 16 of the
Table 3. Satellite images in 109 analysed studies. In some studies more than one satellite type
was used. In six studies the source of satellite sensor data was not given.
Satellite type No. of studies

NOAA Advanced Very High Resolution Radiometer (AVHRR) 22
European Remote Sensing Satellite (ERS) radar image 1
Indian Remote Sensing (IRS) 2
Landsat Multi-Spectral Scanner (MSS) 11
Landsat Thematic Mapper (TM) 66
Meteosat High Resolution Radiometer (HRR) 1
Syste`me Probatoire pour lObservation de la Terre (SPOT) 9
Table 4. Census techniques and bird data structure in 109 analysed studies. In some studies
more than one data source was used. In 35 studies detailed information on bird census data
was not given.
Number of studies
Presence/ Relative
Avian census technique/ Presence absence abundance True abundance
Source of bird data data only data (index counts) (densities)
Counts of individuals/ 14 10 4 1
nests/colonies (systematic
and random)
PSC 1 8 20
Transect counts 1 4 15 1
investigated studies were based on presence only data. Census techniques that use
counts of bird detection as an index of relative abundance (Rosenstock et al. 2002)
were used 39 times. Distance Sampling (Buckland et al. 1993), an empirical
modelling technique for survey data that directly estimates bird densities (absolute
abundance) by taking the variation in species detectability into consideration, were
only used in the study by Kastdalen et al. (2003). In eight studies the source and
census methodology for the bird data was not given.
Little research has been carried out on multi-species and avian community
analyses. Most of the research using remote sensing was carried out to find habitat
use, preferences or to predict species distribution either for a single species (41%) or
for two to five bird species (11%). More than 20 bird species were used in 23% of all
studies investigated.
3.2.3 Statistical analyses of wildlifeenvironment relationships. For the study of
relationships between bird species distribution patterns and environmental variables
the 109 studies used a wide array of statistical methods. Some of them have been
specifically designed for these studies, for example Aspinall and Veitch (1993) and
Tucker et al. (1997) who used specific Bayesian approaches. Other studies applied
common statistical techniques. Many papers did mention statistical tests, but were
not really specific about it. Mostly univariate and bivariate tests as well as
multivariate statistics were carried out. Bayesian models were used in Aspinall and
Veitch (1993), Griffiths et al. (1993), Tucker et al. (1997), Hepinstall and Sader
(1997) and expert opinion techniques were applied by Pearce et al. (2001). Chi-
square test, T-test and MannWhitney U-test were commonly carried out to
compare single habitat variables between study plots with and without the species,
and between two study areas containing species. The use of logistic and multiple
regressions to predict species distribution has strongly increased since the 1990s (e.g.
Avery and Haines-Young 1990, Lauga and Joachim 1992, Griffiths et al. 1993,
Austin et al. 1996, Nhr and Jrgensen 1997, Saveraid et al. 2001, Huettmann and
Diamond 2001, Osborne et al. 2001, Meyer et al. 2002, Jepsen et al. 2002, Suarez-
Seoane et al. 2002, Jeganathan et al. 2004, Venier et al. 2004, Yen et al. 2004).
4. Discussion
4.1 Topics related to the remote sensing imagery
The selection of the right satellite sensor is a crucial decision which affects the
outcome of the habitat description and the wildlifehabitat relationship. So far, no
methods are known to us that provide guidance for such decisions in detail. The
reviewed studies did not show many considerations in regards to spatial scale and
choice of appropriate satellite sensor. Topics like mismatch of scales between remote
sensing data and avian data were discussed in only few studies. Thibault et al. (1998)
and Venier et al. (2004) compared different satellite sensors and showed how the
choice of satellite sensor may affect results in a bird specieshabitat study. However,
the preferred usage of Landsat in almost 80% of all analysed studies is probably due
to the fact that the Landsat programme was the first major satellite programme for
public use. In addition, the high spectral resolution of this sensor, its robust data
structure, and a good priceperformance ratio contributed to its success.
ONeill et al. (1996) recommend that the spatial resolution should be two to five
times smaller than the spatial features of interest. That involves knowing the
landscape features relevant to the species of interest beforehand, which is often
impossible. Additionally, studies containing more fine-grained habitat information
need adequate species knowledge or are fraught with many assumptions about the
species ecology than on studies that have fewer details and are more general in
design (Pearson and Simons 2002). Morrison et al. (1992) suggest a hierarchical
approach viewing habitat analysis first from the broadest scale, and then working
down to the finest level of scale necessary to answer the question of interest.
For some bird species the habitat classification based on satellite images can be of
insufficient resolution (see examples in table 5). Depending on the spatial resolution
of the sensor, specific hard edge features such as narrow linear grooves, hedgerows,
fences and small patches of land or single trees cannot be (precisely) identified in
many satellite images, although, they may be of major importance to specific wildlife
species (see Mack et al. 1997). For these species the features are suggested to be
identified and be mapped by ground surveys or by specific image analysis techniques
such as contextual analysis or edge detection (Andries et al. 1994). Additionally,
satellite imagery may not capture important events affecting the habitat (Jenkins
et al. 2003b). Several non-habitat factors, like historical reason, intraspecific and
interspecific interaction, microhabitat characteristics or the three-dimensional archi-
tecture of vegetation are beyond the scope of satellite remote sensing technology.
Table 5. Examples of limitations of bird specieshabitat relationship studies based on satellite

images.
Reference Bird species Satellite sensor Main limitations mentioned

Palmeirim Ruffed grouse Landsat TM Certain pertinent landscape
(1985) (Bonasa umbellus) features and local variables
could not be detected
Herr and Greater sandhill crane Landsat TM Certain pertinent landscape
Queen (1993) (Grus canadensis tabida) features and local variables
could not be detected
Gratto-Trevor 10 shorebird species Landsat TM Edge habitats could not be
(1996) identified
Tucker et al. Coal tit (Parus major), Landsat TM Satellite-based classes extended
(1997) golden plover (Pluvialis over a range of habitats
apricaria), snipe
(Gallinago gallinago)
Saveraid et al. 11 passerine bird species Landsat TM Spatial resolution was too
(2001) and SPOT coarse and did not contain
enough information
Generally, the finer a satellite-derived habitat map is classified, the more bird
specieshabitat relationships can potentially be revealed. The new generation of high
resolution satellite systems such as Ikonos, Quickbird and Orbview may advance
possibilities to achieve a detailed habitat map, as they provide multi-spectral
imagery of 2.84 m spatial resolution with an effective revisit capability of just a few
days. Their characteristics offer the possibility of doing research that was previously
only possible by ground-based studies or using airborne sensors (Clark et al. 2004,
Ehlers 2004). Although their low spectral resolution (lacking of short-wave infrared)
was identified by Mehner et al. (2004) as the main weakness of the data quality, they
found that the high resolution satellite systems were capable of higher accuracy
than achieved with previous approaches. However, usually the accuracy of the
classification increases with a decrease in the number of classes used (Rees 1990). A
higher-resolution satellite image could improve habitat separability as they
introduce a substantial amount of extra detail and spatial variation into the
classification process, and thus provide a higher number of habitats on a finer scale.
Working at a too detailed level of scale could mean that the number of points per
habitat type with bird records would significantly decrease thereby diminishing the
power of the statistical tests to distinguish differences between habitat use and
availability (Garshelis 2000). Stoms et al. (1992) tested the effects of coarser spatial
resolution of a satellite-based habitat map. They eliminated polygons of a habitat
map that are less than 20 ha and observed no significant effects on a habitat
suitability model for the California condor Gymnogyps californianus. Venier et al.
(2004) demonstrated that the finer resolution Landsat MSS land cover were not
more useful than the coarser AVHRR land cover at identifying local habitat
associations of 10 forest songbirds. Seoane et al. (2004) compared the predictive
ability of bird distribution models derived from 30 m and 250 m resolution satellite-
based maps. As well, they found no significant differences and suggested that
improving vegetation maps above a certain limit has no effect in their power to
predict bird distribution.
Many sources of bias stem from the image processing algorithms employed, such
as atmospheric correction, supervised or unsupervised classification techniques, and
image transformations. These choices can be driven by the processing software
available to the user. Additionally high cloud cover and shadow can restrict the
choice of imagery available to the research project. As a consequence, this restriction
can produce a significant temporal difference between bird and satellite sensor data.
For example, Justice and Hiernaux (1986) stated that the 18 day orbital period of
the Landsat platform is insufficient to provide regular cloud-free coverage during
the 2 month growing and breeding season of the Sahelian Zone.
Assigning habitat types to pixels can cause errors (White et al. 1997) as habitat
classes determined from spectral signatures of the image may not directly
correspond to specific and relevant avian units or habitats for birds. Certain bird
species are likely to be better represented than others by the habitat classes. A
difference between specialist and generalists species was found by Jones et al. (2000),
as landscape characteristics derived from Landsat TM explained a greater
proportion of the specialist species richness than the generalist species richness.
Generally, it is recommended to partition the habitat types in terms of relevant
features according to the habitat preferences (Garshelis 2000). In order to map
habitats under species specific criteria a post-processing of the classified image is
sometimes necessary. Unfortunately, this post-processing is highly dependent on the
prior knowledge of the interpreter and can lead to subjective biases. Thus, extracting
land cover information from remote sensing images through a classifying process
involves much human intervention and judgement, rather than being driven by
objective biological requirements of the species. Drawing boundaries in natural
vegetation areas can be highly problematic (see, for example, Fortin et al. 2000,
Janssen 2000). Habitat types can display gradual change in community type. This is
characterized in the satellite imagery through high frequency variation in spectral
values and in a high proportion of mixed pixels (see Aspinall and Veitch 1993).
Obviously, the level of variation has a limiting effect on the correlation between
habitat and species (Nagendra 2001). Although sometimes a separation of transi-
tional stages might be necessary, this separation can result in too many habitat
classes. That again would increase bird census efforts, as a higher density of bird
records would be necessary to achieve significant speciesenvironment relationships.
If only few habitat classes are defined, each is likely to have considerable variation in
plant species composition and other attributes (Cully and Winter 2000) and thus
reduce the possibility to detect significant bird specieshabitat relationships.
In a specieshabitat study, map accuracy can usually be confirmed efficiently by
using the ground data from the bird sample plots (Berry et al. 1998, Seoane et al.
2004). The importance of a high accuracy of image classification is illustrated by
Lyon et al. (1987). The authors showed that even a 5% change in classification
accuracy of a land-cover map caused significant differences in levels of a habitat
quality index. However the study did not focus on general sources of noise. Hunsacker
et al. (2002) compared relationships between occurrence of California spotted owl
Strix occidentalis and proportion of canopy-cover derived from Landsat TM imagery
and derived from aerial photography. They found that differences in canopy-cover
composition of less than 10% can significantly affect occupancy by the owl.
In several studies computer-assisted image classification was not undertaken;
instead raw reflectance data were used, or images were hand-traced. The dis-
advantage of this approach might be that relationships between reflectance data and
species distribution can be identified but habitats are not described and stratified in
ecologically meaningful terms. In addition, speciesreflectance studies tend not to
provide reliable answers when applied to areas other than the ones in which they
were developed for because season, weather and soil condition may adversely affect
the reliability of extrapolation to another image (Nagendra 2001). Furthermore,
exact relationships between bird occurrence and satellite-derived land cover can
fluctuate. Avery and Haines-Young (1990) and Lavers et al. (1996) found
correlations between dunlin (Calidiris alpina) abundance and near-infrared
reflectance of the Landsat image in Caithness and Sutherland, Britain, but not for
the nearby Shetland Islands as on that island the easily detectable pools were
missing (Lavers and Haines-Young 1997). Further, Morrison (1997) concluded that
mapping wildlife habitats of the Arctic requires constant ground truthing and re-
calibration of habitat classes in different regions.
The importance of a large number of environmental variables increases with the
complexity of the animals and their habitats, and with the degree of accuracy of the
speciesenvironment relationships to be studied. Vertical habitat structure features
can severely affect wildlife distribution and abundance. Severe limitations exist in
woodlands and forests, as most satellites, except for microwave imagery and
Synthetic Aperture Radar (SAR), are unable to detect or characterize the structure
of lower layers, which are likely to be of importance for some species (see, for
instance, Palmeirim 1985). In some of the reviewed studies, the additional ground-
based data used along with satellite-based data were biased by the fact that the data
were chosen because they were available. However, to find explanatory and
meaningful variables determining wildlife occurrence is a difficult endeavour and
depends much on the biotope of interest. In order to measure for instance forest
habitat structures, Morrison et al. (1992) list 29 different variables, such as woody
foliage profile density or tree stump size. To analyse habitats or to build models of
species habitat relationships using remote sensing data and GIS methods, a species
must be either common enough and/or habitat-specific enough to exhibit a sig-
nificant relationship with one or more remotely sensed habitat types (Debinski et al.
1999). Significant speciesenvironment relationship depends on the bird species
itself, and whether it tends to be ubiquitous or rare and to the degree of habitat map
detail. The main habitat features, which are assumed to be closely related to the
objectives, should be mapped, as far as time and financial resources are available.
Such specific habitat features can usually indicate habitat preferences (e.g. Aebischer
and Roberston 1992, Jones 2001, Manly et al. 2002). Some of these habitat features
are helpful as reference data for digital image classification. Roseberry and
Sudkamp (1998) selected variables that were theoretically meaningful, predictive,
and consistent with the resolution and accuracy of the land-cover data. Pearlstine
et al. (2002) suggested that predicted species models only based on land cover classes
fail to incorporate many of the basic ecological characteristics of the species. By
applying ground-based habitat data, Mack et al. (1997) found that speciesarea
relationships described the bird species numbers better than those from remotely
sensed data. However, they also showed that remotely sensed data are of sufficient
spatial resolution for coarse scale estimates of speciesarea relationships. To assess
the relative contribution of ground-based data and satellite sensor data in explaining
differences in data of 62 bird species, Gottschalk (2003) used the coefficients of
determination (R2) to compare different regression models. The best fit of the
models was achieved by combining satellite- and ground-based variables (R250.26)
compared to the sole use of satellite sensor data (R250.18). Nagendra (2001)
pointed out that ancillary data may best to predict species distribution in natural
areas. Keeping in mind that for large scale studies exhaustive terrestrial data are
often not available, Venier et al. (2004) showed that significant improvements to
model bird species distribution can also be made by adding worldwide available
NOAA AVHRR climate data to land cover maps.
4.2 Temporal scale and seasonality

In many ecological studies a source of bias is presented by the relatively short census
period, e.g. within a season and rarely more than over a period of 3 years. In
addition, an analysis based on multi-temporal imagery has greater discriminatory
power than the analysis of a single-date image. Firstly, habitat separability and
identification can be improved by multi-date image based classifications. Secondly,
it is unlikely that short duration studies adequately reveal the variability inherent in
habitat use by animals. Temporal shifts of habitat use and habitat suitability can
occur within time of day, and within and between seasons and years. These shifts
can result from numerous factors such as weather, food abundance, and densities of
conspecifics, competitors and predators (Block and Brennan 1993). For example,
extreme seasonal weather events, such as El Nino, can force animals into habitats
not used during typical years (Homer et al. 1993) and thus bias wildlifehabitat
investigations. Therefore multiyear replication is necessary to accurately identify

most of the species present and the image should normally be closely matched in time
to the period(s) of the bird survey. By contrast, some ground cover characteristics may
change little from year to year (see, for example, Lavers and Haines-Young 1997).
4.3 Bird census

In many of the reviewed studies, non-systematic and non-standardized censuses and
surveys were used. These techniques, however, have the disadvantages that they are
not transparent and that results can be biased and based on incomplete and not
representative data. Of the standardized census techniques, geo-referenced PSC and
further line transect counts seem to be the most appropriate methods, particularly
for studies in remote or difficult to access areas. This is because they are less walking
intensive and thus less time-consuming and more effective. These census techniques
are useful for a broad spectrum of both, natural and non-natural habitats. Geo-
referenced PSC and transect counts can be spatially adjusted to the required scale of
the remote sensing imagery. If data on distributions and true abundances (densities)
are needed, especially for accurate modelling approaches, for many applications,
distance sampling (see Buckland et al. 1993, Bibby et al. 2000) is usually recom-
mended. Its use allows correction for the possibility of detecting birds as it takes
into account that some birds are detectable over much greater distances than others,
and that a species may be more easily spotted in one habitat than another.
General sources of errors caused by bird census techniques are discussed, for
example, in Ralph and Scott (1981), Bibby et al. (2000) and Farnsworth et al. (2002).
The number of survey points should be adjusted, as well as the number of required
repeats of surveys in order to obtain robust estimates. No study was found that
tested different efforts on bird census to show the effect on speciesenvironment
relationships. However, Bibby et al. (1998) recommended a minimum of about 50
points per habitat. For rare species, a higher bird census effort, e.g. density and
repeats of surveys, might be required in order to obtain robust estimates.
If a model is based on relative or absolute densities, as well as on presence
absence data, distortion of concern can be caused by autocorrelation (Schneider
1994). Double recordings of the same bird at two or more (neighbouring) points can
be possible for high, mobile vagrant or large birds visible from two or more points.
Although the number of papers that involve satellite tracking techniques have
increased (e.g. Ginati et al. 1995, Hatch et al. 2000, Berthold et al. 2001, Hashmi
et al. 2001, Watzke et al. 2001), only three of the papers reviewed by us referred to
these techniques (Jouventin et al. 1994, Guinet et al. 1995, Rodhouse et al. 1996).
This may be due to current inaccuracies in location determination, which actually
lies at best around 30150 m and would only allow broad-scale analyses. It is to be
expected, however, that future developments will improve satellite tracking techniques
and will allow a cheaper and more precise determination of animal locations. Linking
of precise bird locations derived by satellite tracking, with habitat maps derived by
space-borne sensors, presents a future challenge that needs to be investigated further.
4.4 Analysing approaches to investigate birdhabitat relationships and model

performance
Remote sensing imagery offers great potential to investigate the subject of habitat
availability which is crucial to address true preferences from habitat use, and for
valid model inferences (e.g. Aebischer and Robertson 1994, Manly et al. 2002). The
reviewed papers did not discuss these issues, and thus appeared to be biased towards
showing statistical relationships but not inferred biologically meaningful habitat
links or true habitat preferences.
An alternative can be the use of resource selection functions (RSF) (e.g. Aebischer
and Robertson 1992, Boyce and McDonald 1999, Manly et al. 2002, Kastdalen et al.
2003), which offer the opportunity that only a small but representative fraction of
the study area needs to be sampled and surveyed for the wildlife species. Once a
robust relationship has been established this relationship can be modelled and
extrapolated to the entire area that is recorded, classified and inventoried by remote
sensing. Additionally, software developments like Biomapper (for presence only
data) and GARP (Genetic Algorithm for Rule-set Prediction) have received
much attention in recent times for spatial analysis and predicting species occur-
rences (Hirzel et al. 2002, Stockwell and Peters 1999). Furthermore, programmes
such as FRAGSTATS or the Patchanalyst are well-used tools in remote sensing
applications for quantifying landscapes/habitats and studying wildlifehabitat
relationships.
One utility of models is to guide further efforts and studies (Garshelis 2000). The
methodology, however, needs to be adjusted by a refinement process that is based
on the specific abiotic and biotic circumstances. Model verification refers to a broad
spectrum of performance standards and criteria including model credibility and
acceptability, realism, generality, precision, breadth, and depth (Marcot et al. 1983).
Key concepts such as threshold-based and non-threshold-based accuracy checking
for models of species habitat or species distribution are given for instance by
Fielding and Bell (1997) and Pearce and Ferrier (2000). So far, however, they have
not been applied very often. The strength of the specieshabitat relationship is
fundamental in a model and also in a monitoring programme based on bio-
indicators. It should therefore be improved through rigorous testing. The value of
ground truthing the relationship between remotely sensed habitat and species on the
ground cannot be overestimated (Debinski and Humphrey 1997).
5. Conclusions and suggestions for future studies

Thirty years of using satellite-based remote sensing and bird data in a GIS
environment has greatly increased our ability to assess causal effects in species
environment relationships. Many of the findings depicted from more than 100 avian
studies using satellite remote sensing are relevant for general wildlife studies and can
therefore used as a template. These techniques allow the convenient presentation,
efficient prediction and monitoring of the distribution of species/communities and
the estimation of the bird population sizes for large and remote areas. Its highest
potential might exist in its application in ecosystems where access is limited and
coarse and quick but quantitative estimates with statistical confidence limits on
biodiversity are urgently needed. In terrestrial environments, combining satellite
sensor data with terrestrially derived measurements seems to be the most effective
way to determine speciesenvironment relationships. For most study sites, the only
alternative to remote sensing is ground data collection, which makes the cost of
covering large areas prohibitively high.
For issues that deal with best habitat classification, the optimum number of habitat
features and the most appropriate scales need to be addressed. This has to be specified
according to the ecology of the target species and the local landscape configuration. To
better understand and to improve the quality of the inference and for comparative
analyses, it is highly recommended that these studies are standardized. Therefore, bird
census techniques, terrestrial measurements, satellite image sources and classification,
accuracy assessment and statistical analyses should be well thought out and
documented in further investigations. This detailed documentation should be given
for data collected by the investigator as well as by others.
More multidisciplinary studies and approaches are needed in order to properly
advance the benefits that remote sensing methods can bring to the research and
conservation on wildlife and its habitats. To date, not all remote sensing
applications in ornithology are mature, and costbenefit analyses of remote sensing
applications for precise avianhabitat applications are somewhat lacking. Neverthe-
less, the method has been well established within the last 30 years and is likely to
play an increasing role in a huge range of applications, especially those determin-
ing the essential spatial elements of species habitats, modelling the distribution of
birds and habitats, and monitoring their changes at a global level.
Acknowledgements
We would like to thank D. Gann, P. Lurz, S. Franklin, M. Wulder, J. Linke, G.
McDermid, Lisa Strecker, and two anonymous reviewers for their input and
suggestions to improve earlier versions of the manuscript. F.H. was supported by
the University of AlaskaFairbanks, and with a Postdoctoral Killam Fund, held
with the Geography Department at the University of Calgary, Canada.
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International Journal of Remote Sensing

Vol. 26, No. 12, 20 June 2005, 26312656

Thirty years of analysing and modelling avian habitat relationships using

T. K. GOTTSCHALK*{, F. HUETTMANN{ and M. EHLERS

(Received 25 July 2003; in final form 13 October 2004 )

The application of remote sensing and Geographic Information Systems (GIS)

*Corresponding author. Email: thomasgottschalk@surfeu.de

Table 1. Classification scheme of avian habitat studies.

Analysing possibilities Possible output Map type

Table 2. Location and area of the studies reviewed.

ID No. Reference Study area (study size, location, country)

ID No. Reference Study area (study size, location, country)

ID No. Reference Study area (study size, location, country)

3.2 Methodologies used in reviewed studies

Satellite type No. of studies

Table 5. Examples of limitations of bird specieshabitat relationship studies based on satellite

Reference Bird species Satellite sensor Main limitations mentioned

4.2 Temporal scale and seasonality

investigations. Therefore multiyear replication is necessary to accurately identify

4.3 Bird census

4.4 Analysing approaches to investigate birdhabitat relationships and model

5. Conclusions and suggestions for future studies

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